The effect of spatial scale on interactions between two weevils and their food plant

The effect of spatial scale on the interactions between the weevils Gymnetron pascuorum Gyll. and Mecinus pyraster Herbst and their host plant, ribwort plantain Plantago lanceolata L., was studied. Both weevils developed in plantain seedheads but occupied different niches within the seedhead. Seedheads were sampled annually from 162 plants at each of two experimental sites consisting of a series of habitat patches of two distinct sizes. Data were analysed from three site-years. Our results suggest that the density of available seedheads varied among years and this had a direct effect on abundance. M. pyraster, which develops in the stem within the seedhead, was more sensitive to changes in seedhead density than was G. pascuorum, which develops within the seeds themselves. The presence of a hedgerow along one side of the experimental site affected the pattern of colonisation of newly-created habitat patches by G. pascuorum but not by M. pyraster. Changes in spatial scale did not affect the variability of seedhead and insect densities. G. pascuorum had an aggregated distribution at all the spatial scales considered, but the distribution of M. pyraster was very scale dependent. The distributions of the two weevil species were positively associated amongst infested plants but not amongst infested seedheads. Behavioural and ecological factors that could explain the results of the data analyses are discussed.     

Adjacent trophic-level effects on spatial density dependence in a herbivore—predator—parasitoid system

Abstract. 1. Spatial density dependence of enemy/victim relationships (predator/prey and host/parasitoid) were examined for the gall-making herbivore Eurosta solidaginis (Diptera: Tephritidae), its predator Mordellistena unicolor (Coleoptera: Mordellidae), and a parasitoid of Mordellistena, Schizoprymnus sp. (Hymenoptera: Braconidae), both before and after experimental perturbation of gall density.
2. Mordellistena predation did not depend on Eurosta density, nor did Schizoprymnus parasitism of Mordellistena depend on Mordellistena density.
3. Schizoprymnus parasitism of Mordellistena depended strongly on the density of Eurosta , the prey of its host.
4. The lack of density dependence in enemy/victim relationships may be explained by effects from adjacent trophic-levels. Ovipositing Schizoprymnus may search for high densities, not of its host Mordellistena , but of its host's prey ( Eurosta galls), and ovipositing Mordellistena may avoid patches of high gall density where the risk of being parasitized is greater.     

Dynamics of parasitism in the organ-pipe wasp, Trypoxylon politurn: effects of spatial scale on parasitoid functional response

Abstract.

• 1 Life tables and rates of parasitism were tabulated from mud nests built by Trypoxylon politum (Hymenoptera: Sphecidae) at nine different nesting sites from Missouri and Mississippi.
• 2 Most developmental mortality occurred either during the first two instars of development, or during the inactive prepupal phase. The majority (76%) of deaths were caused by insect parasitoids and cleptoparasites. Levels of parasitism and survivorship varied among nesting sites, and among locations within the two sites surveyed at a fine spatial scale.
• 3 Total developmental mortality, K, was positively associated with the number of hosts (immature T.politum) per site. Within one of two sites sampled at a fine spatial scale, K was negatively associated with the local density of hosts. Levels of total parasitism were positively associated with host population size, and negatively associated with local host density within one of the two sites sampled at a fine spatial scale.
• 4 Levels of parasitism by Melittobia (Hymenoptera: Eulophidae) were positively associated with the number of hosts per site, but negatively associated with the local density of hosts within sites. Melittobia parasitism was also negatively associated with the local density of old nesting material within sites.
• 5 Parasitism by Melittobia was a function of both the numbers of nests per quadrat and the mean nest size per quadrat at one of the two sites surveyed at a fine scale. At the other site, parasitism by Melittobia was a function of mean nest size per quadrat.
• 6 The life cycle and nesting behaviour of T.politum, in relation to the regulation of its numbers, is discussed.

