Delayed density-dependent parasitism of eggs and pupae as a contributor to the cyclic population dynamics of the autumnal moth

Many populations of forest Lepidoptera exhibit 10-year cycles in densities, with impressive outbreaks across large regions. Delayed density-dependent interactions with natural enemies are recognized as key factors driving these cyclic population dynamics, but emphasis has typically been on the larval stages. Eggs, pupae and adults also suffer mortality from predators, parasitoids and pathogens, but little is known about possible density relationships between mortality factors and these non-feeding life stages. In a long-term field study, we experimentally deployed autumnal moth (Epirrita autumnata) eggs and pupae to their natural enemies yearly throughout the 10-year population cycle in northern Norway. The abundance of another geometrid, the winter moth (Operophtera brumata), increased in the study area, permitting comparisons between the two moth species in predation and parasitism. Survival of autumnal moth eggs and pupae was related to the moth abundance in an inverse and delayed manner. Egg and pupal parasitoids dominated as density-dependent mortality factors and predicted the subsequent growth rate of the host population size. In contrast, effects of egg and pupal predators were weakly density dependent, and generally predation remained low. Parasitism rates did not differ between the autumnal and winter moth pupae, whereas predators preferred winter moth pupae over those of the autumnal moth. We conclude that parasitism of the autumnal moth by egg and pupal parasitoids can be related to the changes of the moth density in a delayed density-dependent manner. Furthermore, egg and pupal parasitoids cannot be overlooked as causal factors for the population cycles of forest Lepidoptera in general.     

Flowers for better pest control? Effects of apple orchard groundcover management on mites (Acari), leafminers (Lepidoptera,Scitellidae), and fruit pests

Flowering companion plants provide nectar, pollen, alternative prey, shelter, and overwintering habitat for arthropod natural enemies and thus might increase their abundance and efficacy in pest control in agricultural fields. We report the results of a 6-year study on the effects of annual and perennial flowering herbs sown in alleys of an apple orchard on phytophagous and predacious mites, leafminer moths and their parasitoids and on pests causing fruit injury. Plots with weed-free bare ground and with regularly mowed grass served as controls. The abundance and diversity of predatory phytoseiid mites increased in the presence of flowering ground cover plants. However, this positive effect on phytoseiids was confined only to spring and autumn and intraguild predation (Typhlodromus pyri gradually displaced Amblyseius andersoni in the presence of flowers) also constrained their enhancement. Spider mites’ (Tetranychidae) abundance was low in all years and did not increase with the alley herb coverage. Incidence of the leafminer Leucoptera malifoliella was similar among the treatments despite the higher parasitism rate and parasitoid diversity in the plot with flowering herbs. We demonstrated that in the presence of flowering ground cover plants, the sex ratio of eulophid parasitoids' (mainly Chrysocharis pentheus) reared from L. malifoliella larvae and pupae shifted towards a male bias. Ground cover management had no effect on fruit injury caused by codling moth (Cydia pomonella) and summer fruit tortrix moth (Adoxophyes orana) and on the percentage of apples without insect damage.     

Habitat use of Bornean Orangutans (<Emphasis Type="Italic">Pongo pygmaeus morio</Emphasis>) in an Industrial Forestry Plantation in East Kalimantan,Indonesia

Many primates now live in anthropogenic landscapes dominated by human activity such as agriculture. Conserving primates in such contexts requires detailed information about habitat use, including landscape features that may influence population viability. We studied Northeast Bornean orangutan (Pongo pygmaeus morio) habitat use in a forestry plantation in East Kalimantan, Indonesia. We conducted camera trapping and nest surveys at 13 locations across three habitat types in the plantation (planted acacia stands, planted eucalyptus stands, and secondary forest patches left uncut or allowed to regenerate) September 2012–March 2013, and calculated four measures of orangutan abundance for each location (independent photo captures/100 camera trap days, or RAI₂; nest encounter rate; nest density; and orangutan density). Orangutans are relatively common in the plantation; they used all three habitat types and exhibited a higher RAI₂ than 70% of other mammal species detected. A logistic regression found that proximity to natural forest areas best predicted orangutan abundance calculated using camera trap data (RAI₂) but that habitat type combined with distance to natural forest best predicted orangutan abundance calculated using nest counts. This suggests that orangutans use planted areas for movement and feeding, but rely on patches of natural forest for resting and access to key resources. Our study and others indicate that orangutans can coexist with some human activities if provided with sufficient access to natural forest. However, we must conduct further research to facilitate effective conservation planning, including gathering additional details about habitat and resource use and possible long-term population impacts.     

