Host locating behaviour of Leschenaultia exul and Patelloa pachypyga: two tachinid parasitoids of the forest tent caterpillar, Malacosoma disstria

Leschenaultia exul (Townsend) and Patelloa pachypyga (Aldrich & Webber) (Diptera: Tachinidae) are the principal larval parasitoids of the forest tent caterpillar (FTC) Malacosoma disstria (Hübner) (Lepidoptera: Lasiocampidae) in Canada. The response of these two fly species to M. disstria differs depending on the tree species on which the host feeds. In wind tunnel experiments, L. exul spent more time on the side of the tunnel containing volatiles from FTC frass and was attracted to the FTC-aspen poplar (Populus tremuloides Michx.) complex preferentially to the FTC-balsam poplar (Populus balsamifera L.) complex. Field bioassays confirmed that this fly species was preferentially attracted to the herbivore-aspen poplar complex as compared to the herbivore-balsam poplar complex. In field bioassays, P. pachypyga was also attracted preferentially to aspen poplar trees containing FTC larvae, compared to balsam poplar trees with host larvae.     

Pros and cons of group living in the forest tent caterpillar: separating the roles of silk and of grouping

Group living can incur both benefits and costs, mediated by different mechanisms. In many gregarious caterpillars, collective use of a network of silk trails is thought to improve foraging. Grouping, i.e., close contact with conspecifics, has been postulated to have both positive (thermoregulation and predator defense) and negative (competition and pathogen transmission) effects. The present experiment distinguishes between silk produced by group members and grouping per se in their effects on growth and development of both early and late larval stadia of the forest tent caterpillar [Malacosoma disstria Hübner (Lepidoptera: Lasiocampidae)] in a laboratory context. For both developmental stadia tested, pre‐established silk trails decreased latency to food finding and hence increased food consumption and growth rate. For younger larvae, pre‐established silk also decreased investment in silk production. Grouping young caterpillars accelerated development at the expense of growth, possibly as a mechanism to avoid intraspecific competition in later larval stadia. In older caterpillars, grouping decreased meal duration, suggesting that competition can indeed occur towards the end of larval development, even in the presence of surplus food. This led to a decrease in growth without any effect on instar duration. The benefits of exogenous silk thus decreased during larval development, whereas the costs associated with crowding increased. Ontogenetic shifts in grouping are common in many taxa: the present study is among the first to provide empirical evidence for an adaptive explanation of observed ontogenetic changes in aggregative behavior.     

Fluctuating asymmetry and survival ability in the forest tent caterpillar mothMalacosoma disstria: implications for pest management

Fluctuating asymmetry of the first tarsal segment of the proleg of the forest tent caterpiller mothMalacosoma disstria Hbn. (Lepidoptera: Lasiocampidae) was significantly inversely related to survival ability in the lab. The monitoring of population levels of fluctuating asymmetry could have important implications in pest management of this and other species by providing an indication of the health of a population.     

Generalist predation on forest tent caterpillar varies with forest stand composition: an experimental study across multiple life stages

1. Generalist enemies can regulate low‐density forest insect populations, and are widely considered to cause greater mortality in more diverse habitats. Forest tent caterpillars (Malacosoma disstria Hübner; FTC) are a major defoliator of aspen (Populus tremuloides Micheaux) in the boreal forest, a region with a mosaic of forest stand types. This heterogeneity may influence FTC outbreaks if generalist predation or parasitism differs among stands of different tree composition. 2. Using exclusion experiments we estimate predation and parasitism of FTC across multiple life‐history stages in low‐density populations occupying both aspen (low diversity) and mixedwood stands (high diversity). 3. Arthropod and avian generalist predators were responsible for most natural enemy‐caused mortality of immature FTC, but their relative impacts varied among FTC life‐history stages. Contrary to expectation, predation on late instar larvae and pupae was higher in the less diverse aspen stands and early instar mortality did not differ. 4. By considering multiple life‐history stages, our results provide a more comprehensive view of natural enemy‐caused morality of immature FTC. Because generalist predation on FTC was higher in aspen than in mixedwood stands, we suggest that FTC populations may be slower to reach outbreak levels in aspen stands.     

