Foraging for solitarily and gregariously feeding caterpillars: A comparison of two related parasitoid species (Hymenoptera: Braconidae)

In the present study we apply a comparative approach, in combination with experimentation, to study behavior of two parasitoid species that attack caterpillar hosts with different feeding strategies (gregarious or solitary). In a semifield setup, consisting of clean cabbage plants and plants infested with one of two host species, the foraging behavior of the specialistCotesia rubecula, on obligate parasitoid of solitarily feedingPieris rapae larvae, was compared to that of the generalistCotesia glomerata, a polyphagous parasitoid of several Pieridae species (mainly the gregariously feedingPieris brassicae).Cotesia glomerata displayed equal propensity to search for and parasitize larvae of both host species. AlthoughC. glomerata exhibited a relatively plastic foraging behavior in that it searched differently under different host distribution conditions, its behavior seems more adapted to search for gregariously feeding hosts. Females exhibited a clear area-restricted search pattern and were more successful in finding the gregariously feeding caterpillars.Cotesia rubecula showed a higher propensity to search forP. rapae than forP. brassicae, i.e., females left the foraging setup significantly earlier when their natural hostP. rapae was not present.C. rubecula showed a more fixed foraging behavior, which seems adapted to foraging for solitarily feeding host larvae. In a setup with onlyP. rapae larvae, the foraging strategies of the two parasitoid species were quite similar. In a choice situationC. glomerata did not show a preference for one of the host species, whileCotesia rubecula showed a clear preference for its natural host species. The latter was shown by several behavioral parameters such as the number of first landings, allocation of search time, and percentage parasitization.     

Demonstration of Cytotoxicity against Wasps by Pierisin-1: A Possible Defense Factor in the Cabbage White Butterfly

The cabbage white butterfly, Pieris rapae, produces pierisin-1, a protein inducing apoptosis of mammalian cells. In the present study, the biological activity of pierisin-1 as a protective agent against parasitic wasps for P. rapae was examined. Pierisin-1 caused detrimental effects on eggs and larvae of non-habitual parasitoids for P. rapae, Glyptapanteles pallipes, Cotesia kariyai and Cotesia plutellae at 1–100 µg/ml, levels essentially equivalent to those found in P. rapae larvae. In contrast, eggs and larvae of the natural parasitoid of P. rapae, Cotesia glomerata proved resistant to the toxicity of pierisin-1 through inhibition of pierisin-1 penetration of the surface layer. The expression level of pierisin-1 mRNA in the larvae of P. rapae was increased by parasitization by C. plutellae, whereas it was decreased by C. glomerata. In addition, C. plutellae was associated with elevation of activated pierisin-1 in the hemolymph. From these observations, it is suggested that pierisin-1 could contribute as a defense factor against parasitization by some type of wasps in P. rapae.     

Coexistence and niche segregation by field populations of the parasitoids Cotesia glomerata and C. rubecula in the Netherlands: predicting field performance from laboratory data

Field experiments with foraging parasitoids are essential to validate the conclusions from laboratory studies and to interpret differences in searching and host selection behaviour of parasitoid species. Furthermore, field experiments can indicate whether the parameters measured in the laboratory are relevant to elucidation of the ecological processes under study, such as adaptation or species interactions. In previous extensive laboratory studies we studied plant- and host-searching behaviour, host acceptance, host suitability; host plant preference, and learning of two congeneric parasitoids of Pieris caterpillars: the generalist Cotesia glomerata, which has been reported to attack several Pieridae species, and C. rubecula, a specialist of the small cabbage white Pieris rapae. In the present field study our aim was to verify the importance of these previous laboratory findings for explaining the performance of these two species in the field. We investigated experimentally whether parasitism on three Pieris species varied with parasitoid species and with food plant of the caterpillars. We exposed different types of host plants, infested with different Pieris species, to parasitism by natural populations of Cotesia species, by setting the experimental plants out in Brussels sprouts cabbage fields. Furthermore we made direct observations of parasitoid foraging in the field. In general, the field results confirmed our predictions on the range of host plant and host species used in the field. The two Cotesia species appear to coexist through niche segregation, since C. glomerata was mainly recovered from P. brassicae and C. rubecula from P. rapae. Although C. glomerata is a generalist at the species level, it can be a specialist at the population level under certain ecological circumstances. Our study shows the importance of variation in host plant attraction and host species acceptance in restricting host plant and host diet in the field. Furthermore the results suggest that, at least in the Netherlands, specialisation of C. glomerata on P. brassicae may occur as a result of C. rubecula outcompeting C. glomerata in P. rapae larvae. Received: 8 July 1999 / Accepted: 31 January 2000     

