Cost-effectiveness analysis of options within an Integrated Crop Management regime against great spruce bark beetle, Dendroctonus micans, Kug. (Coleoptera: Scolytidae)

1 Great spruce bark beetle Dendroctonus micans (Kug.) (Coleoptera: Scolytidae) has been subject to an Integrated Crop Management (ICM) regime in Great Britain since its first discovery in 1982. The elements of the ICM approach are sanitation felling of the initial infestations, restriction on movement of potentially infested conifers to prevent spread to uninfested parts of the country and biological control through rearing and release of the specific predatory beetle Rhizophagus grandis Gyll. (Coleoptera: Rhizophagidae). Such a programme carries costs that have to be evaluated against the benefits to the ICM strategy. 2 This paper presents a cost-effectiveness analysis of three options – (i) continue with the current policy of restriction on movement of infested timber and use of the imported predator R. grandis, (ii) do nothing (i.e. drop all attempts to reduce the spread of D. micans), or, (iii) introduce a new east–west Dendroctonus Micans Control Area (DMCA) to define the management area. Assumptions on rates of mortality and natural spread were based on research into D. micans in Britain’s spruce forests. 3 Appraisal indicated that the current policy was the most cost-effective, even when subject to sensitivity analysis to test the limits of the assumptions included in the models. It is concluded that the current policy should remain in force.     

Past attacks influence host selection by the solitary bark beetle Dendroctonus micans

1. A spatio‐temporal study of host selection and local spread of a solitary bark beetle attacking live spruce Dendroctonus micans (Kugelann) was carried out using a combination of standard statistical methods, geostatistical analyses, and modelling. The study was based on data from three plots (150–300 trees, 0.3–1 ha) from 1978 to 1993. All trees were mapped and successful and abortive bark‐beetle attacks on each tree were counted annually. Because the attacked trees usually survived, temporal attack patterns as well as spatial patterns could be analysed. 2. The distribution of successful insect attacks on the trees was slightly aggregative, indicating some degree of choice rather than totally random establishment. 3. The level of yearly individual attacks per tree was very stable, suggesting that D. micans usually leave the host in which they develop. 4. The attacked trees were distributed randomly in the plots; at the study's spatial scale, the insects dispersed freely throughout the plot (no spatial dependence). 5. On the other hand, time dependence was strong; some trees were attacked repeatedly while others were left untouched. 6. Among a choice of scenarios (random attack, fixed variability in individual host susceptibility, induced host susceptibility following random attack), the best fit was obtained with the model involving induced individual host susceptibility. This type of relation to the host tree contrasts strongly with patterns generally described in host–plant relationships (including gregarious, tree‐killing bark beetles), where local herbivore damage results in induced resistance. 7. These results suggest that the first attacks in a new stand are made at random, that all or most of the beetles emerging from a tree disperse and resample the stand, and that they settle preferentially on trees that were colonised successfully by previous generations.     

Isolation and virulence of entomopathogenic fungi against the great spruce bark beetle,Dendroctonus micans (Kugelann) (Coleoptera: Scolytidae)

Twelve fungal strains including Lecanicillium muscarium (Petch.) Zare and Gams, Isaria farinosa (Holmsk.) Fr., Fusarium sp., Beauveria bassiana Sensu Lato and Beauveria sp. were isolated from larvae and adults of D. micans. In addition, virulence of these isolates against this pest was determined. Conidia suspensions of 1×10⁶ conidia mL^–1 were applied to larvae and adults. The highest mortality and mycosis for larvae were obtained from isolate ARSEF 9271 (Beauveria bassiana) with 90% mortality and mycosis within 10 days. ARSEF 9271 also produced 93% mortality and mycosis in adults. On the other hand, the highest mortality and mycosis for adults were obtained with isolate ARSEF 9272 (Beauveria sp.), with 100% mortality and 80% mycosis within 10 days. These results indicate that isolates ARSEF 9271 and ARSEF 9272 seem to be the most promising potential fungal biocontrol agents against D. micans.     