Nucleopolyhedroviruses of forest and western tent caterpillars: cross-infectivity and evidence for activation of latent virus in high-density field populations

Abstract. 1. Cyclic population dynamics of forest caterpillars are often associated with epizootics of nucleopolyhedrovirus, but it is not known how these viruses persist between generations or through the fluctuations in host population density. 2. To explore the question of virus persistence at different phases of the population cycle, the nucleopolyhedroviruses of two species of tent caterpillar that co‐occur in British Columbia, Canada, Malacosoma californicum pluviale (western tent caterpillar) and Malacosoma disstria (forest tent caterpillar), were characterised. The cross‐infectivity of the viruses in these two host species was investigated to determine whether there might be a route for virus persistence via the alternative host species. Any virus produced in the cross‐infections was characterised to confirm true cross‐infection or to ascertain whether cross‐inoculation triggered latent virus persisting within the population. 3. The virus associated with forest tent caterpillars (MadiNPV) did not infect western tent caterpillars from low‐density populations, nor did it trigger a latent virus infection; however, inoculation of forest tent caterpillars from high‐density populations with virus from western tent caterpillars (McplNPV) resulted in viral infection, but without a dose–response relationship. 4. Analysis of DNA profiles of virus resulting from cross‐infection of the forest tent caterpillar with McplNPV, revealed that 88% of these infections were caused by MadiNPV rather than McplNPV; however the virus from all 44 infected individuals was identical and differed in DNA profile from the stock MadiNPV used for cross‐infection. This suggests strongly that forest tent caterpillars from high‐density field populations harbour a latent, persistent, or sublethal form of MadiNPV that was triggered by exposure to nucleopolyhedrovirus from the western tent caterpillar. 5. Virus was not activated in western tent caterpillars collected over 2 years of late population decline and the first year of population increase.     

The effect of variation in host plant on the growth of an oligophagous insect, Malacosoma americanum and its polyphagous relative, Malacosoma disstria

Larvae of Malacosoma americanum (F.)(Lepidoptera: Lasiocampidae) an oligophagous species that feeds primarily on Prunus and other rosaceous trees, were compared to larvae of the more highly polyphagous congener M. disstria Hb., with respect to their sensitivity to variation in the foliage of a common host plant, Prunus serotina Ehrh. Pupal weight and time to pupation were measured on larvae reared on foliage from open-grown and from shaded saplings. The difference in foliage had a pronounced effect, but no difference was evident between the species in their response to the variation in foliage. This comparison implies that sensitivity to intraspecific variation in host quality does not differ between host-specific and generalized species. However, results from other species suggest that some species of insects do differ in this respect.

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Zusammenfassung Raupen von Malacosoma americanum (F.), einer oligophagen Art, die sich vor allem auf Prunus und andern baumartigen Rosaceen entwickelt, wurden verglichen mit Raupen der polyphageren Verwandten M. disstria Hb. und zwar im Hinblick auf deren Empfindlichkeit auf Unterschiede im Blatt ihrer gemeinsamen Wirtspflanze, Prunus serotina Ehrh. Das Puppengewicht und die Entwicklungszeit bis zur Verpuppung wurden gemessen bei Raupen, welche auf Blättern von freiwachsenden und von beschatteten Jungpflanzen gezüchtet worden waren. Die Blattunterschiede hatten eine ausgesprochene Wirkung, aber es gab keine Unterschiede in der Reaktion der beiden Arten. Dieser Vergleich lässt vermuten, dass die Empfindlichkeit auf intraspezifische Unterschiede der Wirtspflanzenqualität bei wirtsspezifischen und polyphagen Arten gleich ist. Indessen dürften sich laut anderen Resultaten einige Insektenarten anders verhalten.