Spruce budworm impact, abundance and parasitism rate in a patchy landscape

The hypothesis that vegetational diversity may lessen the impact of forest insect pests by favoring natural enemies is appealing to those who seek ecologically sound solutions to pest problems. We investigated the effect of forest diversity on the impact of the spruce budworm Choristoneurafumiferana following the last outbreak, as well as the budworm's current abundance and parasitism rate, in the boreal forest of northwestern Québec. Mortality of balsam fir caused by the budworm was greater in extensive conifer stands than either in “habitat islands” of fir surrounded by deciduous forest or on true islands in the middle of a lake. Adult spruce budworm abundance, assessed by pheromone traps, did not differ significantly between the three types of sites. Larval and pupal parasitism rates were examined by transferring cohorts of laboratory-reared larvae and pupae to trees in the three site types and later collecting and rearing them. The tachinid Actiainterrupta, a parasitoid of fifth and sixth instar larvae, as well as the ichneumonid pupal parasitoids Itoplectesconquisitor, Ephialtesontario and Phaeogenesmaculicornis, caused higher mortality in the habitat islands than on true islands or in extensive stands. Exochusnigripalpistectulum, an ichneumonid that attacks the larvae and emerges from the pupae, caused greater mortality in the extensive stands of conifers. Received: 17 March 1997 / Accepted: 18 November 1997     

Impact of predators on artificially augmented populations of Lymantria dispar L. pupae (Lep., Lymantriidae)

The impact of predators on artificially augmented populations of Lymantria dispar pupae (Lep., Lymantriidae) and the species composition of the predator community were investigated in two oak stands in eastern Austria. The population density of L. dispar has been at innocuous levels for several years at both sites. From mid-June to mid-July, we created artificial prey populations by mounting gypsy moth pupae with beeswax on burlap bands placed at the base, 25, 50 and 100 cm height on 25 trees at each site. A total of 6600 pupae were exposed at each site. A total of 92% of exposed pupae were destroyed by predators, at site I (with dense understorey vegetation) whereas 67% were destroyed on site II (with sparse understorey vegetation). Initially, pupal mortality was highest at the base of trees, but differences in mortality among the four locations where pupae were exposed decreased during the course of the study period. Spatial differences in predation were ascertained, suggesting that the aggregation of small mammals, for example, is related to patches of dense shrub vegetation. Pupal mortality was primarily caused by mice (46.3% at site I and 36.4% at site II). Many exposed pupae simply disappeared (40.4% at site I and 22.8% at site II). Calosoma spp. and other invertebrate predators caused very little mortality among L. dispar pupae (5.5% at site I and 7.8% at site II). Trap catches on separate study plots at both sites revealed that Apodemus flavicollis (Rodentia, Muridae) was the most important predator species present, whereas Apodemus sylvaticus (Rodentia, Muridae) and Clethrionomys glareolus (Rodentia, Arvicolidae) were captured in low numbers.     

The influence of extrafloral nectaries on parasitism of an insect herbivore

The extrafloral nectaries of many plants promote ant defense against insect herbivores. We examined the influence of extrafloral nectaries on the levels of parasitism of a generalist insect herbivore, the gypsy moth (Lymantria dispar L.). Larvae and pupae of the moth were collected from trees with and without extrafloral nectaries growing in the same forests in South Korea and reared to evaluate parasitism. More parasitism occurred on plants with extrafloral nectaries in seven of the nine season-long collections at the six sites and in four out of five collecting periods. Parasitism was higher on the four main genera of plants with extrafloral nectaries than on any of five main genera of plants without extrafloral nectaries. There was no difference in parasitoid richness; nine species occurred in each group, eight of which were the same. There was a positive and almost significant correlation between the abundance of plants with extrafloral nectaries and the parasitism of gypsy moth at the sites. Extrafloral nectaries may reduce herbivory by inducing more parasitism of the insect herbivores that attack plants bearing the glands.     

