Wood ants and a geometrid defoliator of birch: predation outweighs beneficial effects through the host plant

Soil amelioration by a wood-ant species and its consequences for the larval performance of autumnal moths feeding on mountain birch were studied at various distances from the nest mound. Soil nitrate and ammonium nitrogen did not show any clear relationship with distance. However, trees growing in the mound had over 20% more foliar nitrogen than more distant trees. When moth larvae were experimentally protected from predation, their survival rate and final weight tended to decrease with increasing distance. In a laboratory experiment with detached leaves, the relative growth rate of larvae was roughly 30% higher on leaves from trees located on the mound. Differences in larval performance refute the Plant Stress Hypothesis proposed by T.C.R. White and support P.W. Price's Plant Vigor Hypothesis. Predation by ants was examined along the same gradient in trees with and without a glue band that excluded ants from the canopy. Reduction in the daily survival rate of larvae attributable to ant predation was about 35% in trees growing in the mound and around 5% at a distance of 20 m. Other things being equal, about 25 times more larvae entering the penultimate instar would achieve the pupal stage outside the wood-ant territory than in the vicinity of the mound. While both the fertilizing and predatory influence of wood ants is clear, the domain of predation is much larger than the area where trees and their herbivores can exploit enhanced nutrient levels in and around ant mounds. The existence of undamaged green islands around ant mounds in otherwise totally defoliated mountain-birch forests cannot be explained by soil amelioration by wood ants but rather by their predatory activity. Received: 21 November 1996 / Accepted: 8 September 1997     

Effects of ant exclusion during outbreaks of a defoliator and a sap-sucker on birch

1. The influence of a wood ant, Formica aquilonia, on the defoliation of the white birch, Betula pubescens, and on its invertebrate community was studied in ant-exclusion experiments during two outbreaks, the decline phase of the autumnal moth, Epirrita autumnata, and the peak year of the birch aphid, Euceraphis punctipennis, not tended by ants. 2. The numbers of the wood ant and Symydobius oblongus, a tended aphid, in birch foliage showed a strong positive correlation, and the former decreased rapidly when the distance from the ant mound increased, confirming that there was a distance-related gradient in arboreal ant predation. There may have been a parallel reduction in soil amelioration by ants through nest construction and food gathering. 3. The application of a glue ring around the trunk excluded ants totally from the canopy, inhibiting predation but not fertilization. 4. Ant-exclusion resulted in a 90–95% reduction in the growth of tended aphid colonies by mid-season. 5. Outbreak populations of the autumnal moth and the birch aphid were reduced by 45–67% and 77%, respectively, in control trees and correlated negatively with ant numbers. 6. The total percentage of leaf area damaged by moth larvae was 34% lower in ant-foraged than in unforaged trees. 7. Neither the distance from the ant mound nor its interaction with the glue treatment had any effect on the herbivores or folivory, indicating that the possible soil-ameliorating effect was weaker than predation, which reduced herbivore numbers at every distance studied (4–20 m). 8. Predation by ants also affected the abundance of syrphid larvae, predatory on both aphids, the percentage parasitism by a wasp, Aleiodes testaceus, on the autumnal moth, and thereby (or directly) its age (size) distribution. The presence of ants had no influence on spider abundance. 9. It is concluded that predation rather than soil amelioration is likely to be the reason why the degree of folivory and, during serious outbreaks, the mortality of trees are lower in the vicinity of wood-ant mounds.     

Testing the enemies hypothesis in forest stands: the important role of tree species composition

Numerous studies conducted in agro-ecosystems support the enemies hypothesis, which states that predators and parasites are more efficient in controlling pest densities in polycultures than in monocultures. Few similar studies, however, have been conducted in forest ecosystems, and we do not yet have evidence as to whether the enemies hypothesis holds true in forests. In a 2-year study, we investigated whether the survival of autumnal moth (Epirrita autumnata) larvae and pupae differs between silver birch monocultures and two-species mixtures of birch with black alder, Norway spruce and Scots pine. We placed young larvae on birch saplings and monitored their survival until the end of the larval period, when we checked whether they had been parasitized. After the larvae had pupated, pupal survival was tested in a field trial. In 2002, the larvae disappeared earlier and their overall survival was lower in birch–pine mixtures than in other stand types. In 2003, survival probability was lowest in birch–pine stands only during the first week and there were no differences between stands in overall survival. Larval parasitism was not affected by tree species composition. Pupal weight and pupal survival were likewise not affected by stand type. Among the predators, wood ants were more abundant on birches growing in birch–pine mixtures than in other stand types probably because colonies of myrmecophilic aphids were common on pines. In contrast, spider numbers did not differ between stand types. Ant exclusion by means of a glue ring around the birch trunk increased larval survival, indicating that ants are important predators of the autumnal moth larvae; differences in larval survival between stands are probably due to differential ant predation. Our results provide only partial support for the enemies hypothesis, and suggest that it is both tree species composition and species diversity which affect herbivore survival and predation.     