     

Parasitoid aggregation and interference in host–parasitoid dynamics

1. Parasitoids do not distribute themselves evenly among available patches, which has an important bearing on the dynamics of host–parasitoid interactions. This study examined the density‐dependent nature of aggregation of the parasitoid Dirhinus giffardii Silvestri on the oriental fruit fly host, Bactrocera dorsalis (Hendel) distributed among discrete patches. 2. Four artificial patches were created in a cage, and the number of hosts in each patch was manipulated. Parasitoids were released into the cage, and whether parasitoid density and host density influence the degree of parasitoid aggregation was examined. 3. Parasitoid aggregation became stronger (e.g. uneven distribution among patches) as the parasitoid density decreased and also as the host density increased. The index of parasitoid aggregation was not influenced by the distribution of hosts among patches. 4. The empirically characterised aggregation pattern was incorporated in a host–parasitoid model that consists of one host and one parasitoid species. The analysis of the model shows that an unstable system (i.e. the coexistence of the host and parasitoid is impossible) can be stabilised (i.e. coexistence is possible) when the parasitoid aggregates in a way that is consistent with the pattern found in the experiment.     

Dispersal and spatial scale affect synchrony in spatial population dynamics

A large body of theoretical studies has shown that synchrony among populations is critical for the long-term persistence of species in fragmented habitats. Although the effects of dispersal and environmental factors on synchrony have been investigated theoretically, empirical studies of these relationships have been lacking. We explored the interplay between environmental and demographic factors (fecundity, survival, dispersal) on population synchrony for 53 species of birds. We show that the interspecific differences in mean synchrony were determined by global environmental factors whose action was probably mediated by the abundance of each species. After removing the effect of these global factors on synchrony, the residual synchrony was strongly correlated with dispersal distance. The relationship between dispersal and synchrony was stronger for the species nesting in wet habitats than for those nesting in dry habitats. Our results indicate that different factors synchronize bird populations at different spatial scales, thus highlighting the role of scale in understanding spatial population dynamics and extinction risks.     

Relationships between parasitoid host range and host defence: a comparative study of egg encapsulation in two related parasitoid species

Abstract. Parasitoid host range may proceed from traits affecting host suitability, traits affecting parasitoid foraging behaviour, or both. We tested the hypothesis that encapsulation can be used as a reliable indicator of parasitoid host range in two closely related larval endoparasitoids of Lepidoptera. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and a generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). We determined the effects of host species ( Pieris brassicae (L.), P. napi (L.) and P. rapae ) (Lepidoptera: Pieridae) and host developmental stage (early first, second and third instar) on encapsulation of parasitoid eggs. Host species and parasitoid species, as well as the resulting interaction between these two factors had significant effects on encapsulation of Cotesia eggs. Encapsulation in Pieris hosts was much lower for C. glomerata (<34%, except for second and third instar of P. rapae ) than for C. rubecula (>32%), even when the latter was parasitizing P. rapae. Encapsulation increased with the age of the larvae, although the only significant difference was for C. glomerata. Overall, P. rapae showed a stronger encapsulation reaction than P. brassicae and P. napi. Encapsulation levels of C. glomerata corresponded well to patterns of female host species and host age preference for oviposition and parasitoid larval performance. In contrast, percentages of encapsulation of C. rubecula were not consistent with host preference and host suitability. We argue that encapsulation alone is unlikely to provide a sufficient explanation for C. glomerata and C. rubecula host range.     

Linking the coevolutionary and population dynamics of host–parasitoid interactions