An ecological study ofPedicularis dendrothauma, an arboreal hemiparasitic epiphyte from Nepal

Pedicularis dendrothauma is an arboreal epiphytic hemiparasite that was recently discovered in Nepal. We present results of preliminary studies on its habitat, demography, hemiparasitism, flower development, and pollination. While showing a strong preference forAbies spectabilis trunks and branches as a substrate,P. dendrothauma also grows on other species of trees and shrubs in the understorey as well as on dead wood on the forest floor, and only very rarely is rooted in soil or on rocks, rooted in a bryophyte layer. Demographic monitoring of the population in 1999 and 2001 allows us to begin to predict longevity, average size of flowering plants, and survivorship and flowering probability on different substrates. We demonstrate that individual plants ofP. dendrothauma live at least four, and probably at least five years. Studies of haustoria show that the species has retained its hemiparasitic habit despite having moved from a terrestrial to an arboreal habitat. Flowering probability increases with plant size; an average flowering plant has five leaves and a longest leaf that measures 21.5 cm. Survivorship varies from substrate to substrate, with the greatest survivorship over a two year period occurring in plants growing on living trees. Flowering probability was the same for plants growing on living trees as it was for plants on dead wood. Studies of flower development suggest that plants growing high in the canopy proceeded through the stages of floral development more rapidly than plants nearer ground level. Despite many hours of field observation, the method of pollination remains a mystery; we suggest different possible pollination mechanisms.     

Fine-scale abundance and distribution of wild silk moth pupae

Although several factors influence herbivore insect distributions at any particular scale, the most important determinants are likely to differ between species with different life histories. Identifying what these factors are and how they relate to life history forms an important component of understanding the population dynamics of species, and the habitat requirements necessary for their conservation. The pupal stage of two wild silk moth species, Gonometa postica Walker and G. rufobrunnea Aurivillius (Lepidoptera: Lasiocampidae), is the target of harvesting practices that are totally dependent on the availability of pupae from natural populations. Consequently, and partly due to poor knowledge of the species' biology, there is substantial interest in the distribution of pupae among and within trees for both these species. It was investigated whether between- and within-tree pupal distributions in these two species are non-random, and if so, whether there are relationships between pupation site use and tree characteristics such as tree size, available pupation space and branch position. Between-tree patterns in pupal abundance were random in terms of absolute spatial position, but markedly non-random with respect to tree characteristics. The apparent G. postica pupae were aggregated on large larval host plants, whereas the cryptic G. rufobrunnea pupae were aggregated on non-host plants. These patterns reflect the life history differences of the two species. In contrast, at the within-tree scale, branch position, aspect and tree shape influenced pupation site choice similarly for both species. These patterns might be related to microclimate. Documenting between-tree and within-tree patterns in Gonometa pupal distributions is the first step towards explaining pupation site selection, as well as identifying possible evolutionarily selective factors in the species, and generating testable hypotheses from these.     

From Forest to Farmland: Species Richness Patterns of Trees and Understorey Plants along a Gradient of Forest Conversion in Southwestern Cameroon

Vegetation surveys were carried out at 24 sampling stations distributed over four land use types, namely near-primary forest, secondary forest, agroforestry systems and annual crop lands in the northeastern part of the Korup region, Cameroon, to assess the impact of forest conversion on trees and understorey plants. Tree species richness decreased significantly with increasing level of habitat modification, being highest and almost equal in secondary and near-primary forests. Understorey plant species richness was significantly higher in annual crop lands than in other land use types. The four land use types differed in tree and understorey plant species composition, the difference being smaller among natural forests. Tree and understorey plant density differed significantly between habitat types. Density was strongly correlated with species richness, both for trees and understorey plants. Five tree and 15 understorey plant species showed significant responses to habitat. A 90% average drop in tree basal area from forest to farmland was registered. Our findings support the view that agroforestry systems with natural shade trees can serve to protect many forest species, but that especially annual crop lands could be redesigned to improve biodiversity conservation in agricultural landscapes of tropical rainforest regions.     