Neither the devil nor the deep blue sea: larval mortality factors fail to explain the abundance and distribution of Tischeria ekebladella

Abstract.  1. The distribution, abundance and population dynamics of herbivorous insects may be affected by trophic interactions, by abiotic influences, or by intra-specific processes. Relatively little is known about how trophic influences vary across space. Here, we investigate spatial variation in mortality in the oak-feeding leaf miner Tischeria ekebladella as attributable to individual causal agents.
2. Leaf miners were experimentally introduced on 67 trees on an island 5 km² in area in south-western Finland. On each tree, some larvae were protected by a muslin bag, others by a glue barrier around the branch and some left exposed.
3. In the bagged transplants, 78.4% of larvae survived, compared with only 9.6% in the other two treatments. Most of the mortality was because of airborne agents: mortality on branches sheltered by a glue barrier was as high as on fully exposed branch tips.
4. We consider mortality caused by parasitoid wasps to be the main source of larval death and the primary factor driving general patterns of survival. The effects of bird predation and premature leaf abscission were negligible.
5. We detected spatial aggregation in larval survival and parasitism rates at the level of individual trees, but not across the landscape.
6. Spatial variation in overall leaf miner survival, parasitism and leaf abscission does not suffice to explain patterns of incidence and abundance of wild T. ekebladella on experimental trees. Rather, we identify metapopulation dynamics as a likely determinant of the spatial distribution of T. ekebladella in the landscape.     

Limiting the success of stem borers (Prodoxus quinquepunctellus) in yuccas: indirect effects of ants, aphids, and fruit position

Abstract.  1. Although the moth–yucca mutualism is often studied as a pairwise interaction, yucca plants are also the sole host for a variety of other visitors. One of these additional visitors is a stem-boring moth, Prodoxus quinquepunctellus.
2. In this study, it is shown how the reproductive success of Prodoxus indirectly depends on the interactions between yuccas and their pollinators ( Tegeticula , Prodoxidae) as well as the indirect effects of ants and aphids.
3. Aggressive wood ants foraging on yuccas will attack adult Prodoxus moths while attempting to oviposit. This reduces the number of eggs laid in yucca stalks, leading to fewer larvae feeding in the stalks.
4. Once in the stalk, the survival of Prodoxus eggs/larvae depends upon the rate at which the flowering stalks dry out during fruit maturation. Portions of the stalk above the highest fruit dry out quickly and survivorship approaches zero in these dry sections, while larvae in green sections of the flowering stalk have significantly higher survival rates. The presence of aphids feeding on the stalk slows down the rate of stalk drying and could lead to increased survival of Prodoxus larvae.
5. Overall, ants have strong indirect effects on P. quinquepunctellus by controlling how many eggs are laid in the stalk and by influencing the distribution of aphids. However, it is primarily the presence and position of the fruit that can affect larval survivorship, and fruit position is a function of pollinator visits and resource limitation. These complex interactions illustrate the importance of studying the yucca–moth mutualism in a community context.     

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.     