     

The understanding of industrial melanism in the peppered moth (Biston betularia) (Lepidoptera: Geometridae)

Melanic polymorphism in B. betularia has been extensively studied. Correlations between high melanic frequency and high levels of air pollution have been demonstrated. Kettlewell and others have shown that differential bird predation has an important effect on the maintenance of the polymorphism, and coefficients of visual selection have been obtained on the assumption that the moth habitually rests on tree trunks. Computer models based on these selective coefficients show that they are not sufficient accurately to explain observed melanic frequencies. Other non-visual selective factors and weak frequency-dependent selection have been invoked to improve fits. Analysis of the resting positions of moths recorded in the wild demonstrates that B. betularia does not usually rest in exposed positions on tree trunks, but rather rests on the underside of branches, on trunks in shaded positions just below major branch joints or on foliate twigs. The results of a pilot selection experiment, while agreeing qualitatively with Kettlewell's results, suggest that fitness estimates that assume trunk-resting are quantitively incorrect. The error is greatest for melanic moths in rural areas. It is suggested that visual selective coefficients based on a true assessment of the resting behaviour of the moths may considerably improve the fit between computer predictions and observed phenotype frequency distributions.     

The danger within: the role of genetic,behavioural and ecological factors in population persistence of colour polymorphic species

Polymorphic species have been the focus of important work in evolutionary biology. It has been suggested that colour polymorphic species have specific evolutionary and population dynamics that enable them to persist through environmental changes better than less variable species. We suggest that recent empirical and theoretical work indicates that polymorphic species may be more vulnerable to extinction than previously thought. This vulnerability arises because these species often have a number of correlated sexual, behavioural, life history and ecological traits, which can have a simple genetic underpinning. When exacerbated by environmental change, these alternate strategies can lead to conflict between morphs at the genomic and population levels, which can directly or indirectly affect population and evolutionary dynamics. In this perspective, we identify a number of ways in which the nature of the correlated traits, their underpinning genetic architecture, and the inevitable interactions between colour morphs can result in a reduction in population fitness. The principles illustrated here apply to all kinds of discrete polymorphism (e.g. behavioural syndromes), but we focus primarily on colour polymorphism because they are well studied. We urge further empirical investigation of the genetic architecture and interactions in polymorphic species to elucidate the impact on population fitness.     

The evolution of phenotypic plasticity in spatially structured environments: implications of intraspecific competition, plasticity costs and environmental characteristics

We model the evolution of reaction norms focusing on three aspects: frequency-dependent selection arising from resource competition, maintenance and production costs of phenotypic plasticity, and three characteristics of environmental heterogeneity (frequency of environments, their intrinsic carrying capacity and the sensitivity to phenotypic maladaptation in these environments). We show that (i) reaction norms evolve so as to trade adaptation for acquiring resources against cost avoidance; (ii) maintenance costs cause reaction norms to better adapt to frequent rather than to infrequent environments, whereas production costs do not; and (iii) evolved reaction norms confer better adaptation to environments with low rather than with high intrinsic carrying capacity. The two previous findings contradict earlier theoretical results and originate from two previously unexplored features that are included in our model. First, production costs of phenotypic plasticity are only incurred when a given phenotype is actually produced. Therefore, they are proportional to the frequency of environments, and these frequencies thus affect the selection pressure to avoid costs just as much as the selection pressure to improve adaptation. This prevents the frequency of environments from affecting the evolving reaction norm. Secondly, our model describes the evolution of plasticity for a phenotype determining an individual's capability to acquire resources, and thus its realized carrying capacity. When individuals are distributed randomly across environments, they cannot avoid experiencing environments with intrinsically low carrying capacity. As selection pressures arising from the need to improve adaptation are stronger under such extreme conditions than under mild ones, better adaptation to environments with low rather than with high intrinsic carrying capacity results.     

Population dynamics in changing environments: the case of an eruptive forest pest species