First detailed report of brood parasitoidism in the invasive population of the paper wasp Polistes dominulus (Hymenoptera, Vespidae) in North America

The European paper wasp Polistes dominulus (Christ) is a model system in the fields of behavioral ecology, ecological immunology, and invasion biology. Since its introduction to the US in 1978, its invasion success has been attributed, in part, to a lack of parasites or parasitoids infecting this population. This is despite the number of parasites which infest the native population and the generalist polistine parasites and parasitoids documented in sympatric North American species. Multiple studies have cited low parasite pressure as evidence that the invasive population of P. dominulus is benefiting from a post-invasion release from enemies. Here, we present the first well documented case of parasitoidism of the invasive population of P. dominulus in North America.     

Persistence in Massachusetts of the veined white butterfly due to use of the invasive form of cuckoo flower

The native pierid butterfly Pieris oleracea underwent a large range reduction in New England in the twentieth century, likely due to the introduction the invasive butterfly Pieris rapae (Lep.: Pieridae) to North America in 1860, and later the oligophagous parasitoid Cotesia glomerata (Hymenoptera: Braconidae) in 1884. Thought extirpated from the state by the 1970s, one large dense population of the butterfly was found in the mid 1980s in a flood plain meadow along the Housatonic River in Lenox, Berkshire Co., Massachusetts. We examined how this native pierid was able to maintain a relatively dense local population by feeding on a novel, invasive host plant, Cardamine pratensis (cuckoo flower), in a meadow habitat despite known parasitoid presence. We approached this question in three ways. First, we deployed trap host plants (cuckoo flower and collards) stocked host larvae (first and second instars of either P. rapae or P. oleracea) at the Lenox site and other locations to determine current rates of C. glomerata attack, for comparison with historical information. Second, we used olfactometer experiments to determine if C. glomerata females could detect the cuckoo flower volatiles released during P. oleracea larval feeding. Third, we used field-cage experiments to determine if the plant architecture found in the flood plain meadow inhibited the ability of C. glomerata females to locate and parasitize hosts. Specifically, we asked if overtopping vegetation prevented or reduced parasitism of P. oleracea larvae feeding on the covered basal rosettes of C. pratensis, which is the physical form of host plant for three of the four butterfly generations at the site.     

Food plant and herbivore host species affect the outcome of intrinsic competition among parasitoid larvae

1. In nature, several parasitoid species often exploit the same stages of a common herbivore host species and are able to coexist despite competitive interactions amongst them. Less is known about the direct effects of resource quality on intrinsic interactions between immature parasitoid stages. The present study is based on the hypothesis that variation in the quality or type of plant resources on which the parasitoids indirectly develop may be complementary and thus facilitate niche segregation favouring different parasitoids in intrinsic competition under different dietary regimes. 2. The present study investigated whether two herbivore species, the cabbage butterflies Pieris brassicae and Pieris rapae (Pieridae), and the quality of two important food plants, Brassica oleracea and Brassica nigra (Brassicaceae), affect the outcome of intrinsic competition between their primary larval endoparasitoids, the gregarious Cotesia glomerata (Braconidae) and the solitary Hyposoter ebeninus (Ichneumonidae). 3. Hyposoter ebeninus is generally an intrinsically superior competitor over C. glomerata. However, C. glomerata survived more antagonistic encounters with H. ebeninus when both developed in P. brassicae rather than in P. rapae caterpillars, and while its host was feeding on B. nigra rather than B. oleracea. Moreover, H. ebeninus benefitted from competition by its higher survival in multiparasitised hosts. 4. These results show that both plant and herbivore species mediate the battleground on which competitive interactions between parasitoids are played out and may affect the outcomes of these interactions in ways that enable parasitoids to segregate their niches. This in turn may promote coexistence among parasitoid species that are associated with the same herbivore host.     