Parasitism and ant protection alter the survival of the lycaenid Hemiargus isola

Abstract. 1. Although the majority of lycaenid–ant associations is facultative, few studies have documented the protection benefits provided by ants to lycaenids that are tended facultatively (Pierce & Easteal, 1986; Peterson, 1993).
2. Larvae of Hemiargus isola (Lepidoptera: Lycaenidae) feeding on Dalea albiflora are tended facultatively by several species of ant. In 1999 and 2000, the levels of parasitism and the identities of attendant ants were determined for larvae of H. isola . In addition, the presence of ants was manipulated experimentally to determine the efficacy of protection provided by attendant ants to H. isola .
3. Lycaenids were parasitised by a braconid wasp, Cotesia cyaniridis (Riley), and a tachinid fly, Aplomya theclarum (Scudder). In 1999 and 2000, 62 and 65% of larvae were parasitised; the percentage of the population parasitised did not differ significantly between years. In both 1999 and 2000, parasitism by the braconid wasp C. cyaniridis accounted for > 99% of all parasitism events.
4. Four species of ant, Crematogaster sp., Dorymyrmex sp., Forelius sp., and Formica sp., were associated with 88–99% of the tended lycaenids collected in both 1999 and 2000. For both years, there was a single, numerically dominant species associated with >80% of the tended larvae collected, but the identity of this numerically dominant ant differed between years.
5. Experimental exclusion of ants from D. albiflora plants resulted in 78% larval mortality as a result of parasitism, nearly twice that of larvae that were tended by ants on unmanipulated plants.     

The growth and survival of Panolis flammea larvae in the absence of predators on Scots pine and lodgepole pine

Abstract. 1. The survival, growth and development of larvae of the pine beauty moth, Panolis flammea (D & S) (Lepidoptera: Noctuidae) were examined as part of a study to determine (i) why damaging outbreaks of this insect in Scotland are frequent on lodgepole pine but do not occur on Scots pine, and (ii) why outbreaks are associated with areas of deep un-flushed peat.
2. Larvae were studied inside predator exclusion cages on mature pine trees in Elchies forest where a major infestation occurred in 1978–79.
3. Surprisingly, larval survival was generally greatest on Scots pine and lowest on lodgepole growing in deep peat. Also, larval growth and development were greater on Scots than lodgepole pine and were unaffected by the type of soil in which the lodgepole pine was growing.
4. Larval performance was generally better on lodgepole pine that had been thinned and received fertilizer.
5. Larval survival was affected by pine shoot stage during egg hatch in one of the two years in which this was examined.
6. Observations on P.flammea pupae were confused by disease, parasitism and probably by differences in weight loss caused by differences in larval development. Parasitism by ichneumonid parasitoids was recorded on Scots pine but not on lodgepole pine.
7. It was concluded that (i) the absence of pine beauty moth outbreaks on Scots pine, and (ii) the occurrence of outbreaks on lodgepole pine growing in deep unflushed peat were not the result of lodgepole pine growing in deep peat being a more suitable host plant than lodgepole pine and Scots pine growing elsewhere. It seems more likely that the observed outbreak behaviour of P.flammea in Scotland is associated with differences in predation and parasitism in plantations of difference host plants and in plantations of lodgepole pine growing in different soils.     

Folivorous larvae on flowers: do autumnal moths benefit from catkins of the mountain birch?

Immature leaves of birches and other deciduous trees and shrubs are the principal food resource for the larvae of the autumnal moth, Epirrita autumnata (Borkhausen) (Lepidoptera: Geometridae). However, the larvae of this geometrid, which displays pronounced population cycles and causes massive forest defoliations in northernmost Europe, are also frequently found on the catkins of mountain birch, Betula pubescens ssp. czerepanovii (Orlova) Hämet-Ahti (Betulaceae). To examine whether autumnal moth larvae visit birch catkins for their high nutritional value, we conducted two growth experiments in which the larval diet was supplemented with male catkins, and a single test where female catkins were used. We found that the presence of male catkins in the larval diet had a positive effect on the larvae, in relation to their pupal mass (reflecting life span fecundity), survival probability, as well as the duration of the larval period. The presence of female catkins in the diet did not affect larval performance. Our findings suggest that young autumnal moth larvae actively move to male catkins, which provide a nutritional benefit and thus lead to a better performance compared with those feeding elsewhere. Another possible benefit for young larvae choosing the catkins instead of leaves may involve predator or parasitoid avoidance.     