Patterns and monitoring of Sesia apiformis infestations in poplar plantations at different spatial scales

Poplar plantations are expanding worldwide but little is known about the hornet clearwing moth, Sesia apiformis, one of their most severe pests. Thirty‐two poplar plantations of I‐214 clone were sampled in Spain, according to a factorial design combining stand age, site condition and understorey management to investigate the main drivers of S. apiformis habitat selection at both tree and stand level. In each plantation, one pheromone trap was activated during the flight season to test the correlation between captures and percentage of attacked trees. The proportion of other poplar forests in surrounding landscapes was calculated and used as a covariate in predictive models of trap catches. There were significantly more attacked trees in older stands. In young stands, the percentage of infested trees increased with the percentage cover of understorey vegetation. There was no significant effect of site quality on the rate of infestation and no difference in tree height, diameter or crown condition between attacked and un‐attacked trees within each stand, suggesting that S. apiformis could behave as primary pest. We hypothesized that the critical stage in the life cycle of the moth was the first instar larvae, which may benefit from protection of deep bark cracks in older stands and understorey vegetation in younger stands. We observed a positive correlation between trap capture and percentage of attacked trees in a radius of 100 m around the trap. The regression of trap catches against percentage of attacked trees was improved when the area of other poplar plantations within a distance of 600 m was incorporated in the model. This suggests that surrounding poplar stands may act as sources of immigrating moths in monitored stands. Our findings confirm that S. apiformis should be considered as a potential threat to poplar plantations and that pheromone trapping provides a suitable monitoring tool.     

Manipulating carabid beetle abundance alters prey removal rates in corn fields

Generalist natural enemies such as carabid beetles have the potential to maintain a variety of pests below outbreak levels in annual crops. To assess the relationship between carabid beetle abundance and field rates of prey removal, we created plots surrounded by different boundaries that selectively affected dispersal of edaphic arthropods, primarily carabids. Three treatments were established: (1) naturally occurring communities, (2) augmented communities using ingress boundaries, and (3) reduced communities using egress boundaries. Selective boundaries altered carabid communities with minimal habitat alteration and without use of insecticides. Three times during the growing season, a fixed number of onion fly pupae were placed in plots to evaluate the impact of carabid abundance on predation rates. A combination of vertebrate and invertebrate exclosures allowed us to evaluate prey removal by invertebrates alone. In comparison to the no boundary treatment, carabids increased 54.2% and decreased 83.1% in plots surrounded by ingress and egress boundaries respectively. Predation rates were positively correlated with carabid abundance (r² = 0.70, p < 0.0001). Significantly more pupae were removed from exclosures allowing access to invertebrates alone than from total exclosures, suggesting that invertebrates represented an important group of predators. Laboratory trials tested the feeding potential of the four most abundant carabid species and showed that they readily consumed onion fly pupae, supporting our hypothesis that carabids were the main predators in field tests. This study corroborates and extends previous observations of the importance of carabid beetles as generalist predators of insect pests in agricultural fields.     

Crop domestication disrupts a native tritrophic interaction associated with the sunflower, Helianthus annuus (Asterales: Asteraceae)

Abstract.  1. Crop domestication has been a largely overlooked factor that may explain why insect herbivores tend to be more abundant and less attacked in agricultural habitats than in native habitats. This study explores how domestication of the sunflower, Helianthus annuus L., affects the sunflower moth, Homoeosoma electellum (Lepidoptera: Pyralidae), and its parasitoids.
2. Common garden experiments were used to assess the effect of domestication on larval abundance in an agricultural and a native habitat. The effect of domestication on parasitism was determined using two studies: one study that manipulated nitrogen according to levels found in agricultural and native habitats, and a second common garden study in the native habitat.
3. At peak infestation in the agricultural common garden, larval abundance was 10 times higher on agricultural plants than on wild plants. In contrast, larval abundance did not differ between plant genotypes in the native habitat.
4. Larvae were four times more likely to be parasitised on wild sunflowers than on agricultural sunflowers, and three times more likely to be parasitised on low nitrogen plants than on high nitrogen plants. Parasitism did not differ between agricultural and wild plants in the native habitat, where flowers were more similar in size.
5. Sunflower domestication has increased larval abundance, accelerated larval development, and lowered parasitism. The magnitude of the effect appears to depend upon nutrient availability. Thus, domestication can disrupt tritrophic interactions, and may help explain why some insect pests are more abundant and less attacked in agro-ecosystems.     