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (±)-catechin

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the “novel weapons” hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the “evolution of increased competitive ability” hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (±)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Response of the wasp (<Emphasis Type="Italic">Cotesia glomerata</Emphasis>) to larvae of the large white butterfly (<Emphasis Type="Italic">Pieris brassicae</Emphasis>)

The large white butterfly (Pieris brassicae L) first invaded northernmost Japan from Siberia around 1994, and after a few years, began to expand its range. The wasp, Cotesia glomerata (L) parasitizes larvae of the small white butterfly (Pieris rapae crucivora Boisduval), a usual host in the same geographic area. Some Pieris brassicae larvae in Hokkaido have been parasitized by Cotesia glomerata, but the parasitism rate of Pieris brassicae larvae tends to be lower than that of Pieris rapae. To examine the process of parasitizing Pieris brassicae larvae, we observed how the parasitoid wasp responded to the host larvae on damaged leaves. Cotesia glomerata females tended to avoid Pieris brassicae larvae, and even when female wasps inserted their ovipositors into Pieris brassicae larvae, none laid eggs. The parasitoids obtained from Pieris rapae larvae failed to parasitize Pieris brassicae during the host-acceptance step.     

Parasitism by Cotesia glomerata Induces Immunosuppression of Pieris rapae: Effects of Ovarian Protein and Polydnavirus

A gregarious endoparasitoid wasp, Cotesia glomerata, parasitizes the cabbage butterfly, Pieris rapae. During wandering larval stage for pupal metamorphosis, the parasitoid larvae egress from the parasitized host to form cocoons thus eventually leading to death of the host. This study focused on the effect of C. glomerata parasitization on cellular immune response of P. rapae. For this purpose, an ideal anticoagulant buffer was formulated to procure the hemocytes in native form with morphological, behavioral, and functional characteristics. The hemocytes selectively encapsulated only DEAE beads under in vitro conditions and a quantitative study revealed about 70% of the beads being encapsulated. On the other hand, calyx fluid from C. glomerata injected to P. rapae markedly inhibited the spreading ability of the hemocytes in a dose-dependent manner and also attenuated the in vitro encapsulation response of the hemocytes against the cationic bead. The calyx fluid contained polydnavirus as well as ovarian proteins. The isolated polydnavirus genome consisted of variously sized-segments with their unequal amounts. The P. rapae injected with the calyx fluid expressed several polydnaviral genes within 2 h. These results suggest that the immunosuppression of the parasitized P. rapae may be induced by the polydnaviral gene products as well as ovarian proteins.     

Effect of belowground herbivory on parasitoid associative learning of plant odours

Root herbivores can influence both the performance and the behaviour of parasitoids of aboveground insect herbivores through changes in aboveground plant quality and in the composition of the plant's odour blend. Here we show that root herbivory by Delia radicum larvae did not influence the innate preferences for plant odours of the two closely related parasitoid species Cotesia glomerata and C. rubecula, but did affect their learned preferences, and did so in an opposite direction. While C. glomerata learned to prefer the odour of plants with intact roots, C. rubecula learned to prefer the odour of root‐infested plants. The learned preference of C. glomerata for the odour of plants with intact roots matches our previously published result of its better performance when developing in P. brassicae hosts feeding on this plant type. In contrast, the relatively stronger learned preference of C. rubecula for the odour of root‐infested plants cannot be merely explained by its performance, as the results of our present study indicate that D. radicum root herbivory did not influence the performance of C. rubecula nor of its host P. rapae. Our results stress the importance of assessing the influence of root herbivores on both innate and learned responses of parasitoids to plant odours.     

