Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Reprint of: Tree diversity promotes predatory wasps and parasitoids but not pollinator bees in a subtropical experimental forest

From regional to global scales, anthropogenic environmental change is causing biodiversity loss and reducing ecosystem functionality. Previous studies have investigated the relationship between plant diversity and functional insect communities in temperate and also in tropical grasslands and forests. However, few studies have explored these dynamics in subtropical forests. Here, cavity-nesting Hymenoptera and associated parasitoids were collected across a controlled tree diversity experiment in subtropical China to test how predatory wasps, bees and parasitoids respond to tree species richness. Abundance and species richness of predatory wasps and parasitoids were positively correlated with tree species richness, while bee abundance and bee species richness were unrelated to tree species richness. Our results indicate that tree species richness increases the abundance and species richness of important communities such as predators and parasitoids. Moreover, the results highlight the importance of subtropical forests in maintaining abundance and species richness of key functional insect groups.     

Vertical stratification of ichneumonid wasp communities: the effects of forest structure and life‐history traits

Parasitoid wasp communities of the canopy of temperate forests are still largely unexplored. Very little is known about the community composition of parasitoids between canopy and understory and how much of this difference is related to forest structure or parasitoid biological strategies. In this study we investigated upon the difference in the community composition of the parasitic wasps Ichneumonidae between canopy and understory in a lowland temperate forest in northern Italy. We used general linear models to test whether parasitic strategy modifies species vertical stratification and the effect of forest structure. We also tested differences in β‐diversity between canopy and understory traps and over time within single forest layers. We found that stand basal area was positively related to species richness, suggesting that the presence of mature trees can influence local wasp diversity, providing a higher number of microhabitats and hosts. The ichneumonid community of the canopy was different from that of the understory, and the β‐diversity analysis showed higher values for the canopy, due to a higher degree of species turnover between traps. In our analyses, the vertical stratification was different between groups of ichneumonids sharing different parasitic strategies. Idiobiont parasitoids of weakly or deeply concealed hosts were more diverse in the understory than in the canopy while parasitoids of spiders were equally distributed between the two layers. Even though the ichneumonid community was not particularly species‐rich in the canopy of the temperate forests, the extension of sampling to that habitat significantly increased the number of species recorded.     

Insect responses to host plant provision beyond natural boundaries: latitudinal and altitudinal variation in a Chinese fig wasp community

Many plants are grown outside their natural ranges. Plantings adjacent to native ranges provide an opportunity to monitor community assembly among associated insects and their parasitoids in novel environments, to determine whether gradients in species richness emerge and to examine their consequences for host plant reproductive success. We recorded the fig wasps (Chalcidoidea) associated with a single plant resource (ovules of Ficus microcarpa) along a 1200 km transect in southwest China that extended for 1000 km beyond the tree's natural northern range margin. The fig wasps included the tree's agaonid pollinator and other species that feed on the ovules or are their parasitoids. Phytophagous fig wasps (12 species) were more numerous than parasitoids (nine species). The proportion of figs occupied by fig wasps declined with increasing latitude, as did the proportion of utilized ovules in occupied figs. Species richness, diversity, and abundance of fig wasps also significantly changed along both latitudinal and altitudinal gradients. Parasitoids declined more steeply with latitude than phytophages. Seed production declined beyond the natural northern range margin, and at high elevation, because pollinator fig wasps became rare or absent. This suggests that pollinator climatic tolerances helped limit the tree's natural distribution, although competition with another species may have excluded pollinators at the highest altitude site. Isolation by distance may prevent colonization of northern sites by some fig wasps and act in combination with direct and host‐mediated climatic effects to generate gradients in community composition, with parasitoids inherently more sensitive because of declines in the abundance of potential hosts.     

Restore it,and they will come: trap-nesting bee and wasp communities (Hymenoptera: Aculeata) are recovered by restoration of riparian forests