In recent decades we have seen rapid and co‐occurring changes in landscape structure, species distributions and even climate as consequences of human activity. Such changes affect the dynamics of the interaction between major forest pest species, such as bark beetles (Coleoptera: Curculionidae, Scolytinae), and their host trees. Normally breeding mostly in broken or severely stressed spruce; at high population densities some bark beetle species can colonise and kill healthy trees on scales ranging from single trees in a stand to multi‐annual landscape‐wide outbreaks. In Eurasia, the largest outbreaks are caused by the spruce bark beetle, Ips typographus (Linnaeus), which is common and shares a wide distribution with its main host, Norway spruce (Picea abies Karst.). A large literature is now available, from which this review aims to synthesize research relevant for the population dynamics of I. typographus and co‐occurring species under changing conditions. We find that spruce bark beetle population dynamics tend to be metastable, but that mixed‐species and age‐heterogeneous forests with good site‐matching tend to be less susceptible to large‐scale outbreaks. While large accumulations of logs should be removed and/or debarked before the next swarming period, intensive removal of all coarse dead wood may be counterproductive, as it reduces the diversity of predators that in some areas may play a role in keeping I. typographus populations below the outbreak threshold, and sanitary logging frequently causes edge effects and root damage, reducing the resistance of remaining trees. It is very hard to predict the outcome of interspecific interactions due to invading beetle species or I. typographus establishing outside its current range, as they can be of varying sign and strength and may fluctuate depending on environmental factors and population phase. Most research indicates that beetle outbreaks will increase in frequency and magnitude as temperature, wind speed and precipitation variability increases, and that mitigating forestry practices should be adopted as soon as possible considering the time lags involved.     

Temperature and population density: interactional effects of environmental factors on phenotypic plasticity,immune defenses,and disease resistance in an insect pest

Temperature and crowding are key environmental factors mediating the transmission and epizooty of infectious disease in ectotherm animals. The host physiology may be altered in a temperature‐dependent manner and thus affects the pathogen development and course of diseases within an individual and host population, or the transmission rates (or infectivity) of pathogens shift linearly with the host population density. To our understanding, the knowledge of interactive and synergistic effects of temperature and population density on the host–pathogen system is limited. Here, we tested the interactional effects of these environmental factors on phenotypic plasticity, immune defenses, and disease resistance in the velvetbean caterpillar Anticarsia gemmatalis. Upon egg hatching, caterpillars were reared in thermostat‐controlled chambers in a 2 × 4 factorial design: density (1 or 8 caterpillars/pot) and temperature (20, 24, 28, or 32°C). Of the immune defenses assessed, encapsulation response was directly affected by none of the environmental factors; capsule melanization increased with temperature in both lone‐ and group‐reared caterpillars, although the lone‐reared ones presented the most evident response, and hemocyte numbers decreased with temperature regardless of the population density. Temperature, but not population density, affected considerably the time from inoculation to death of velvetbean caterpillar. Thus, velvetbean caterpillars succumbed to Anticarsia gemmatalis multiple nucleopolyhedrovirus (AgMNPV) more quickly at higher temperatures than at lower temperatures. As hypothesized, temperature likely affected caterpillars' movement rates, and thus the contact between conspecifics, which in turn affected the phenotypic expression of group‐reared caterpillars. Our results suggest that environmental factors, mainly temperature, strongly affect both the course of disease in velvetbean caterpillar population and its defenses against pathogens. As a soybean pest, velvetbean caterpillar may increase its damage on soybean fields under a scenario of global warming as caterpillars may reach the developmental resistance faster, and thus decrease their susceptibility to biological control by AgMNPV.     

Factors affecting the initiation and growth of aboveground adventitious roots in a tropical cloud forest tree: an experimental approach

Some of the proximate factors that would induce aboveground stems to produce adventitious roots were investigated experimentally on Senecio cooperi, a tropical cloud forest tree. Stem segments were air-layered with different treatments to promote root formation, and the number of roots initiated and rates of root growth were monitored for 20 weeks. Treatments were the application of wet epiphytes or dry epiphytes plus associated humus, sponges wetted with either water or nutrient solutions, or dry sponges. Controls (stem segments with nothing applied) were also monitored. Numbers of adventitious roots formed and rates of subsequent root growth differed among treatments. Wet epiphyte/humus and nutrient solutions were most effective in producing roots, which suggests that epiphytes and the nutrients they intercept and retain within the canopy may cue adjacent host tissue to exploit this resource.     