The role of host species, age and defensive behaviour on ovipositional decisions in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

The main objective of this study was to determine the extent to which host acceptance behaviour as related to host species, age, and defensive behaviour might explain the differences in host use that exist between two congeneric and sympatric species of parasitic wasps. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). Cotesia species differed in their responses to host species (P. brassicae (L.), P. napi (L.) and P. rapae) and developmental stage (early and late 1st, 2nd and 3rd instars). In no-choice tests, host acceptance by C. rubecula was higher for p. rapae and females did not distinguish among the 6 host ages. In contrast, when foraging for P. brassicae and P. napi, C. rubecula females more readily attacked early first instar. Cotesia glomerata showed a higher degree of behavioural plasticity towards acceptance of Pieris host species and host age than did C. rubecula. Cotesia glomerata females parasitized the three Pieris species and showed higher acceptance of first and second instars over third instar. Oviposition success was also influenced by host defensive behaviour. The frequency and the effectiveness of defensive behaviour rose with increasing age of the host, P. brassicae being the most aggressive Pieris species. Furthermore, the mean duration of C. glomerata oviposition was significantly reduced by the defensive reactions of P. brassicae, which would likely affect parasitoid fitness as oviposition time is positively correlated to clutch size in C. glomerata. Acceptance frequencies corresponded well to field reports of Pieris-Cotesia associations and to patterns of parasitoid larval performance, suggesting that the acceptance phase might be used as a reliable indicator of Cotesia host-specificity.     

The use of oviposition‐induced plant cues by Trichogramma egg parasitoids

1. Female parasitoids have evolved various foraging strategies in order to find suitable hosts. Egg parasitoids have been shown to exploit plant cues induced by the deposition of host eggs. 2. The tiny wasp Trichogramma brassicae uses oviposition‐induced cues from Brussels sprouts to locate eggs of the cabbage white butterflies Pieris brassicae and Pieris rapae that differ in their egg‐laying behaviour. These plant cues are elicited by male‐derived anti‐aphrodisiac pheromones in the accessory reproductive gland (ARG) secretions of mated female butterflies. However, the closely related generalist species Trichogramma evanescens does not respond to Brussels sprout cues induced by the deposition of P. brassicae egg clutches. 3. Here we showed in two‐choice bioassays that T. evanescens wasps respond to Brussels sprout cues induced by (i) the deposition of single eggs by P. rapae, and (ii) the application of ARG secretions from either mated P. rapae females, or from virgin female butterflies in combination with P. rapae's anti‐aphrodisiac compound indole. The wasps only associatively learned to respond to Brussels sprout cues after applying indole alone by linking those cues with the presence of P. rapae eggs. 4. Our results indicate that Trichogramma wasps more commonly exploit oviposition‐induced plant cues to locate their host eggs. Generalist wasps show less specificity in their response than specialists and employ associative learning.     

