High Host Density Research Articles

The impact of patch encounter rate on patch residence time of female parasitoids increases with patch quality

Abstract 1. For animal species that forage on patchily distributed resources, patch time allocation is of prime importance to their reproductive success. According to Charnov’s marginal value theorem (MVT), the rate of patch encounter should influence negatively the patch residence time: as the rate of patch encounter decreases, the patch residence time increases. Moreover, the MVT predicts that animals should stay longer in high quality patches.2. Using the aphid parasitoid Aphidius rhopalosiphi (Hymenoptera: Aphidiinae), the effects of these two factors (patch encounter rate and host density) were combined in order to test if the increment in patch residence time for a given decrease in patch encounter rate was larger for high quality patches than for low quality patches.3. The results show a significant effect of the interaction between the two factors. In high host density patches, parasitoids spent more time if they experienced a low patch encounter rate, while in low host density patches, patch encounter rate had no significant effect on the patch residence time. This suggests that the response of A. rhopalosiphi females to patch encounter rate varied with host density in the patch. Moreover, the same interaction effect was observed for the number of ovipositor contacts on aphids.4. Parasitoid females can use patch encounter rate to estimate patch density in the habitat but the effect of this estimate on their patch residence time is modulated by patch quality. Staying longer in a patch when patches are rare is more advantageous when the fitness gained by doing so is large. In low quality patches, the expected fitness gain is small and the female may gain more by leaving and taking her chance at finding another patch.

Short-distance dispersal behavior and establishment of the parasitoid Gonatocerus ashmeadi (Hymenoptera: Mymaridae) in Tahiti: Implications for its use as a biological control agent against Homalodisca vitripennis (Hemiptera: Cicadellidae)

The egg parasitoid Gonatocerus ashmeadi Girault (Hymenoptera: Mymaridae), was introduced into French Polynesia as a biological control agent to control the invasive plant feeding pest Homalodisca vitripennis (Germar) (Hemiptera: Cicadellidae). The short-distance dispersal of G. ashmeadi was monitored as part of the biological control program. G. ashmeadi showed exponential dispersal capacity with 47 m/day being a minimum estimate of its natural rate of spread at high host densities (>150 nymphs per minute of sweep net sampling) in urbanized areas at sea level, which were characterized by a high diversity of exotic ornamental plants. This rate of spread contrasted starkly with almost nonexistent establishment and dispersal where host densities were very low (<2 nymphs per minute of sweep net sampling) at high elevation (800 m) with relatively undisturbed native vegetation. Survey results across different altitudes revealed an effect of vegetative diversity and host density on the measurable mobility and establishment of G. ashmeadi. In contrast, no significant influence of wind direction was found on G. ashmeadi dispersal rate or direction. Survey results for G. ashmeadi from French Polynesia suggest that the best release establishment strategies for classical biological control of H. vitripennis are: (1) many small releases where host density is high, or (2) larger and fewer releases where host densities are low.

INVADING PARASITOIDS SUFFER NO ALLEE EFFECT: A MANIPULATIVE FIELD EXPERIMENT

One frequent explanation for the failure of biological invasions is the Allee effect: due to positive density dependence, initially small invading populations may fail to establish and spread. Populations released for biological control are similar to fortuitous invading populations and may therefore suffer from Allee effects. However, unlike fortuitous invasions, biological control allows the experimental manipulation of initial population size and, thus, offers a unique opportunity to test for the occurrence of Allee effects. We manipulated the initial size of 45 populations of a parasitoid wasp introduced for the biological control of a phytophagous insect and followed the population dynamics of both parasitoids and hosts during three years. Our results suggest an absence of Allee effects but clear negative density dependence instead: (1) the probability of establishment after three years was not affected by initial population size; (2) net reproductive rate was highest at low parasitoid density and high host density; (3) the sex ratio, reflecting the proportion of virgin females, did not increase at low density, suggesting that low densities did not impede mate-finding; (4) the depression of host populations did not depend upon the number of parasitoids introduced. This is, to our knowledge, the first experimental test of the Allee effect in an invading parasitoid. It leads us to propose that a number of behavioral and life-history features of many parasitoids could protect them from Allee effects.