Population density and group composition of Japanese sika deer (<Emphasis Type="Italic">Cervus nippon yakushimae</Emphasis>) in an evergreen broad-leaved forest in Yakushima,southern Japan

The population density of Japanese sika deer (Cervus nippon yakushimae Kuroda and Okada) in an evergreen broad-leaved forest in Yakushima, southern Japan, was surveyed over 4years from 1998 to 2001. Two approximately 50ha study sites, Hanyama and Kawahara, were established with a total of 4km of census trails at each site. The estimated densities of sika deer at the two sites were 43–70 deerkm^–2 at Hanyama and 63–78 deerkm^–2 at Kawahara, although these values might be underestimates. The adult sex ratio (number of adult males:number of adult females) ranged from 0.6 to 1.0 at Hanyama, and from 0.4 to 0.9 at Kawahara. Mean group size was 1.9 deer (male group, 1.5 deer; female group, 1.6 deer; mixed group, 3.6 deer). The population density of sika deer was relatively high compared to other sites in Japan, with the exception of very small (<10km²) islands. Possible explanations for this naturally high density of sika deer in an evergreen broad-leaved forest in Yakushima are discussed.     

特大干旱对树木死亡的影响——以美国德克萨斯州东部森林为例

美国德克萨斯州在2011年经历了史上最严重的干旱,这一事件造成约3亿多株树木死亡。在大时空尺度上(面积约9×10~6 hm~2,时间跨度近20年),基于近1800个森林样地,4次周期性调查中的约209663株树木,使用主成份分析(PCA)和广义线性混合效应模型(GLM)回归,对树木死亡的时空差异及其干旱强度与长度对树木死亡造成的中长期复杂影响进行了研究。采用树木密度、树木基面积、林地年龄、样地调查时间间隔作为树木间的竞争指标,分析了造成大旱前后周期水平和年度水平上的样地树木死亡差异的原因。综合分析了不同地理区域、树木种组、胸径大小和林地起源的4个划分标准下树木死亡对死亡率的相对贡献。结果表明：松属树木的死亡率最低(7.92%);高度低、胸径小的树木的死亡率较大,分别为29.79%和26.00%。人工林的树木死亡率(10.26%)低于天然林(13.47%);西海湾平原生态区树木的死亡率在干旱后达到最大(22.27%);西南区的树木死亡率在干旱后也达到最大(13.78%);海拔和纬度对树木死亡率影响不明显。德州东部森林整体死亡格局形成原因较为复杂,各地理区域、林地起源、树木大小和不同树种,...     

The influence of sex gland components on mating by the summerfruit tortrix moth, Adoxophyes orana, in the laboratory

Couples of Adoxophyes orana F.v.R. were isolated for one week in small 1.25 l rearing boxes in the laboratory with sex gland components deposited on rubber caps. At the end of the trials females were dissected and spermatophores counted. Reduction of mating increased with the dose of a mixture of the attractant components Z9-14:Ac and Z11-14:Ac (9:1) but decreased with increasing density of 1 to 5, 10 and 20 couples per box. Each of the attractant components alone produced a mating inhibition equal to the mixture. The minor components Z9-12:Ac and 12:Ac had approximately the same effect as the main components, but 14:Ac and Z11-14:OH had no effect.Identical trials conducted on a strain of summerfruit moth newly introduced to our rearing showed that ten times less product was needed to achieve the same degree of mating inhibition as with the old strain. With adaptation to laboratory conditions the proportion of unmated females diminished and repeated matings occurred more frequently. In order to achieve meaningful results one must work with material from regularly renewed rearings, or better with the direct descendants of individuals from orchards. The results are discussed in relation to mating disruption of A. orana in orchards.

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Etude en laboratoire de l'influence de six composantes phéromonales de la tordeuse de la pelure Adoxophyes orana sur les accouplements