Nectar exploitation by herbivores and their parasitoids is a function of flower species and relative humidity

In conservation biological control, diversification of the agro ecosystem with flowering vegetation is seen as an important tool to support the broad range of predators and parasitoids that require nectar and pollen sources to survive and reproduce. In order to identify flowering plants that provide suitable food sources for natural enemies without supporting the pest species, we analyzed the exploitation of 19 flowering plants by two important lepidopteran cabbage pests, Pieris rapae and Plutella xylostella, and their hymenopteran parasitoids, Cotesia glomerata and Diadegma semiclausum. The experiments were conducted at 90% r.h., while Pieris rapae was tested both at 45% r.h. and at 90% r.h. At 45 ± 5% r.h., corresponding with field conditions at which P. rapae is predominantly active, the butterfly was unable to feed on a number of exposed floral nectar sources whose nectar was successfully exploited at 90% r.h. The broader nectar exploitation by P. rapae at the high humidity is presumably explained by the resulting decrease in nectar viscosity. When comparing D. semiclausum and its herbivorous host P. xylostella, the herbivore exploited a broader range of plants. However, those plants that benefited both the parasitoid and the herbivore had a much stronger effect on the longevity of the parasitoid. The results from the accessibility bioassay suggest that flowers where nectar is not accessible can have a negative impact on insect survival presumably by stimulating foraging without providing accessible nectar. Our results underline the importance of considering species-specific environmental conditions when fine-tuning the choice of nectar sources to be used in conservation biological control programs.     

Extended phylogeny of Aquilegia: the biogeographical and ecological patterns of two simultaneous but contrasting radiations

Studies of the North American columbines (Aquilegia, Ranunculaceae) have supported the view that adaptive radiations in animal-pollinated plants proceed through pollinator specialisation and floral differentiation. However, although the diversity of pollinators and floral morphology is much lower in Europe and Asia than in North America, the number of columbine species is similar in the three continents. This supports the hypothesis that habitat and pollinator specialisation have contributed differently to the radiation of columbines in different continents. To establish the basic background to test this hypothesis, we expanded the molecular phylogeny of the genus to include a representative set of species from each continent. Our results suggest that the diversity of the genus is the result of two independent events of radiation, one involving Asiatic and North American species and the other involving Asiatic and European species. The ancestors of both lineages probably occupied the mountains of south-central Siberia. North American and European columbines are monophyletic within their respective lineages. The genus originated between 6.18 and 6.57 million years (Myr) ago, with the main pulses of diversification starting around 3 Myr ago both in Europe (1.25–3.96 Myr ago) and North America (1.42–5.01 Myr ago). The type of habitat occupied shifted more often in the Euroasiatic lineage, while pollination vectors shifted more often in the Asiatic-North American lineage. Moreover, while allopatric speciation predominated in the European lineage, sympatric speciation acted in the North American one. In conclusion, the radiation of columbines in Europe and North America involved similar rates of diversification and took place simultaneously and independently. However, the ecological drivers of radiation were different: geographic isolation and shifts in habitat use were more important in Europe while reproductive isolation linked to shifts in pollinator specialisation additionally acted in North America.     

Evidence for multiple introductions of Centaurea stoebe micranthos (spotted knapweed,Asteraceae) to North America

Invasive species’ success may depend strongly on the genetic resources they maintain through the invasion process. We ask how many introductions have occurred in the North American weed Centaurea stoebe micranthos (Asteraceae), and explore whether genetic diversity and population structure have changed as a result of introduction. We surveyed individuals from 15 European native range sites and 11 North American introduced range sites at six polymorphic microsatellite loci. No significant difference existed in the total number of alleles or in the number of private alleles found in each range. Shannon–Weaver diversity of phenotype frequencies was also not significantly different between the ranges, while expected heterozygosity was significantly higher in the invasive range. Population structure was similar between the native range and the invasive range, and isolation by distance was not significant in either range. Traditional assignment methods did not allocate any North American individuals to the sampled European populations, while Bayesian assignment methods grouped individuals into nine genetic clusters, with three of them shared between North America and Europe. Invasive individuals tended to have genetically admixed profiles, while natives tended to assign more strongly to a single cluster. Many North American individuals share assignment with Romania and Bulgaria, suggesting two separate invasions that have undergone gene flow in North America. Samples from three other invasive range sites were genetically distinct, possibly representing three other unique introductions. Multiple introductions and the maintenance of high genetic diversity through the introduction process may be partially responsible for the invasive success of C. stoebe micranthos.     