Riparian forests have been greatly affected by anthropogenic actions with formerly continuous riparian forests being slowly converted into small and isolated patches. Riparian forests are extremely important habitats for many groups of insects, including bees and wasps, because they are sources of shelter and food for them and their offspring. There is a growing body of evidence of success in the restoration of riparian forest plant communities; however, little research has been done on the associated invertebrate communities. We test whether restoring plant communities is sufficient for restoring the taxonomic composition of trap-nesting bees and wasps and which functional traits are favored in different sites. We predict that species richness, abundance, and community composition of trap-nesting bees and wasps of riparian sites undergoing restoration will converge on the “target” of a reference site with increasing time, since restoration increases habitat complexity. We also predict that the width of restored patches will also influence the species richness, abundance and community composition of trap-nesting bees and wasps. Bee richness and abundance, and wasp richness, were strongly related to fragment width, but not to age since restoration. Our results indicate that although restored sites are relatively small and scattered in a fragmented landscape, they provide suitable habitat for re-colonization by community assemblages of trap-nesting bees and wasps and the traits selected captured the responses to the habitat restoration. Hence, restored riparian areas can be considered important habitats for invertebrates, thus contributing to an increase in local biodiversity and, possibly, the restoration of some of the ecosystem services they originally provided.     

Recovery of species diversity and composition of braconid parasitic wasps after reforestation of degraded grasslands in lowland East Kalimantan

Human-induced disturbances and wildfires can transform areas of tropical rainforest into Imperata-dominated grassland, but it may be possible that recovery of biodiversity is facilitated by reforestation with fast-growing trees. We compared the assemblages of braconid wasps as parasitoids of taxonomically diverse groups of insects among Imperata grasslands, young and mature plantations of Acacia mangium, young secondary forests after wildfires, and old secondary forests in the lowland of East Kalimantan. The abundance and species richness of braconids, which had declined in Imperata grasslands, somewhat increased in Acacia plantations, and also the species composition of braconids in Acacia plantations was transitional between Imperata grasslands and old secondary forests. Parasitoids of detritivores and wood borers increased markedly after plantation, while those of herbivores showed a distinct turnover of species all over the range from grasslands to old secondary forests. The plantation of A. mangium had most likely facilitated the recovery of the diversity of host forest and their parasitoids, but the recovery was just at the rudimentary stage even in mature plantations. Monitoring of parasitic wasps would be useful to test the continuous recovery of forest biodiversity in plantation stands.     

Insect attraction to herbivore-induced beech volatiles under different forest management regimes

Insect herbivore enemies such as parasitoids and predators are important in controlling herbivore pests. From agricultural systems we know that land-use intensification can negatively impact biological control as an important ecosystem service. The aim of our study was to investigate the importance of management regime for natural enemy pressure and biological control possibilities in forests dominated by European beech. We hypothesize that the volatile blend released from herbivore-infested beech trees functions as a signal, attracting parasitoids and herbivore enemies. Furthermore, we hypothesize that forest management regime influences the composition of species attracted by these herbivore-induced beech volatiles. We installed flight-interception traps next to Lymantria dispar caterpillar-infested young beech trees releasing herbivore-induced volatiles and next to non-infested control trees. Significantly more parasitoids were captured next to caterpillar-infested trees compared to non-infested controls, irrespective of forest type. However, the composition of the trophic guilds in the traps did vary in response to forest management regime. While the proportion of chewing insects was highest in non-managed forests, the proportion of sucking insects peaked in forests with low management and of parasitoids in young, highly managed, forest stands. Neither the number of naturally occurring beech saplings nor herbivory levels in the proximity of our experiment affected the abundance and diversity of parasitoids caught. Our data show that herbivore-induced beech volatiles attract herbivore enemies under field conditions. They further suggest that differences in the structural complexity of forests as a consequence of management regime only play a minor role in parasitoid activity and thus in indirect tree defense.     

Differences in endemic insect assemblages among vegetation types on a small island of the oceanic Ogasawara Islands

Natural vegetation is often replaced by invasive alien plants on isolated oceanic islands. To determine how invasive alien plants affect insect diversity, we compared flying insects captured using Malaise traps among different vegetation types on a small island (Nishijima; 0.49 km²) in the oceanic Ogasawara (Bonin) Islands in the north‐western Pacific. The numbers of individuals and species, and the species composition of pollinators (bees), predators (wasps) and wood borers (cerambycid, mordellid and elaterid beetles) were compared among three vegetation types: Casuarina equisetifolia (an invasive alien tree) forest, natural forest and natural grassland (forest edge), during two seasons (June and October–November 2005). In traps, 80.0, 66.7, 87.5, 85.7 and 100.0% of bee, wasp, cerambycid, mordellid and elaterid beetle species, respectively, were endemic to the Ogasawara Islands. Grassland had the highest wasp and bee species richness, whereas natural forest had the highest species richness of wood‐boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups (except mordellid beetles). More individuals of most insect groups (except bees) were captured in June than in October–November. More individual bees and wasps were captured in grassland than in forests, whereas more individual mordellid and elaterid beetles were captured in forests than in grassland. The number of cerambycid individuals did not differ among vegetation types. Redundancy analysis suggested that most insect species preferred natural forest or grassland to alien forest. Therefore, further invasion of natural grassland and forest by the alien tree C. equisetifolia may negatively affect the endemic insect fauna of Nishijima.     