Temperature‐dependent development of the great European spruce bark beetle Dendroctonus micans (Kug.) (Coleoptera: Curculionidae: Scolytinae) and its predator Rhizophagus grandis Gyll. (Coleoptera: Monotomidae: Rhizophaginae)

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Turbulence, trees and semiochemicals: wind-tunnel orientation of the predator, Rhizophagus grandis, to its barkbeetle prey, Dendroctonus micans

Abstract. Insects apparently responding to the visual stimulus offered by a tree or other object may also be responding to the variety of physical effects caused by the obstruction, including turbulence, a reduction in wind velocity and eddies. The relative importance of prey odour associated with the bark beetle Dendroctonus micans (Kugelann) (Scolytidae), prey-host-tree odour, silhouette, and physical barrier to the wind, in the orientation and landing behaviour of a predatory beetle, Rhizophagus grandis Gyllenhall (Rhizophagidae), were investigated. R. grandis responded very positively to the frass of its prey when presented on a 'tree' but not to the same quantity of frass presented alone. Frass on black plastic uPVC pipes was significantly more attractive than frass on real host tree logs, suggesting that host-tree volatiles do not enhance the attractiveness of prey frass.
The beetles responded to 'transparent' Mylar three-dimensional cylinders in the same way as they did to the black plastic pipes. Beetles were also tested with frass and two-dimensional stimuli offering a visual stimulus or physical barrier alone or in combination. Frass with a two-dimensional visual stimulus was no better than frass alone: few insects landed. Significantly more insects were attracted to the same two-dimensional visual stimulus with a hidden upwind barrier producing turbulence, but a similar number were attracted to the frass in front of a hidden barrier alone. As many plants are similar in size to the logs used in these experiments, comparable effects might influence the behaviour of other phytophagous insects, their predators and parasitoids.     

Differential impact of egg predation by Zetzellia mali (Acari: Stigmaeidae) on Metaseiulus occidentalis and Typhlodromus pyri (Acari: Phytoseiidae)

The differential impact of Zetzellia mali on the phytoseiids Metaseiulus occidentalis and Typhlodromus pyri was studied in the laboratory and by analysis of population from orchard plots that contained either phytoseiid, similar numbers of prey mites and high or low densities of Z. mali. Five hypotheses were evaluated to explain why Z. mali had more impact on M. occidentalis in the field than on T. pyri. Given equal opportunity, Z. mali adult females did not consume more M. occidentalis eggs than T. pyri eggs nor did adult females of either phytoseiid inflict greater mortality on Z. mali eggs or larvae through attack or consumption. There was no difference in the within-tree association of Z. mali adult females with eggs of either phytoseiid species nor were there differences in the way prey mites (all life stages) were spatially partitioned between adult female Z. mali as compared with adults and deutonymphs (combined) of either phytoseiid. The foraging area of adult female Z. mali and the oviposition locations of the two phytoseiids from both field and laboratory data were compared using spatial statistics. Metaseiulus occidentalis laid significantly more eggs in the primary foraging area of adult female Z. mali than T. pyri. This was the only factor identified which may explain the greater impact of Z. mali on M. occidentalis. The impact of these interspecific effects on the persistence of predatory mite guilds and biological control are discussed.     

The influence of aphid-produced honeydew on parasitoid fitness and nutritional state: A comparative study

Honeydew is a sugar-rich resource excreted by many hemipteran species and is a key food source for other insect species such as ants and parasitoid wasps. Here, we evaluated the nutritional value of 14 honeydews excreted by 13 aphid species for the generalist aphid parasitoid Lysiphlebus testaceipes to test a series of hypotheses concerning variation in the nutritional value of honeydew. There was a positive correlation between the body sugar content of honeydew-fed parasitoids and their longevity. This information is valuable for biological control researchers because it demonstrates that the nutritional state of honeydew-fed parasitoids in the wild can indicate their fitness, independently of the honeydew source they have fed on.Although the carbohydrate content and longevity of L. testaceipes differed greatly among the different honeydews, we did not find a significant effect of aphid or host plant phylogeny on these traits. This result suggests that honeydew is evolutionarily labile and may be particularly subject to ecological selection pressures. This becomes apparent when considering host aphid suitability: Schizaphis graminum, one of the most suitable and commonly used hosts of L. testaceipes, produced honeydew of the poorest quality for the parasitoid whereas Uroleucon sonchi, one of the few aphids tested that cannot be parasitized by L. testaceipes, excreted the honeydew with the highest nutritional value. These data are consistent with the hypothesis that hemipterans are subject to selection pressure to minimize honeydew quality for the parasitoids that attack them.     