Parasitoids deter foraging by Argentine ants (Linepithema humile) in their native habitat in Brazil

The Argentine ant, Linepithema humile, has invaded sites across Africa, Australia, Europe, and North America. In its introduced ranges it eliminates native ants and tends agricultural pests. Few studies have examined the ecology of Argentine ants in their native habitat. This study examined the effects of parasitoid flies, genus Pseudacteon, on the foraging behavior of Argentine ants in part of their native range in southern Brazil. Pseudacteon parasitoids commonly attacked Argentine ants, but not other ant species, in daylight at temperatures above 18°C. Argentine ants abandoned food resources and returned underground in the presence of parasitoids. Parasitoid attack rates diminished as Argentine ants retreated underground. Where parasitoids were present, Argentine ants were abundant at food resources only during times of day when parasitoids were inactive. Where parasitoids were absent, Argentine ants were abundant at food resources throughout the day. Overall, the presence of parasitoids explained observed variation in Argentine ant foraging far better than temperature, although temperature had some effect. The results suggest that Pseudacteon parasitoids inhibit the ability of Argentine ants to gather food resources in their native habitat in Brazil. Received: 11 December 1997 / Accepted: 12 June 1998     

AN ECOLOGICAL REVIEW OF CLADOPHORA GLOMERATA (CHLOROPHYTA) IN THE LAURENTIAN GREAT LAKES1

Cladophora glomerata (L.) Kütz. is, potentially, the most widely distributed macroalga throughout the world’s freshwater ecosystems. C. glomerata has been described throughout North America, Europe, the Atlantic Islands, the Caribbean Islands, Asia, Africa, Australia and New Zealand, and the Pacific Islands. Cladophora blooms were a common feature of the lower North American Great Lakes (Erie, Michigan, Ontario) from the 1950s through the early 1980s and were largely eradicated through the implementation of a multibillion‐dollar phosphorus (P) abatement program. The return of widespread blooms in these lakes since the mid‐1990s, however, was not associated with increases in P loading. Instead, current evidence indicates that the resurgence in blooms was directly related to ecosystem level changes in substratum availability, water clarity, and P recycling associated with the establishment of dense colonies of invasive dreissenid mussels. These results support the hypothesis that dreissenid mussel invasions may induce dramatic shifts in energy and nutrient flow from pelagic zones to the benthic zone.     

Invasion history of North American Canada thistle,Cirsium arvense

Aim Canada thistle (Cirsium arvense– Cardueae, Asteraceae) is one of the worst invasive plants world‐wide. Native to Eurasia, its unintentional introduction into North America now threatens the native flora and is responsible for enormous agricultural losses. The goals of this study are to: (1) reconstruct the evolutionary history of C. arvense and estimate how often it may have colonized North America, (2) compare the genetic diversity between European and North American populations to detect signs of demographic bottlenecks and/or patterns of population admixture, and (3) conduct bioclimatic comparisons to infer eventual niche shifts following this species’ introduction into North America. Location Europe and North America. Methods A total of 1522 individuals from 58 populations were investigated with six microsatellite markers. Estimates of heterozygosity (H_E) and allelic richness (R_S) were quantified for each population, and population structure was inferred via analyses of molecular variance (AMOVAs), principal components analyses (PCAs), Mantel tests and Bayesian clustering analyses. Climatic niche spaces were based on 19 bioclimatic variables extracted from approximately 32,000 locations covering the entire range, and compared using PCA and hierarchical cluster analysis. Results Although there is evidence of multiple introductions from divergent European lineages, North American populations of C. arvense exhibited significantly lower levels of genetic diversity than their putative ancestors. Bioclimatic comparisons pointed to a high degree of niche conservatism during invasion, but indicated that genotypes from the former USSR and Central European mountain chains were probably best adapted to invade North America upon entry into the continent. Main conclusions Genetic and historical data suggest that C. arvense first entered North America from Western Europe with the first European settlers, and was later introduced from Eastern Europe into the prairie states during the agricultural boom. The species went through a significant bottleneck following its introduction into the New World, but the level of genetic diversity remained high owing to admixture between genetically differentiated lineages and to a highly efficient outcrossing breeding system.     