Classification and ordination of communities of soil arthropods in an urban park of Osaka City

To assess the effect of management of a park on soil arthropods, communities of Oribatida and Collembola were analyzed at 11 sites of different vegetation in Tsurumi Park, an urban park of Osaka City. The type of canopy layer and soil density strongly affected the community parameters, such as species diversity. Ordination revealed that soil density, contents of organic matter, and shrub layer were important for variety in the oribatid community; the shrub layer was important for the collembolan community. Species richness of both arthropod groups was highest in a mixed forest and lowest on bare land, while the abundance of Collembola was highest on a lawn site. Areas of a common vegetation type had a similar oribatid community;Trichogalumma nipponica dominated in deciduous forests,Eohypochthonius crassisetiger in mixed forests and one of coniferous forests. On the other hand, collembolan communities did not correspond with the vegetation.Sminthurinus sp. was collected from every site, and the most abundant species wasCryptopygus thermophilus that exhibited an outbreak on lawn sites. A significant correlation existed between species diversities but not between abundances of Oribatida and Collembola.     

Does forest loss affect the communities of trap-nesting wasps (Hymenoptera: Aculeata) in forests? Landscape vs. local habitat conditions

We investigated changes in the communities of trap-nesting Hymenoptera in forests in relation to forest loss on a landscape scale and understory conditions on a local habitat scale. Two specific questions were addressed. (1) Do the communities change with degrees of forest loss? (2) Do the communities change with varying local environmental conditions of understory habitats? The study was made in a landscape characterized by distributed forest patches within intensively managed agricultural surroundings. We deployed trap-nests at eight randomly selected sites in forests in summer. To quantify forest loss, the amount of forest coverage was calculated using GIS. To indicate local habitat conditions, the species richness of understory flowering plants was used. All together, 12 species of wasps and no bees were captured. Regression analyses showed that both abundance and species richness of the wasps were not significantly related to forest coverage. However, abundance of trap-nesting wasps was significantly related to species richness of understory plants, but species richness of the wasps was not significantly related to the plants. These results suggest that communities of trap-nesting wasps in forests are influenced more by the local habitat conditions than by forest loss.     

Richness and Abundance of Ichneumonidae in a Fragmented Tropical Rain Forest

Because of the magnitude of land use currently occurring in tropical regions, the local loss of animal species due to habitat fragmentation has been widely studied, particularly in the case of vertebrates. Many invertebrate groups and the ichneumonid wasps in particular, however, have been poorly studied in this context, despite the fact that they are one of the most species-rich groups and play an important role as regulators of other insect populations. Here, we recorded the taxonomic composition of ichneumonid parasitoids and assessed their species richness, abundance, similarity, and dominance in the Los Tuxtlas tropical rain forest, Mexico. We compared two forest types: a continuous forest (640 ha) and a forest fragment (19 ha). We sampled ichneumonids using four malaise traps in both forest types during the dry (September–October) and rainy (March–April) seasons. A total of 104 individuals of Ichneumonidae belonging to 11 subfamilies, 18 genera, and 42 species were collected in the continuous forest and 11 subfamilies, 15 genera, and 24 species were collected in the forest fragment. Species richness, abundance, and diversity of ichneumonids were greater in the continuous forest than in the forest fragment. We did not detect differences between seasons. Species rank/abundance curves showed that the ichneumonid community between the forest types was different. Species similarity between forest types was low. The most dominant species in continuous forest was Neotheronia sp., whereas in the forest fragment, it was Orthocentrus sp. Changes in the ichneumonid wasp community may compromise important tropical ecosystem processes.     