The interplay between coevolutionary and population or community dynamics is currently the focus of much empirical and theoretical consideration. Here, we develop a simulation model to study the coevolutionary and population dynamics of a hypothetical host–parasitoid interaction. In the model, host resistance and parasitoid virulence are allowed to coevolve. We investigate how trade-offs associated with these traits modify the system's coevolutionary and population dynamics. The most important influence on these dynamics comes from the incorporation of density-dependent costs of resistance ability. We find three main outcomes. First, if the costs of resistance are high, then one or both of the players go extinct. Second, when the costs of resistance are intermediate to low, cycling population and coevolutionary dynamics are found, with slower evolutionary changes observed when the costs of virulence are also low. Third, when the costs associated with resistance and virulence are both high, the hosts trade-off resistance against fecundity and invest little in resistance. However, the parasitoids continue to invest in virulence, leading to stable host and parasitoid population sizes. These results support the hypothesis that costs associated with resistance and virulence will maintain the heritable variation in these traits found in natural populations and that the nature of these trade-offs will greatly influence the population dynamics of the interacting species. Received: December 20, 1999 / Accepted: July 17, 2000     

Effects of variation among plant species on the interaction between a herbivore and its parasitoid

Abstract.  1. Previous studies have demonstrated that phenotypic traits of plants have the potential to affect interactions between herbivores and their natural enemies. Consequently, the impact of natural enemies on herbivore vital rates and population dynamics may vary among plant species. This study was designed to investigate the potential for density-dependent parasitism of an aphid herbivore feeding on six different host plant species.
2. Population densities of the aphid Aphis nerii B de F (Homoptera: Aphididae) and its parasitoid Lysiphlebus testaceipes Cresson (Hymenoptera: Braconidae) were recorded within a single growing season on six different species of milkweed in the genus Asclepias L. (Asclepiadaceae). Asclepias species are known to vary in their quality as food for herbivores. Although data on plant quality were not available in this study, population data were analysed to determine the effects of different Asclepias species on rates of parasitism and aphid population growth.
3. Parasitism rates of A. nerii varied among Asclepias species but were temporally density dependent over at least some range of aphid density on all plant species. Aphid population growth rates also varied among Asclepias species, and declined with an increase in the maximum parasitism rates among plant species; however, in no case was density-dependent parasitism sufficient to prevent exponential population growth of aphids within the growing season. The results serve to emphasise that, if natural enemies are to regulate herbivore populations, density-dependent mortality is a necessary, but not sufficient, condition for regulation.     

Additive and non-additive effects from a larger spatial scale determine small-scale densities in a web spider <Emphasis Type="Italic">Neriene brongersmai</Emphasis>

We examined 15 populations of Neriene brongersmai, a common sheet-web spider inhabiting cedar forest floor, to find out how density at a small spatial scale (patch level) is determined by processes operating at this scale as well as those from a larger spatial scale (population level). Here we focus on two types of large-scale effects that may influence small-scale density: an additive effect that changes the density at patch level; and a non-additive effect that changes the relationship between the density and its limiting factor at the patch level. ANCOVA showed that patch-level density of this spider was positively correlated with web-site availability at this level, but the density with a given amount of web-site differed among populations (cedar forests), indicating the existence of an additive large-scale effect. Multiple regression analysis showed that web-site availability at a population level explained the additive large-scale effect well, but prey availability and forest size did not. It seemed likely that increased web-site availability may have reduced the mortality of spiders while moving to a new web-site, and hence increased population density. A non-additive large-scale effect was also revealed: i.e. the relationship between density and web-site availability at the patch level tended to be stronger in populations with a greater additive large-scale effect. Higher intraspecific competition for web-sites in these populations appeared to have strengthened this relationship.     

The effects of parasitoid fecundity and host taxon on the biological control of insect pests: the relationship between theory and data

1. A simple, intuitive argument and the tenets of the biological control literature both suggest that, in general, parasitoids with a greater fecundity will provide better control of their hosts, and will thus be better biological control agents. 2. A model of host-parasitoid dynamics, based on the standard Thompson–Nicholson–Bailey approach and incorporating the effects of parasitoid fecundity-limitation and host density-dependence, also indicates that as parasitoid fecundity decreases so does local stability and the degree of host suppression. 3. A taxonomically diverse data set obtained from the biological control record failed to support this theoretical prediction, but at the same time indicated a strong effect of host taxon on the outcome of biological control. 4. The hypothesis that the fecundity of parasitoids is correlated positively with their ability to suppress host populations is supported by data exclusively from the host order Lepidoptera. 5. Possible explanations for the divergence between the fecundity-limitation hypothesis and the complete data set include: the ability of parasitoids to provide long-term control of pests without the presence of a stable host–parasitoid equilibrium; differences between the concepts of successful control in theory and practice; evolutionary trade-offs between fecundity and other parasitoid life-history features, such as search efficiency, leading to better pest control by parasitoids with low fecundity; and differing windows of vulnerability to parasitoid attack between host taxa.     