Take-off capacity as a criterion for quality control in mass-produced predators,Rhizophagus grandis (Col.: Rhizophagidae) for the biocontrol of bark beetles,Dendroctonus micans (Col.: Scolytidae)

Time spent by adult beetles in cold storage at 3–7°C accounted for 81% of the loss of take-off capacity inRhizophagus grandis Gyllenhal in windtunnel experiments. At the age of three weeks, the insects were at their highest take-off capacity at about 80%. This was followed by a steady decrease, 7% of the insects failing to take-off each month. Changes in the fat reserves during cold storage could explain at least partly this reduction of flight capacity although there was no significant difference in fresh weight between insects that were able/unable to take-off. Sex had a significant influence on take-off rates, with an 8.7% higher take-off rate in females. Take-off capacity was further reduced when the insects were mass-produced in cultures using parent beetles submitted to a blend of synthetic oviposition stimulants instead of live prey larvae. Response to synthetic attractants by those insects which took-off, however, was not influenced by cold storage or by the use of synthetic stimulants in the culture medium.

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Résumé Le temps passé en conservation à basse température (3–7°C) depuis la métamorphose est intervenu pour 81% dans la perte de la capacité d'envol chez le coléoptère prédateurRhizophagus grandis Gyll. lors d'expériences en tunnel de vol. A l'age de trois semaines, les insectes sont à leur plus haut niveau d'envol (envol d'environ 80% des insectes). Par la suite, il y a un déclin constant de la capacité d'envol, à raison de 7% des insectes chaque mois. Des changements dans les réserves lipidiques peuvent partiellement expliquer cette réduction, bien qu'il n'y ait pas eu de différence entre le poids frais d'insectes capables de s'envoler et celui d'individus qui en étaient incapables. Le sexe a une influence sur le taux d'envol, avec un taux d'envol significativement plus élevé de 8.7% chez les femelles. La capacité d'envol est encore réduite chez des insectes qui ont été produits dans des élevages de masse où les parents étaient soumis à un mélange de stimuli de ponte de synthèse au lieu d'être mis en présence de larves deD. micans vivantes. Chez les insectes qui prennent leur vol, cependant, la réponse aux attractifs de synthèse est indépendante de l'age ainsi que des conditions d'élevage.

     

Nestbox orientation: a species-specific influence on occupation and breeding success in woodland passerines

Capsule Nestbox orientation has species-specific influences on nestbox occupation and breeding success for woodland passerines.

Aims To determine if nestbox orientation had an influence upon nestbox selection or breeding success for three co-occurring woodland passerines.

Methods We analysed 15 consecutive years of breeding data (1990–2004) from 295 nestboxes in the UK using circular statistical analyses to examine the influence of orientation upon nestbox occupation and breeding success for three species, Blue Tit Cyanistes caeruleus, Great Tit Parus major and Pied Flycatcher Ficedula hypoleuca.

Results The three species used nestboxes of all orientations during the 15-year period. The frequency of nestbox occupation by Great Tits correlated with orientation (the mean number of nests in boxes oriented south-southwest was lower than the mean number of nests in boxes facing other directions). There was no such relationship for Blue Tits or Pied Flycatchers. Nestbox orientation influenced the breeding success of Pied Flycatcher (the mean number of young to fledge from boxes oriented south-southwest was lower than from boxes facing other directions). There was no such relationship for Blue or Great Tits.

Conclusion Nestbox orientation can be an important influence on occupation and breeding success, but this differed between species. Intriguingly, although the directionality reduced nestbox occupation (Great Tit) and breeding success (Pied Flycatcher) was the same (south-southwest), there was a disparity in the influence of orientation for Great Tit (orientation influenced the frequency of occupation but not success) and Pied Flycatcher (orientation did not influence occupation but did affect success). We discuss these disparities, considering the possible influences of mating strategy, breeding phenology, nestbox microclimate and offspring quality.     