Cocoon predation on diprionid sawflies: the effect of forest fertility

Predation by small mammals is thought to be one of the main regulators of outbreaking sawfly species. It has been suggested that predation may be lower in poor and dryish forests, and this is the reason why outbreaks often begin from this type of environment. We studied experimentally how fertility of the forest site affects cocoon predation experienced by two sawfly species, the common pine sawfly Diprion pini (Linnaeus) and the European pine sawfly Neodiprion sertifer (Geoffroy). We applied a fertilization treatment to selected pine-dominated barren forest sites in Finland, and 2–4 years later monitored predation on the sawfly cocoons in fertilized and control areas. The results did not support the idea that forest fertility was related to cocoon predation. We also could not verify that small mammal abundance was related to fertility of the forest. The most obvious pattern we observed was that the two sawfly species differed dramatically in predation experienced. N. sertifer has its cocoon phase in mid-summer and experienced only moderate predation (37%) whereas D. pini, with its cocoon phase in autumn, suffered from very heavy predation (96%). Our observations suggest that if predation is important in controlling the population dynamics of the species, its impact depends more on the sawfly species and season than on the fertility of the forest site. Received: 1 March 1998 / Accepted: 25 May 1998     

Saproxylic insects and fungi in deciduous forests along a rural–urban gradient

Urbanization is increasing worldwide and is regarded a major threat to biodiversity in forests. As consequences of intensive human use, the vegetation structure of naturally growing urban forests and their amount of deadwood can be reduced. Deadwood is an essential resource for various saproxylic insects and fungi. We assessed the effects of urbanization and forest characteristics on saproxylic insects and fungi. We exposed standardized bundles consisting of each three freshly cut beech and oak branches in 25 forests along a rural–urban gradient in Basel (Switzerland). After an exposure of 8 months, we extracted the saproxylic insects for 10 months using an emergence trap for each bundle. We used drilling chips from each branch to determine fungal operational taxonomic units (OTUs). In all, 193,534 insect individuals emerged from the experimental bundles. Our study showed that the abundance of total saproxylic insects, bark beetles, longhorn beetles, total flies, moths, and ichneumonid wasps decreased with increasing degree of urbanization, but not their species richness. However, the taxonomic composition of all insect groups combined was altered by wood moisture of branches and that of saproxylic beetles was influenced by the degree of urbanization. Unexpectedly, forest size and local forest characteristics had a minor effect on saproxylic insects. ITS (internal transcribed spacer of rDNA) analysis with fungal specific primers revealed a total of 97 fungal OTUs on the bundles. The number of total fungal OTUs decreased with increasing degree of urbanization and was affected by the volume of naturally occurring fine woody debris. The composition of fungal OTUs was altered by the degree of urbanization and pH of the branch wood. As a consequence of the altered compositions of saproxylics, the association between total saproxylic insects and fungi changed along the rural–urban gradient. Our study shows that urbanization can negatively impact saproxylic insects and fungi.     

Community structure, survival and mortality factors in arctic populations of Eupontania leaf gallers

We studied survival, mortality factors, and community structure of nine species of leaf-galling sawflies, Eupontania spp., living on ten willow species (Salix spp.) at six sites on the Russian arctic tundra. The sawfly species represented two different gall types: the viminalis-type, which forms pea-shaped galls on the underside of leaf blades, and the vesicator-type, which forms bean-shaped galls on both sides of the leaf blade. Gall communities in the northernmost site had only one parasitoid species, but up to six parasitoids were found at the southernmost site. Inquiline parasitoids were encountered only in the two southern sites. Survival of the larvae varied between 20.0 and 82.8% among galler species at different sites. Parasitoids were the most important mortality factor for the sawflies. They caused mortality of 7.8-65.4%, depending on galler species and site, and it was highest in the northernmost site. Plant-specific mortality varied from 1.7 to 28.4% by galler species and it tended to decrease towards the north. Mortality from parasitoids was greater in the vesicator-type gallers than in the viminalis-type gallers. The total mortality caused by parasitoids in the arctic communities does not appear to differ from that in the diverse southern communities of Eupontania in Middle Europe, Scandinavia and North America, despite the assemblage having only a few members in the Arctic. The largest difference between the southern and the northern communities was the lack of inquiline parasitoids in the north. Our data do not support the hypothesis that abiotic, rather than biotic, factors would be more important in determining the abundance of populations of herbivorous insects in the harsh arctic environment.     