The Physiological Ecology of the Zebra Mussel, Dreissena polymorpha, in North America and Europe

SYNOPSIS. The zebra mussel, Dreissena polymorpha (Pallas), wasintroduced into North America in 1986. Initial North American(N.A.) studies suggested that physiological responses variedbetween N.A. and European populations. However, literature reviewindicates agreement on most aspects of physiological adaptationincluding: respiratory responses; hypoxia/anoxia tolerance;salinity limits; emersion tolerance; freezing resistance; environmentalpH limits; calcium limits; starvation responses; and bioenergeticpartitioning. The main differences among N.A. and European musselsappear to be elevated upper thermal limits and temperaturesfor optimum growth among N.A. populations. N.A. zebra musselsprobably originated from the northern shore of the Black Seain the warmest portion of the mussel's European range. However,most European physiological data come from northern Europe wherepopulations may be adapted to colder temperatures. Alternatively,N.A. research suggests that mussels may have a capacity forseasonal temperature acclimatization such that responses recordedin warmer N.A. waters may be different from those recorded innorthern Europe even after short-term laboratory acclimation.Studies of genetic variation and physiological response amongEuropean and N.A. D. polymorpha populations are required toelucidate the basis for physiological differentiation. Recentlyevolved D. polymorpha has poor resistance adaptations comparedto unionacean and sphaeriid bivalves with longer freshwaterfossil histories. Poor resistance adaptations make it less suitedfor stable habitats, instead, its high fecundities, early maturity,and rapid growth are adaptations to unstable habitats whereextensive resistance adaptations are of little value.     

Dynamic effects of parasitism by an endoparasitoid wasp on the development of two host species: implications for host quality and parasitoid fitness

1. The study reported here examined growth and developmental interactions between the gregarious larval koinobiont endoparasitoid Cotesia glomerata (Hymenoptera: Braconidae) and two of its hosts that vary considerably in growth potential: Pieris rapae and the larger P. brassicae (Lepidoptera: Pieridae). At pupation, healthy larvae of P. brassicae are over twice as large, in terms of fresh body mass, as those of P. rapae. 2. Clutch size of C. glomerata was manipulated artificially, and the relationship between parasitoid burden and the maximum weight of the parasitised host (= host–parasitoid complex) was measured. In both hosts, the maximum complex weight was correlated positively with parasitoid burden. Compared with unparasitised hosts, however, the growth of P. rapae was increased at significantly lower parasitoid burdens than in P. brassicae. Emerging wasp size was correlated negatively with parasitoid burden in both host species, whereas development time was less affected. 3. After larval parasitoid egress, the weight of the host carcass increased slightly, but not significantly, with parasitoid burden, although there was a strong correlation between the proportion of host mass consumed by C. glomerata larvae during development and parasitoid burden. 4. Clutch size was generally correlated positively with instar parasitised in both hosts, and greater in P. brassicae than in P. rapae. Sex ratios were much more female biased in L1 and L2 P. rapae than in all other host classes. Adult parasitoid size was correlated inversely with host instar at parasitism, and wasps emerging from P. brassicae were larger, and completed development faster, than conspecifics emerging from P. rapae. 5. The data reveal that parasitism by C. glomerata has profound species‐specific effects on the growth of both host species. Consequently, optimality models in which host quality is often based on host size at parasitism or unparasitised growth potential may have little utility in describing the development of gregarious koinobiont endoparasitoids. The results of this investigation are discussed in relation to the potential effectiveness of gregarious koinobionts in biological control programmes.