Greenhouse and field evaluations of transgenic canola against diamondback moth, shape Plutella xylostella, and corn earworm, shape Helicoverpa zea

Canola (Brassica napus L.) cultivars Oscar and Westar, engineered with a Bacillus thuringiensis (Bt) cryIA(c) gene, were evaluated for resistance to lepidopterous pests, diamondback moth, Plutella xylostella L. (Plutellidae) and corn earworm, Helicoverpa zea (Boddie) (Noctuidae) in greenhouse and field conditions. In greenhouse preference assays conducted at vegetative and flowering plant stages, transgenic plants recorded very low levels of damage. A 100% diamondback moth mortality and 90% corn earworm mortality were obtained on transgenic plants in greenhouse antibiosis assays. The surviving corn earworm larvae on transgenic plants had reduced head capsule width and body weight. Mortality of diamondback moth and corn earworm were 100% and 95%, respectively, at different growth stages (seedling, vegetative, bolting, and flowering) on the transgenic plants in greenhouse tests. In field tests conducted during 1995–1997, plots were artificially infested with neonates of diamondback moth or corn earworm or left for natural infestation. Transgenic plants in all the treatments were highly resistant to diamondback moth and corn earworm larvae and had very low levels of defoliation. Plots infested with diamondback moth larvae had greater damage in both seasons as compared with corn earworm infested plots and plots under natural infestation. After exposure to defoliators, transgenic plants usually had higher final plant stand and produced more pods and seeds than non-transgenic plants. Diamondback moth injury caused the most pronounced difference in plant stand and pod and seed number between transgenic and non-transgenic plants. Our results suggest that transgenic canola could be used for effective management of diamondback moth and corn earworm on canola.     

Plant characteristics associated with natural enemy abundance at Michigan native plants

Habitat management is a type of conservation biological control that focuses on increasing natural enemy populations by providing them with plant resources such as pollen and nectar. Insects are known to respond to a variety of plant characteristics in their search for plant-provided resources. A better understanding of the specific characteristics used by natural enemy insects in selecting these resources could greatly improve efficiency in screening plants for habitat management. We examined 5 previously tested and widely recommended resource plants and 43 candidate plants to test whether the number and type of natural enemies and herbivores at each plant were predicted by plant characteristics including: period of peak bloom, floral area, maximum flower height, hue, chroma, and corolla size. Natural enemy abundance increased with week of peak bloom and greater floral area across all plants tested. Ordination of plant characteristics indicated that increasing floral area, period of peak bloom, maximum flower height, and decreasing corolla width grouped together into a single principal component. Both natural enemy and herbivore abundance increased significantly with the principal component for this set of characteristics, but the relationship with herbivore abundance was weaker. These results indicate that, for a given time of the season, selection of plants with the largest floral area has potential to increase natural enemy abundance in habitat management plantings and streamline plant selection for habitat management.     

农田非作物生境调控与害虫综合治理

就害虫综合治理的研究而言,农田生态系统可以区分出作物生境和邻近作物的非作物生境。昆虫从作物生境迁移到非作物生境,与作物生境缺乏食物或受人类栽培活动干扰而引起的自然迁移、转换寄主和寻求庇护场所有关。许多研究表明,与特定作物田块相联系的植被类型和结构可影响害虫及其天敌迁居的种类、数量和时间。因此,我们可以通过改变大田周围非作物生境的植被组成及特征来调控农业生态系统中害虫与天敌的相互关系,提高天敌对害虫的控制效能。由于景观的空间格局对节肢动物的生物学特性有直接或间接的影响,所以,以景观为单元要比以同类作物的田块为单元更加适于害虫综合治理的研究和实施。在现代农业景观区域内重新引入和相嵌一些非栽培植物或廊道,可为众多有益节肢动物的繁殖、取食和避害提供多种类型的非作物栖境。     

Assessment of the natural control of Neoleucinodes elegantalis in tomato cultivation using ecological life tables