Effect of the canopy complexity of trees on the rate of predation of insects

The effect of the complexity of tree canopies on the effectiveness of biological control of tree‐dwelling insect herbivores by predators has been neglected. A complex canopy provides a greater variety of resources than a simple canopy, which may result in an increase in the abundance of both herbivores and predators in complex canopies and a higher predation rate. On the other hand, it may be more difficult to locate prey in trees with a complex canopy, which may lower the predation rate. The main aim of this study is to determine the relationship between predation rate and canopy complexity. We have used a system consisting of lime trees, larvae of blowflies as prey and naturally occurring predators (mainly birds and ants). The complexity of the canopy of 12 of the lime trees was reduced by pruning, and the predation rate was determined by recording the fate of prey pinned on each of the trees for 30 min several times during the course of a year. The predation rate was negatively associated with canopy complexity. We compare these results with those reported in the literature and contrary to the widely held view conclude that a negative relationship between predation rate and canopy complexity is a common phenomenon and can be expected to be reported in studies not only on ambush but also on predators that actively search for prey.     

Performance of a spring-feeding moth in relation to time of oviposition and bud-burst phenology of different host species

Abstract. 1. As a spring-feeding moth committed to immature foliage, the autumnal moth Epirrita autumnata (Lepidoptera, Geometridae) must have egg hatch synchronised with the bud-burst of its host plants. Due to large individual variation in the length of the pupal period, however, E. autumnata populations exhibit a prolonged period of flight and oviposition. Because the timing of oviposition in autumn is associated with the timing of egg hatch in the following spring, the time window for egg hatch expands and more potential hosts may become attainable. This suggestion was evaluated under field conditions by rearing E. autumnata eggs and larvae on four different hosts.
2. The performance of E. autumnata was measured by using estimates for fecundity (pupal mass) as well as survivorship of eggs and larvae. Based on the availability of foliage and phenological synchrony between larval and leaf development, early-laid eggs and the larvae originating from them were predicted to perform better on the hosts that have early-flushing leaves. On the late-flushing hosts, the larvae that hatched later were predicted to perform better than the larvae that hatched earlier. Half of the trials were exposed to predators and parasitoids, while the rest were conducted inside mesh-bags preventing larval dispersal and mortality due to natural enemies.
3. The results of the experiment did not support the simple predictions. In particular, host-plant quality and natural enemies appeared to operate discordantly between early- and late-laid eggs. Larvae from the late-laid eggs had rapid development during the larval stages and pupated at the same time and with the same pupal mass as the larvae hatched from the early-laid eggs.
4. The results indicate an occurrence of several, unknown selective forces in E. autumnata populations maintaining variation in the length of the pupal period, timing of oviposition, and timing of egg hatch.     

Effect of habitat complexity on biological control by the red imported fire ant (Hymenoptera: Formicidae) in collards

A potentially important and understudied biological control agent in US agroecosystems is the red imported fire ant, Solenopsis invicta Buren. Red imported fire ants may be particularly important biological control agents because we can manipulate their abundance with changes in habitat complexity. The effect of habitat complexity on biological control by fire ants was determined using plots of collards intercropped with white clover (complex habitat) and simple collard monocrops. The most economically significant pests of collards are larvae of the diamondback moth (DBM), Plutella xylostella (L.). Predation of DBM larvae by fire ants was more rapid and efficient in the intercrop than the monocrop. Red imported fire ants were 23% less abundant in the intercrop than the monocrop, however, suggesting that fire ants had a greater per capita effect on DBM survival in the complex habitat. Red imported fire ant predation of DBM larvae was significantly affected by larval density. Red imported fire ants also reduced the survival of leaf beetles, another economically significant pest taxa, by 45%. Furthermore, collard leaf damage tended to be inversely related to fire ant density and fire ants were more effective at reducing crop damage in the complex intercrop. Our study indicates the ability of red imported fire ants to be effective biological control agents and suggests that increasing habitat complexity can enhance red imported fire ant efficacy and herbivore control.     

Oviposition decreased in response to enriched water: a field study of the pitcher-plant mosquito, Wyeomyia smithii

Abstract.  1. Environmental cues are known to influence oviposition behaviour in mosquitoes, with important consequences for larval survival and insect population dynamics. Enriched microhabitats have been shown to be preferred oviposition sites.
2. In a field experiment designed to determine whether ovipositing mosquitoes are sensitive to different levels of nutrient enrichment, new pitcher-plant ( Sarracenia purpurea ) leaves were opened and enriched with 0, 2, or 20 dead ants, and the number of pitcher-plant mosquito ( Wyeomyia smithii ) larvae resulting from subsequent oviposition were measured.
3. Oviposition rates were higher in leaves with low levels of enrichment (0 and 2 ants per leaf), although larval development was enhanced at the highest enrichment level.
4. Results suggest that, although these mosquito larvae are nutrient limited, ovipositing females preferentially avoid highly enriched leaves. This counterintuitive result may be due to low oxygen concentrations or a masked cue in enriched leaves, and contrasts with other oviposition studies.     