A lagged, density-dependent relationship between jack pine budworm Choristoneura pinus pinus and its host tree Pinus banksiana

Abstract. 1. Survival of newly emerged jack pine budworm Choristoneura pinus pinus is related to the density of available pollen cones (microsporangiate strobili) produced by its host tree, jack pine Pinus banksiana. 2. A 7‐year time series of observations from a plot network in Ontario, Canada, compared the propensity of jack pine to produce pollen cones, τ, on trees that were either defoliated or undisturbed by the jack pine budworm. 3. Non‐defoliated jack pine trees have a high propensity to produce pollen cones. More than one‐third of these trees produced pollen cones in every year of the series. Propensity varied significantly among plots and trees. Temporal patterns in propensity were also highly variable but within a plot propensity was often autocorrelated in time. 4. Defoliation by the jack pine budworm was associated with forest plots composed of the oldest and the largest trees and with the fewest trees per hectare. Within a plot, outbreaks lasted 3 or 4 years although individual trees were only defoliated in 1 or 2 years. 5. The propensity to produce pollen cones in jack pine was reduced in the years after defoliation. The most pronounced reductions in propensity occurred where defoliation was most severe. 6. The reduction in propensity to produce pollen cones resulting from previous defoliation, coupled with the dependence of jack pine budworm survival on the availability of pollen cones, induces a lagged, negative feedback between the density of the consumer and that of its resource. 7. The lagged, density‐dependent relationship between jack pine budworm and its jack pine host contributes to oscillatory dynamics of the jack pine budworm. Comparison of the outbreak behaviour of jack pine budworm with that of the closely related eastern spruce budworm C. fumiferana suggests that differences in the strength of the host‐plant interaction may account for differences in the relative frequency of outbreaks in the respective systems.     

The influence of cultivar and cultivar-aphid odours on the olfactory response of the parasitoid Aphidius colemani

The olfactory response of the parasitoid Aphidius colemani (Viereck) (Hymenoptera: Braconidae) to odours in a tritrophic system involving three cultivars of common cabbage, Brassica oleracea var capitata, characterized by different levels of susceptibility to Myzus persicae (Sulzer) (Hemiptera: Aphididae) was studied in a four‐way olfactometer. Odours influenced A. colemani response in the olfactometer to varying degrees. The magnitude of parasitoid response to odours of uninfested cabbage depended on cultivar, with Derby Day [green‐leaved, susceptible to M. persicae and the crucifer specialist, Brevicoryne brassicae (Linnaeus) (Hemiptera: Aphididae)] and Minicole (green‐leaved, partially resistant with known antibiosis factors for B. brassicae) preferred over Ruby Ball (red‐leaved with antixenosis factors for M. persicae and B. brassicae). The odour of the cabbage cultivar on which the parasitoid had been reared was preferred over the other cultivars. However, when provided with a choice between odours of infested plants, parasitoids did not show a significant preference for the cultivar on which they were reared. Results from the study show that parasitioids differentiated between odour of the three cultivars in dependence of their rearing history when the plant is uninfested.     