Artificial diet and rearing methods for Thanasimus dubius (Coleoptera: Cleridae), a predator of bark beetles (Coleoptera: Scolytidae)

Clerid beetles are common natural enemies of bark beetles, and could potentially be used as biological control agents if they could be reared in sufficient numbers. We developed an artificial diet devoid of insect components for rearing Thanasimus dubius (Fabricius), a clerid that attacks several economically important bark beetles in eastern North America. We reared larvae of this predator using the artificial diet, and then used either natural or factitious prey to feed the adults so produced. Two different methods of presenting the diet were also examined. We then compared the performance of T. dubius reared on the artificial diet with newly-emerged wild individuals collected from the field. Our results suggest that adult predators reared on the diet are near in quality to wild ones, and high R₀ values can be obtained. No difference in prey preference was found between wild and diet-reared individuals after five generations in the laboratory. Sufficient numbers of predators could be generated using these techniques to permit limited field trials of augmentative biological control.     

Herbivory below ground and biological weed control: life history of a root-boring weevil on purple loosestrife

The life history of the root-boring weevil Hylobius transversovittatus was studied in north and central Europe. The weevil develops on Lythrum salicaria, a perennial marshland plant that has become a problem weed in North America. It was found in all habitats of its host plant with the exception of permanently flooded sites. It also attacked L. salicaria in an early successional stage, devastating large parts of the storage tissue. The beetle was found in two-thirds of the field populations examined with a mean attack rate of 76.3%. Larvae developed according to a 1- or 2-year generation cycle depending on the time of oviposition. Adult beetles developing within 1 year emerged between July and October, whereas beetles with a 2-year larval period emerged within 3 weeks in July and August. Adults were long-lived and could hibernate several times. Beetles of the new generation mated right on the spot and some oviposited prior to overwintering. Hibernated females had an oviposition period from May to early September and produced 3–4 eggs/day during the peak oviposition period. Specific mortality factors were scarce. Dispersal flights ensure the regular occurrence of H. transversovittatus in scattered L. salicaria populations. The severe impact of the weevil is expected to reduce the competitive ability of its host plant after introduction into North America.     

Small-scale spatial genetic structure in the Central African rainforest tree species Aucoumea klaineana: a stepwise approach to infer the impact of limited gene dispersal, population history and habitat fragmentation

Under the isolation-by-distance model, the strength of spatial genetic structure (SGS) depends on seed and pollen dispersal and genetic drift, which in turn depends on local demographic structure. SGS can also be influenced by historical events such as admixture of differentiated gene pools. We analysed the fine-scale SGS in six populations of a pioneer tree species endemic to Central Africa, Aucoumea klaineana. To infer the impacts of limited gene dispersal, population history and habitat fragmentation on isolation by distance, we followed a stepwise approach consisting of a Bayesian clustering method to detect differentiated gene pools followed by the analysis of kinship-distance curves. Interestingly, despite considerable variation in density, the five populations situated under continuous forest cover displayed very similar extent of SGS. This is likely due to an increase in dispersal distance with decreased tree density. Admixture between two gene pools was detected in one of these five populations creating a distinctive pattern of SGS. In the last population sampled in open habitat, the genetic diversity was in the same range as in the other populations despite a recent habitat fragmentation. This result may due to the increase of gene dispersal compensating the effect of the disturbance as suggested by the reduced extent of SGS estimated in this population. Thus, in A. klaineana, the balance between drift and dispersal may facilitate the maintenance of genetic diversity. Finally, from the strength of the SGS and population density, an indirect estimate of gene dispersal distances was obtained for one site: the quadratic mean parent-offspring distance, sigma(g), ranged between 210 m and 570 m.     

Biology ofTyta luctuosa [Lep.: Noctuidae] and its potential value as a biological control agent for the weedConvolvulus arvensis

The biology of the noctuid,Tyta luctuosa (Denis & Schiffermüller) (Lep.: Noctuidae), a defoliator of field bindweed,Convolvulus arvensis L., was studied in southern Europe.T. luctuosa is widely distributed and feeds on both thick stands and scattered host populations growing in a diversity of habitats. It undergoes 2 and perhaps a partial 3rd generation/year in southern Europe and is active during most of the growing season ofC. arvensis. In the laboratory the total time from egg to adult averages 45.6 days. There are 5 larval instars. Adult females deposit on average over 400 eggs. The larvae being able to feed and develop on native North AmericanCalystegia spp. in the laboratory, there are some reservations about its release in North America. However, the moth has potential value as a biological control agent for field bindeed in the western USA where it would fill an almost unoccupied niche.      