Life table studies allow for a better understanding of mortality factors against insect pests. It assesses each mortality factor and its parameters in population suppression. Neoleucinodes elegantalis (Guenée) (Lepidoptera: Crambidae) is one of the major pests of solanaceous crops in South America. The background information provided by life table studies might be used for developing management strategies for controlling N. elegantalis. Thus, the aim of this study was to identify and quantify the mortality factors of N. elegantalis in tomato cultivation by using life tables. The results were obtained from 64 ecological life tables. The mortality factors for eggs, larvae and pupae were due to rain, fruit drop, predation, parasitism and unviability caused by physiological disorder and diseases. The percentage of total death for eggs, larvae and pupae were 64.2%, 24.9% and 5.93%, respectively. The highest rates of mortality were caused by predation and parasitism. Thus, mortality of N. elegantalis caused by natural enemies might be used as a basis for determining which methods are better for controlling N. elegantalis while having the lowest possible impact on its natural enemies.     

Diversity and abundance of lepidopteran stem borers and their host plants in Ethiopia

Lepidopteran stem borers are among the most important insect pests of maize, sorghum and sugarcane in sub‐Saharan Africa. Except for Chilo partellus, the other stem borer pests in Ethiopia are indigenous to Africa and are assumed to have coevolved with some native grasses and sedges in the natural habitat. In addition to pest species, natural habitats harbour diverse non‐economic stem borer species, some of which are new to science. However, with the growing threats to natural habitats, some non‐economic stem borer species may switch or expand their host ranges to include cultivated crops and evolve as “new” pests. Besides host switch, some of the unknown species currently limited to natural habitats may disappear. We examined the diversity, abundance and interactions of lepidopteran stem borers and their wild host plants in five different vegetation mosaics in Ethiopia. The stem borer species diversity varied among vegetation mosaics and host plants. Forty‐four stem borer species belonging to 14 different genera in the families of Noctuidae, Crambidae, Pyralidae and Tortricidae were recorded from 34 wild host plants and through light trap. Among these families, Noctuidae was the highest in species richness in which 31 species were identified, out of which 15 species and two genera were new to science. This paper discusses the ecological interpretation of host plant–stem borer species interactions, particularly in relation to habitat disturbances.     

Life history, life table, habitat, and conservation of Byasa impediens (Lepidoptera: Papilionidae)

This article investigates the biology of Byasa impediens, presenting its life-table data and analyzing its habitat requirements and the key factors threatening the survival of this species. This study also aims to detect specific protection methods to guarantee the long-term survival of Byasa impediens in Baishuijiang Reserve. Byasa impediens is bivoltine in Baishuijiang Reserve. The pupae overwinter on shrubs or on branches of trees. The eclosion of the first generation starts in mid-April. The adults of the first generation emerge in large numbers in mid-late May, and the second generation emerges from late June to mid-July. The two generations overlap. The adult males emerge 7–10 days earlier than the adult females. Their flight behavior is determined by factors such as perching along small rivers and gullies, and visiting flowers. The males are strong fliers. On the contrary, the flying ability of the females is weak; therefore, they just visit flowers, mate, and lay eggs near the natal area. The ratio of female to male is 1:4.1. The maximum lifespan of the males is 26 days, with an average of 6.9 days, whereas the maximum lifespan of the females is 21 days, with an average of 7.6 days. The pregnant eggs per female are 31.5 on average. The incubation period of the first generation is 12–14 days, whereas that of the second generation is 7–9 days. The larvae feed mainly on Aristolochia heterophylla. The larval period of the first generation lasts for 30 days with five instars and that of the second generation lasts for 30–40 days with five or six instars. The pupal period of the first generation begins in early June and lasts for 20–26 days whereas for the second generation, it begins between late July and late September. The adults prefer the following nectar plants: Albizzia julibrissin, Bauhinia glauca, Clerodendrum bungei, and Sambucus chinensis. The plant Aristolochia heterophylla is distributed at an altitude of 900–1680 m, and the most suitable range is 1200–1500 m. The host plants grow mainly along the paths and along the borders of forests where the canopy is rather open and shrubby undergrowth is found. The plant can hardly be found when the canopy density of the forest is greater than 80%. The elevation range most suitable for the larvae is 1200–1500 m. The ideal habitat of host plants also seems to be the ideal habitat of Byasa impediens. The key factors that adversely affect the population of Byasa impediens are loss and deterioration of habitats. The loss and deterioration of the habitats result in a decrease in the numbers of host plants and a more restricted distribution of the potential habitats. The habitat is easily influenced by anthropogenic activities, such as herding, cultivating, and using pesticides, which in turn influences the growth of the host plant Aristolochia heterophylla and the larvae of butterflies. Abnormal climatic conditions and natural enemies are the key factors affecting population density. The hot and dry weather in summer and the heavy rain in autumn considerably reduce the survival rate of eggs and larvae. The ichneumon parasitoids reduce the survival rate of the over-wintering pupae. The primary natural enemies of the larvae include spiders, earwigs, wasps, bugs, and ichneumon parasitoids. Other natural enemies of pupae and adults are birds. The most important conservation measures are preservation and reconstruction of the natural habitat, which includes rebuilding forests, enhancing management, enforcing existing laws, and developing eco-tourism. Creating a core-patch near all patches at a central area is also an important measure for conservation. In the most ideal habitat, appropriate shrub cutting can increase the growth of host plants, thereby promoting expansion of the Byasa impediens population.     