Résumé Des essais ont été effectués en laboratoire en mettant des couples de papillons d'Adoxophyes orana F.v.R. pendant une semaine dans des boîtes d'élevage de 1,25 l avec ou sans composantes phéromonales. A la fin de l'essai, les femelles sont disséquées pour la recherche des spermatophores. L'inhibition des accouplements s'accroît avec l'augmentation de la charge des deux composantes principales de l'attractif, le Z9-14:Ac et le Z11-14:Ac (9:1) et elle décroît lorsque la densité de popultion passe de 1 à 5, 10, 20 couples de papillons par boîte. Chacune de ces deux composantes principales isolée provoque une réduction d'accouplements aussi importante que le mélange lui-même. Les composantes mineures Z9-12:Ac et 12:Ac ont pratiquement la même efficacité que les composantes principales, mais le 14:Ac et le Z11-14:OH sont sans effet.Des essais identiques effectués avec une souche d'A. orana récemment introduite en élevage continu, montrent qu'il suffit d'environ dix fois moins d'attractif pour provoquer le même effet que sur l'ancienne souche. Parallèlement à l'adaptation aux conditions d'élevage, le nombre de femelles non accouplées dans les témoins diminue progressivement et la fréquence des accouplements multiples augmente. II est donc vivement recommandé de travailler avec des souches continuellement renouvelées en élevage, ou avec les descendants directs d'individus prélevés en vergers. Les résultats obtenus sont commentés en relation avec la lutte par la technique de confusion.

     

The effects of regeneration by fragmentation upon clonal diversity in the tropical forest shrub Poikilacanthus macranthus: random amplified polymorphic DNA (RAPD) results

Poikilacanthus macranthus (Acanthaceae), like many shrubs in the neotropical cloud forest of Monteverde, Costa Rica, is capable of regeneration through stem and twig fragments. Our field sampling indicated that reproduction in P. macranthus is primarily through fragmentation in the Monteverde region. Random amplified polymorphic DNAs (RAPDs) were employed to examine clonal diversity in four populations of P. macranthus , and two primers yielded 11 variable markers that were used to identify genets. Sixty-eight multilocus genotypes were identified out of 277 plants sampled. Clonal diversity values were higher than the average for clonal plant species, as the number of genets per ramet was 0.25, while the mean D value was 0.87 and the mean E value was 0.84.     

从恢复生态学视角透析防护林衰退及其防治对策：以章古台地区樟子松林为例

针对防护林衰退问题,以章古台樟子松固沙林为案例,应用生态演替、干扰、种群密度等关键性恢复生态学理论和原则,对防护林在决策层面的设计要素的科学性及其与衰退的关系进行了分析和评价,认为大面积造林与地带性顶极的不吻合（即偏离生态学原则）,使防护林建设的目标、步骤、树种的组成和密度等出现了偏颇,成为防护林衰退的最深层次的原因,疏于管理和粗放经营以及频频发生的自然和人为干扰也是致衰的重要因素;据此提出深化对受损生态系统的认识、强化物质和能量的投入、建立干扰的防控体系等对策.文中还就人工植被建设目标与生态演替顶极类型的判别,以及偏途顶极在人工植被建设中的应用问题进行了讨论,一并作为对策以供参考.     

Bird richness and composition along an agricultural gradient in New Guinea: The influence of land use,habitat heterogeneity and proximity to intact forest

Abstract We examined variation in bird species richness, abundance and guild composition along an agricultural gradient in New Guinea, and looked for any additive influence of habitat heterogeneity on these variables. The study was based on a grid of survey plots, six plots wide and 24 plots long with the long axis running from a settlement 2.4 km through active and abandoned agricultural plots towards a large area of forest. Each circular survey plot (25 m radius) was assigned to a broad habitat type, ten habitat measures taken, and birds counted for 1 h in each plot. Principal component analysis (PCA) habitat axis 1 described an axis of decreasing forest alteration (larger trees, greater tree densities, fuller canopy) that was positively correlated with distance from the settlement. Bird richness and abundance were highest at intermediate disturbance levels (plots with mid‐range axis 1 scores). Proportions of insectivores and frugivores increased with decreasing forest alteration, while proportions of nectarivores decreased. We calculated three measures of habitat heterogeneity by comparing each plot's PCA score to those of eight neighbouring plots (50–110 m away). These measures reflected how different the plot was to its neighbours, how variable the habitat was around the plot, and the degree to which the plot bordered less disturbed forest. We related these measures to plot bird variable scores independently, and to residuals following regressions of bird scores against PCA scores. Heterogeneity measures had no significant influence on abundance or richness measures, but there were greater proportions of frugivores in plots showing a given degree of habitat alteration if they bordered more pristine habitat. While we readily identified differences in bird communities along the agricultural gradient, the influences of habitat heterogeneity were not striking for birds at this fine scale.