Life table and predation of Oligota pygmaea (Coleoptera: Staphylinidae) a major predator of the red spider mite, Oligonychus coffeae (Acarina: Tetranychidae) infesting tea

The staphylinid beetle, Oligota pygmaea (Solier) is an important predator of the red spider mite, Oligonychus coffeae (Nietner) infesting tea. Biology, life table and predatory efficiency of O. pygmaea were studied under laboratory conditions. Duration of developmental stages of O. pygmaea was 3.2, 5.7 and 12.5 d for eggs, larvae and pupae, respectively with an average of 23.0 d from egg to adult emergence. After a mean preovipostion period of 2.9 d, each female laid an average of 400.5 eggs in its life span. Adult O. pygmaea lived for an average of 54.1 d. Adult females lived for a longer period of 58.8 d compared to the longevity of 49.4 d of adult male. Studies revealed that its life table characterized by an intrinsic rate of natural population increase (r) of 0.118 d, net reproductive rates (R_o) of 243.693 eggs/female, gross reproduction rate (Σm_x) of 245.313 eggs/female, generation time (T) of 46.575 d, doubling time (DT) of 5.874 d and finite rate of increase (λ) of 1.125 d. Seasonal abundance of O. pygmaea and its prey, O. coffeae was monitored by sampling 25 tea leaves randomly from each experimental block grown under the prevailing field conditions. O. pygmaea showed a typical pattern of population dynamics with a peak during January to March and low incidence during June to September. Peak in the population of O. pygmaea coincided with the abundance of O. coffeae in the tea fields. Weather factors such as high temperature, low relative humidity and low sunshine hours adversely affected the populations of O. pygmaea. The first to third instar larvae of O. pygmaea consumed 31.0–133.2 eggs of mites per day. Third instar larva of O. pygmaea consumed an average of 133.2 eggs, 46.4 hexapod larvae, 39.6 nymphs and 11.4 adults per day. Adult females consumed more number of red spider mites compared to the males.     

The potential for indirect effects between a weed, one of its biocontrol agents and native herbivores: A food web approach

The safety of biological control is a contentious issue. We suggest that constructing and analyzing food webs may be a valuable addition to standard biological control research techniques, as they offer a means of assessing the post-release safety of control agents. Using preliminary data to demonstrate the value of food webs in biocontrol programs, we quantified the extent to which a key agent has infiltrated natural communities in Australia and, potentially, impacted on non-target species. Using these data, we also demonstrate how food webs can be used to generate testable hypotheses regarding indirect interactions between introduced agents and non-target species. We developed food webs in communities invaded to varying degrees by an exotic weed, bitou bush, Chrysanthemoides monilifera ssp. rotundata, and a key biocontrol agent for this weed in Australia, the tephritid fly, Mesoclanis polana. Three food webs were constructed during springtime showing the interactions between plants, seed-feeding insects and their parasitoids. One food web was constructed in a plot of native Australian vegetation that was free of bitou bush (‘bitou-free’), another in a plot of Australian vegetation surrounded by an invasion of bitou bush (‘bitou-threatened’) and a third from a plot infested with a monoculture of bitou bush (‘bitou-infested’). The bitou-free web contained 36 species, the bitou-threatened plot 9 species and the bitou-infested web contained 6 species. One native Australian herbivore attacked the seeds of bitou bush. M. polana, a seed-feeding fly, was heavily attacked by native parasitoids, these being more abundant than the parasitoids feeding on the native seed feeders. A surprising result is that none of the three species of native parasitoids reared from M. polana were reared from any of the native herbivores. The food webs revealed how a highly host-specific biocontrol agent, such as M. polana has the potential to change community structure by increasing the abundance of native parasitoids. The webs also suggest that indirect interactions between M. polana and native non-target species are possible, these been mediated by shared parasitoids. The experiments necessary to determine the presence of these interactions are outlined.