The temporal correlation and spatial synchrony in the stemborer and parasitoid system of Coast Kenya with climate effects

The spatial synchrony and the temporal auto-correlationship of the exotic stemborer Chilo partellus (Swinhoe) (Lepidoptera: Crambidae) and the indigenous Sesamia calamistis Hampson (Lepidoptera: Noctuidae) and Chilo orichalcociliellus (Strand) (Lepidoptera: Crambidae), and crosscorrelationship with their indigenous and introduced larval parasitoids Cotesia sesamiae (Cameron) and Cotesia flavipes Cameron (Hymenoptera: Braconidae) was studied using 3-year data collected in coastal Kenya. An autoregressive model was used to study the effect of climatic stochasticity or population density-dependent factors on stemborer populations. It appeared that rainfall did have a direct impact on stemborers in the south coast and an indirect one in the north coast. Spatial nonparametric correlation functions (SNCF) and cross-correlation functions (SNCCF) were applied for spatial synchrony analysis. The regional synchrony of Ch. partellus and S. calamistis decreased and that of Ch. orichalcociliellus increased after the introduction of Co. flavipes. The positive crosscorrelation coefficient between stemborers and parasitoids suggests a synchrony between the pest and its natural enemy.     

The geographical variation of network structure is scale dependent: understanding the biotic specialization of host–parasitoid networks

Research on the structure of ecological networks suggests that a number of universal patterns exist. Historically, biotic specialization has been thought to increase towards the Equator. Yet, recent studies have challenged this view showing non‐conclusive results. Most studies analysing the geographical variation in biotic specialization focus, however, only on the local scale. Little is known about how the geographical variation of network structure depends on the spatial scale of observation (i.e. from local to regional spatial scales). This should be remedied, as network structure changes as the spatial scale of observation changes, and the magnitude and shape of these changes can elucidate the mechanisms behind the geographical variation in biotic specialization. Here we analyse four facets of biotic specialization in host–parasitoid networks along gradients of climatic constancy, classifying the networks according to their spatial extension (local or regional). Namely, we analyse network connectance, consumer diet overlap, consumer diet breadth, and resource vulnerability at both local and regional scales along the gradients of both current climatic constancy and historical climatic change. While at the regional scale none of the climatic variables are associated to biotic specialization, at the local scale, network connectance, consumer diet overlap, and resource vulnerability decrease with current climatic constancy, whereas consumer generalism increases (i.e. broader diet breadths in tropical areas). Similar patterns are observed along the gradient of historical climatic change. We provide an explanation based on different beta‐diversity for consumers and resources across the geographical gradients. Our results show that the geographical gradient of biotic specialization is not universal. It depends on both the facet of biotic specialization and the spatial scale of observation.     

Spatial variability of hosts,parasitoids and their interactions across a homogeneous landscape

Species assemblages and their interactions vary through space, generating diversity patterns at different spatial scales. Here, we study the local‐scale spatial variation of a cavity‐nesting bee and wasp community (hosts), their nest associates (parasitoids), and the resulting antagonistic network over a continuous and homogeneous habitat. To obtain bee/wasp nests, we placed trap‐nests at 25 sites over a 32 km² area. We obtained 1,541 nests (4,954 cells) belonging to 40 host species and containing 27 parasitoid species. The most abundant host species tended to have higher parasitism rate. Community composition dissimilarity was relatively high for both hosts and parasitoids, and the main component of this variability was species turnover, with a very minor contribution of ordered species loss (nestedness). That is, local species richness tended to be similar across the study area and community composition tended to differ between sites. Interestingly, the spatial matching between host and parasitoid composition was low. Host β‐diversity was weakly (positively) but significantly related to geographic distance. On the other hand, parasitoid and host‐parasitoid interaction β‐diversities were not significantly related to geographic distance. Interaction β‐diversity was even higher than host and parasitoid β‐diversity, and mostly due to species turnover. Interaction rewiring between plots and between local webs and the regional metaweb was very low. In sum, species composition was rather idiosyncratic to each site causing a relevant mismatch between hosts and parasitoid composition. However, pairs of host and parasitoid species tended to interact similarly wherever they co‐occurred. Our results additionally show that interaction β‐diversity is better explained by parasitoid than by host β‐diversity. We discuss the importance of identifying the sources of variation to understand the drivers of the observed heterogeneity.