Life-history consequences of predation for a subantarctic beetle: evaluating the contribution of direct and indirect effects

1. Recently, a small predatory beetle, Trechisibus antarcticus (Carabidae), was accidentally introduced onto the island of South Georgia, sub-Antarctic.
2. From the presumed site of introduction the beetle is invading the coastal lowland area, building up high densities locally in the tussock-forming grass Parodiochloa flabellata .
3. In the coastal area the endemic detritivorous/herbivorous beetle Hydromedion sparsutum (Perimylopidae) is common, especially in and beneath the tussocks.
4. The first three, out of six, larval instars of H. sparsutum are easily taken prey by the carabid.
5. In sites colonized by the carabid, total abundance and the ratio between larval and adult numbers of H. sparsutum are far lower, and its adult body size clearly larger, than in comparable sites where the carabid is absent.
6. Two hypotheses are proposed for explaining the increase in adult body size of H. sparsutum : (i) the increase is a direct effect of predation: selection by the predator favours large hatchlings and/or larvae with a high growth rate; and (ii) the increase is an indirect effect of predation: by lowering the density of H. sparsutum , predation has increased its per capita food supply, enabling a higher growth rate and a larger adult body size.
7. A food addition experiment in a carabid-free site showed availability of high quality food to be insufficient for sustaining the initial larval population.
8. In the laboratory, females from the predator-infested sites produced larger eggs and hatchlings than females from the carabid-free sites, but mass specific growth rates of the larvae were not higher.
9. Field and laboratory data give stronger support to the food hypothesis than to the size selectivity hypothesis.     

Influence of native ants on arthropod communities in a vineyard

• 1 Ants can have a range of effects on arthropods in crops, including suppressing herbivores such as caterpillars. However, ants can also increase hemipteran densities while reducing natural enemy numbers. In vineyard ecosystem, the effects of native ants and their interactions with other arthropods are poorly understood.
• 2 An ant‐exclusion experiment was designed to test the impact of native ants on both canopy and ground arthropods concurrently. The potential influence of ants on predation and parasitism of light brown apple moth (LBAM) eggs, a grape pest, was also examined. Adult grapevine scale insects and earwigs under bark were counted after a season of ant‐exclusion.
• 3 Among 23 ground ant species collected, six were found to forage in the canopy, with two Iridomyrmex species being the most commonly encountered.
• 4 There was no difference in the abundance of most arthropod orders and feeding groups between ant‐excluded and control vines, although ground spiders were more abundant under ant‐excluded vines, despite increased ground ant foraging pressure. LBAM egg parasitism and predation were low and probably affected by weather and other arthropods. Ant exclusion did not reduce survival of scale insects, although the distribution and abundance of scale insects were negatively associated with earwigs.
• 5 In conclusion, native ants did not consistently suppress arthropod assemblages, including natural enemies, and they did not promote the survival of scale insects. Interactions among native ant species within a vineyard might minimize their effects on other arthropods, although this needs further study.

     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Oviposition site preference and larval mortality in a leaf-mining moth

Abstract. 1. The univoltine leaf-mining moth, Lithocolletis quercus Ams., is endemic to Israel, where it spends its 10.5 month larval period feeding only in the leaves of Quercus calliprinos Webb.
2. We compared patterns of egg deposition and sources of larval mortality to test whether oviposition patterns and site preferences confer an enhanced likelihood of larval survival.
3. Dominant sources of larval mortality were premature leaf abscission and death from unknown causes, whereas predation, parasitism and intraspecific interference accounted for relatively little larval mortality.
4. Eggs, and thus mines, were aggregated among leaves of host trees even though premature leaf abscission was positively correlated with density of mines per leaf. Interference competition among larvae was the only other density-dependent mortality factor.
5. Oviposition patterns within leaves mitigated the probability of death from larval interference, and probably also from early leaf abscission.
6. Despite these density-dependent mortality factors, overall probability of larval survival to pupation was independent of initial density of mines on a leaf.
7. The long larval period allows synchrony between oviposition flights and times of predictable resource availability.     