Restructuring of Lepidoptera communities by introduced Vespula wasps in a New Zealand beech forest

Introduced social wasps (Vespula vulgaris) reach high densities in some New Zealand beech forests, because honeydew provides an abundant high-energy food source. We manipulated wasp density to estimate an “ecological damage threshold” for large, free-living Lepidoptera larvae. There will be a continuum of ecological damage thresholds for wasp density depending on the prey species or habitat. Experimentally placed small caterpillars had a significantly higher survival rate than large caterpillars, and the survival rate of both groups decreased with increasing wasp density. Spring-occurring caterpillars have a probability of surviving of 0.90–0.95, assuming wasps are the only source of mortality. However, at the peak of the wasp season we predict caterpillars would have virtually no chance (probability of 10⁻⁷⁸ to 10⁻⁴⁰) of surviving to adults. Wasp abundance must be reduced by at least 88% to conserve the more vulnerable species of free-living caterpillars at wasp densities similar to those observed in our study sites. This equates to a damage threshold of 2.7 wasps per Malaise trap per day. It was exceeded for about 5 months of the year in non-poisoned sites. There are currently no biological or chemical control techniques available in New Zealand that will reduce wasp abundance below this damage threshold throughout the year. Our models show that most Lepidoptera with spring caterpillars will be able to persist, but species with caterpillars occurring in the peak wasp season will be eliminated. Received: 5 January 1998 / Accepted: 10 February 1999     

Responses of parasitoids to saproxylic hosts and habitat: a multi-scale study using experimental logs

Species belonging to higher trophic levels are particularly vulnerable to habitat loss and consequential host population declines, but detection of effects depends on observation scale. We investigated the effects of habitat and host availability at multiple scales on parasitoids of early successional saproxylic beetles in middle boreal Sweden, where forestry has led to habitat fragmentation and coarse woody debris (CWD) loss. Parasitoid wasps and beetle hosts were collected from nine locations, each containing three spruce-dominated stand types (clear-cut, mature managed and unmanaged stands), using emergence traps on experimental CWD. We measured local CWD volumes and determined the availability of forests of a suitable age within the landscape. We tested parasitoid responses to stand type, CWD volume, abundance of known and probable hosts and longitude. Additionally, we tested whether parasitoids responded to the area of habitat of a suitable age within radii from 0.2 to 10 km. Stand type appeared in best-fit models for all common species, suggesting that wasps respond strongly to habitat at local scales. Longitude (largely climate) featured commonly, but CWD volume was never significant. Host abundance appeared in best-fit models for three of five common species, proving significant only for Bracon obscurator, the abundance of which correlated with that of Orthotomicus laricis at both trap and site levels. Rhimphoctona spp. also correlated significantly with its known host Tetropium castaneum at the trap level. B. obscurator responded to habitat area at scales of 0.6–1 km and Cosmophorus regius responded at radii greater than 7 km, while the larger species did not respond strongly to habitat area. The role of habitat area at greater scales thus varied greatly amongst species, but our data suggest that dispersal of these common early successional species may not be strongly restricted at the current scale of fragmentation of their boreal habitats.     

Vegetation increases the abundance of natural enemies in vineyards

Non-crop areas can increase the abundance of natural invertebrate enemies on farmland and assist in invertebrate pest control, but the relative benefits of different types of vegetation are often unclear. Here, we investigated abundance of natural enemies in vineyards with edges consisting of different types of vegetation: remnant native forests, wooded margins planted after establishment of the crop (hereafter called shelterbelts), or pasture. Invertebrates were sampled four times using canopy sticky traps and ground level pitfall traps, replicated across two seasons at one of the sites. The distribution and abundance of natural enemies in relation to edges with adjacent vegetation or pasture were mapped by distance indices (SADIE) and compared with ANOVAs. There was a positive influence of adjacent wooded vegetation on staphylinids, predatory thrips, predatory mites, spiders, ladybird beetles and hymenopteran parasitoids including Trichogramma egg parasitoids in the canopy and/or at ground level, although there were significant differences among sites and groups of organisms. In contrast, pasture edges had no effect or a negative effect on numbers of natural enemies in vineyards. To directly assess potential beneficial effects of adjacent vegetation, predation and parasitism of eggs of a vineyard insect pest, Epiphyas postvittana Walker (Lepidoptera: Tortricidae), was measured. Parasitism by Trichogramma was higher adjacent to remnant vegetation while predation was not affected. These results indicate that the abundance and distribution of vineyard natural enemies and parasitism of pest moth eggs is increased adjacent to edges with wooded vegetation, leading to beneficial effects for pest control. The conservation of remnant woodland and planting of shelterbelts around vineyards may therefore have direct economic benefits in terms of pest control, whereas non-crop pasture may not produce such benefits.     