Gall-forming and free-feeding herbivory along vertical gradients in a lowland tropical rainforest: the importance of leaf sclerophylly

In contrast to most insect guilds, gall-forming insects are thought to reach highest diversity on sclerophyllous vegetation, such as Neotropical savannas and Mediterranean vegetation types. The water and nutrient stress endured by meristems of canopy trees in tall wet tropical rainforests may cause leaf sclerophylly. Hence, the upper canopies of such ecosystems may represent a suitable habitat for gall-forming insects. At the San Lorenzo Protected Area, Panama, we estimated free-feeding herbivory and gall densities within five sites in 2003 and 2004, by surveying leaves in vertical and horizontal transects. In each sample, we recorded leaf density (mature and young foliage), free-feeding herbivore damage and number of galls, including the presence of live larvae, parasitoids or fungi. We surveyed 43 994 leaves, including 231 plants and 73 tree and liana species. We collected 5014 galls from 17 host-plant species, including 32 gall species of which 59% were restricted to the canopy (overall infestation rates: 2.4% in 2003, 5.5% in 2004). In 2003, 16% of the galls were occupied by live larvae, against 5% in 2004. About 17–20% of leaves surveyed suffered from free-feeding herbivory. Leaf sclerophylly increased significantly with sampling height, while free-feeding herbivory decreased inversely. Conversely, the number of live galls collected in the canopy was 13–16 times higher than in the understorey, a pattern consistent across sites and years. Hence, the probability of gall survivorship increased with increasing leaf sclerophylly as death by fungi, parasitoids or accidental chewing were greater in the understorey. Increasing harsh ecophysiological conditions towards the upper canopy appear favourable to galls-forming population maintenance, in support of the hypothesis of harsh environment. Hence, gall diversity and abundance in the upper canopy of tall tropical forests are perhaps among the highest in the world.     

Type 3 functional response of mice to gypsy moth pupae: is it stabilizing?

We conducted field experiments in 2002 and 2003 to determine whether the functional response of white-footed mice ( Peromyscus leucopus ) to gypsy moth ( Lymantria dispar ) pupae is decelerating (e.g. type 2) or accelerating (e.g. type 3) at low pupal density. In both experiments, live gypsy moth pupae were deployed in June (prior to the appearance of natural pupae) at densities of approximately 1, 8, and 35 pupae per mouse home range in oak-forest grids in upstate New York and monitored over 10 days for signs of predation. Pupae were deployed 1.5-m high on tree boles in 2002, whereas in 2003 the three density treatments were crossed with a height treatment: ground level vs 1.5-m high. The relationship between daily predation rate (proportion of pupae eaten/day) and pupal density was significantly positive in both years, indicating an accelerating functional response. Daily predation rates on ground-level pupae were substantial in the lowest density treatment, suggesting that dense mouse populations could drive gypsy moths to extinction despite an accelerating functional response. Daily predation rates on elevated pupae increased over several days in the medium and high density treatments, suggesting a lagged shift from ground- to tree-level foraging by mice. Within the high-density treatments, predation rates on pupae showed no apparent relationship with the number of pupae on a tree. Our results disagree qualitatively with simple models of type 3 functional response, in which predation rate of prey approaches zero as prey approach extinction, and support the contention that an accelerating functional response alone may be insufficient to prevent prey extinction.