Predation risk perception and food scarcity induce alterations of life-cycle traits of the mosquito Culex pipiens

Abstract.  1. Predators may affect prey populations by direct consumption, and by inducing defensive reactions of prey to the predation risk. Food scarcity frequently has effects on the inducible defences of prey, but no consistent pattern of food–predation risk interaction is known.
2. In this study the combined effect of food shortage and predation-risk perception in larvae of the mosquito Culex pipiens was investigated. Water exposed to the aquatic predator bug Notonecta glauca was used as a source of predation intimidation. Mosquito larvae were reared in three different media containing either no predator cues or the cues of N. glauca that had been fed on either C. pipiens larvae or on Daphnia magna . Food was provided in favourable or limited amount for these set-ups.
3. The results showed that chemical cues from the predators fed with prey's conspecifics caused a decreased survival, delayed pre-imaginal development, and reduction in body size of emerged mosquitoes, whereas chemical cues from predators fed with D. magna caused only delayed development. Food scarcity significantly exacerbates the negative effect of the predator cues on pre-imaginal development of C. pipiens . Effects of the cues on larval development and body size of imagoes are significantly stronger for females than for males.
4. The present study suggests that when food is limited, predators can affect population dynamics of prey not only by direct predation, but also by inducing lethal and sublethal effects due to perception of risk imposed by chemical cues. To understand the effects of predators on mosquito population dynamics, environmental parameters such as food deficiency should be considered.     

Spatial responses of two herbivore groups to a geometrid larva on mountain birch

Direct or plant-mediated interactions between herbivores may modify their spatial distribution among and within plants. In this study, we examined the effect of a leaf-chewing geometrid, the autumnal moth (Epirrita autumnata), on two different herbivore groups, leaf rolling Deporaus betulae weevils and Eriocrania spp. leafminers, both feeding on mountain birch (Betula pubescens ssp. czerepanovii). The exact locations of herbivores within tree canopies were mapped during three successive summers. In the first 2 years, some trees were artificially colonized by eggs of the autumnal moth to induce both rapid and delayed resistance in the foliage. The natural infection levels of the pathogenic rust fungus (Melampsoridium betulinum), potentially involved in species interactions, were also recorded. At the level of the whole tree, the density of D. betulae leaf rolls was lower in trees infested by the autumnal moth in the same year. However, the feeding locations within trees were partly segregated: D. betulae favoured shadier branches, while E. autumnata preferred the sunny parts of the canopy. The autumnal moth did not affect current- or following-year density of leafminers at the tree or branch level. Trees infected by rust had fewer leafminers in the same summer than noninfected trees. There were no interaction effects between defoliation by the autumnal moth and rust infection, and no delayed effects on the abundance of other herbivores the following year. Taken together, these findings suggest that the autumnal moth has a negative, partially plant-mediated impact on D. betulae, and can reduce the extent of current-year defoliation caused by D. betulae. This may be beneficial for the mountain birch, since the greater part of D. betulae damage occurs around or after the end of the larval period of the autumnal moth, which may be a critical time for tree recovery after moth outbreaks.     

Population dynamics of the white pine weevil, Pissodes strobi, infesting jack pine, Pinus banksiana, in Ontario, Canada

1. Eight generations of white pine weevil, Pissodes strobi (Peck) , infesting the terminal shoots of young jack pine trees, were sampled as larvae, pupae, and adults between 1988 and 1995.
2. The density of adult weevils increased rapidly for the first 3 years of the study, then declined in the next 4 years. Between-generation rates of change in density of weevils were related to mean number of weevils produced per terminal shoot of the host tree. Net increases in population density followed years in which a relatively high number of weevils emerged per terminal shoot and vice versa.
3. The mean number of weevils emerging per terminal shoot was determined by survival of weevils between the end of the larval stage and the successful emergence of adult weevils.
4. There was no clear relationship between survival of the post-feeding stages of the weevil and rates of parasitism or bird predation on these same stages. There was, however, a strong negative relationship between survival of weevils within the terminal shoot and the abundance of the facultative dipteran predator Lonchaea corticis Taylor.
5. The relationship among generational rates of change in populations, survival, and predation was evident irrespective of the age of the trees infested, suggesting that population dynamics of the weevil may be influenced as much by local predation pressure as by physical and biotic changes associated with growth of the young host trees.