Volatiles from green‐lipped mussel as a lead to vespid wasp attractants

Vespid wasps (Vespula vulgaris L. and V. germanica Fab. Hymenoptera; Vespidae) are highly abundant in 1 million ha of New Zealand's indigenous beech forests (Nothofagus spp.) and have had detrimental effects on the New Zealand native fauna. This hyperabundance is due in part to the vast supply of carbohydrate‐rich honeydew produced by scale insects Ultracoelostoma spp. native to New Zealand. Current control methods include the use of wet cat food as a protein source with insecticide as a lure‐and‐kill‐based system, but there are problems with fresh baits degrading rapidly, and a more durable formulation would enable the expansion and longevity of wasp control. Four crude protein baits were tested for vespid attraction. Green‐lipped mussels had the highest vespid catch of the crude baits tested, and aged and fresh mussels were equally attractive. From headspace analysis of the green‐lipped mussel volatiles, a series of butanoate esters, 3‐octanone and 1‐octen‐3‐ol were identified as possible attractants. These compounds were tested individually and in various blend combinations for the attraction of Vespula wasps in matagouri vegetation at the edge of beech forests. We found synergistic effects between single attractive compounds when tested in various combinations, and the multicomponent lures were more attractive to these wasps than heptyl and octyl butanoate, previously identified attractants for vespid species. The new multicomponent lures could form the basis for a new generation of attractants for social wasps that can provide sustained control methods for invasive vespid wasps.     

The role of host plant species in the phenotypic differentiation of sympatric populations of Aleiodes nolophanae and Cotesia marginiventris

The ecology of parasitoids is strongly influenced by their host plant species. Parasitoid fitness can be affected by a variety of plant traits that could promote phenotypic differentiation among populations of parasitoids. Generalist parasitoids are expected to be more affected by plant traits (e.g., plant defensive traits) than specialist parasitoids. Data are presented on phenotypic differences of two braconid parasitoid wasps ovipositing on the same insect host species on two different host plant species. Adult mass, adult longevity, and percent parasitism are compared for the generalist parasitoid Cotesia marginiventris Cresson and the specialist parasitoid Aleiodes nolophanae Ashmead (both Hymenoptera: Braconidae) emerging from green cloverworms, Hypena scabra Fabricius (Lepidoptera: Noctuidae), feeding on two host plant species, alfalfa (Medicago sativa L.) and soybean (Glycine max L. Merr.) (both Fabaceae), at three locations. Specialist wasps that parasitized the green cloverworm on alfalfa had a significantly larger mass than the ones that parasitized the green cloverworm on soybean at the three study sites. Generalist wasps that parasitized green cloverworms on alfalfa had a larger mass than wasps parasitizing green cloverworms on soybean only at one of the study sites (i.e., Prince George's County, MD, USA). Similarly, both specialist and generalist wasps lived longer when parasitizing green cloverworms on alfalfa than when parasitizing them on soybean at only one of the study sites (i.e., Prince George's County). In Prince George's County, percent parasitism on alfalfa by the specialist parasitoid was higher than on soybean for three consecutive years and percent parasitism by the generalist parasitoid was the same on alfalfa and soybean every year. Thus, phenotypic differences among populations associated with different host plant species vary geographically (i.e., parasitoid phenotype associated with different host plant species differ at some sites while it is the same at other sites). The implications of geographic variation for biological control are discussed.     

Impact of cocoon predation and parasitism on endemic populations of the common pine sawfly, Diprion pini (L.) (Hymenoptera, Diprionidae) in different forest types

1 Predation and parasitism on litter‐buried cocoons of the common pine sawfly Diprion pini (L.) were compared in different forest types with endemic sawfly populations by field exposure of laboratory‐reared cocoons during three consecutive years (1993–1995). 2 The impact of cocoon predation was dependent on season and forest type. The highest predation (up to 95%) was found during autumn in forest stands with a dense understory vegetation. 3 Cocoon parasitism varied between year, season and forest type. The highest parasitoid attack was observed in pure pine forests with more or less barren soils, but did not exceed 24% of exposed cocoons. 4 Cocoons were exposed in small patches. Predators tended to exploit all cocoons of a patch, whereas parasitoids only attacked a few cocoons of a patch. Predation was similar on cocoons placed in the litter and those buried more deeply in the soil, whereas parasitism of soil‐buried cocoons was rare. 5 These results indicate that predators can have a remarkable potential for limiting endemic sawfly densities, if habitat conditions in a forest maintain their population and support their foraging behaviour. A notable effect of parasitoids on sawfly cocoons deposited in the litter is obviously restricted to typical pure and barren pine forests, but may play there a similar role as predation.