A Simulation Model for Low-Pathogenicity Avian Influenza Viruses in Dabbling Ducks in Europe

Anseriformes are the reservoir of low-pathogenicity avian influenza viruses (LPAIV). Studies have shown a high LPAIV prevalence associated with low antibody detection in a wild duck population in northern European countries, whereas in winter areas (Mediterranean basin), low viral detection and high seroprevalence were observed. In order to gain insight into the role played by both population recruitment and migration on AIV persistence, an epidemiological model was developed. A susceptible, infectious and removed (immune or dead)-individuals model coupling population and infection dynamics was developed to simulate LPAIV circulation in dabbling ducks throughout the entire year. The transmission coefficient (beta) was calculated using the original dataset of published works, whereas dabbling duck demographic parameters were obtained from the literature. The estimated host density threshold for virus persistence is 380 susceptible individuals per day whereas the critical community size needed for maintaining the virus throughout the winter has been estimated to be about 1200 individuals. The model showed peaks of viral prevalence after nesting and during the moult period because of population recruitment and high host density, respectively. During the winter and spring periods, the viruses reach the minimal endemic level and local extinction is highly probable because of stochastic phenomena, respectively 80% and 90% of probabilities. The most sensitive parameters of the model are the recruitment rate of young susceptible animals and the duration of virus shedding.

A SIMULATION MODEL FOR THE LPAI VIRUSES IN DABBLING DUCKS IN EUROPE

Anseriformes are the reservoir of low pathogenic avian influenza viruses (LPAIV). Studies have shown a high LPAIV prevalence associated with low antibody detection in a wild duck population in northern European countries, whereas in winter areas (Mediterranean basin), low viral detection and high seroprevalence were observed. In order to gain insight into the role played by both population recruitment and migration on AIV persistence, an epidemiological model was developed. The model showed peaks of viral prevalence after nesting and during the moult period due to population recruitment and high host density, respectively. During the winter and spring periods, the viruses reach the minimal endemic level and local extinction is highly probable due to stochastic phenomena, respectively 80% and 90% of probabilities. The most sensitive parameters of the model are the recruitment rate of young susceptible animals and the duration of virus shedding.

Natural control ofDiuraphis noxiaandRhopalosiphum maidis(Aphidoidea) by parasitic entomophthorales (Zygomycota) in Slovakia

Seasonal development of entomophthoralean infection in colonies of Diuraphis noxia and Rhopalosiphum maidis was evaluated on summer barley under conditions of south-western Slovakia in 2001 and 2002. D. noxia was infesting summer barley during both years, while R. maidis was only recorded in 2002. Further three cereal aphids, Rhopalosiphum padi, Metopolophium dirhodum and Sitobion avenae, were also observed on summer barley. Complex of entomophthoralean fungi identified in cereal aphid colonies included four species, Pandora neoaphidis, Entomophthora planchoniana, Conidiobolus obscurus and Neozygites fresenii, with P. neoaphidis as the most prevalent fungus during both years. Two parasitic fungi, P. neoaphidis and E. planchoniana, were identified from D. noxia, but P. neoaphidis was predominant. Infection rate in D. noxia colonies was low, not exceeding 15% and 10% in 2001 and 2002, respectively. Only nine R. maidis individuals were killed with entomophthoralean fungi. The fungi were identified as P. neoaphidis and C. obscurus. No epizootics were observed in the aphid colonies. The fungi could not control development of the aphid populations since they did not appear until high host densities had been established in the crops.

Functional response and progeny production of the Madeira mealybug parasitoid, Anagyrus sp. nov. nr. sinope: The effect of host stage preference

Anagyrus sp. nov. nr. sinope Noyes & Menezes (Hymenoptera: Encyrtidae) is a koinobiont parasitoid with an oviposition preference for the third-instar and pre-reproductive adult female Madeira mealybug, Phenacoccus madeirensis Green (Hemiptera: Pseudococcidae). In this study, we report that host stage preference significantly influenced the parasitoid’s functional response and progeny production. The parasitoids exhibited a type II functional response when offered either second-instar or pre-reproductive adult female P. madeirensis at densities of 2–70 mealybugs per patch. The number of mealybugs parasitized within a 24-h period increased with host density up to the estimated maximum of 13 adults and 28 second-instar nymphs. The total number of progeny produced was higher from adult mealybugs and increased with host density until threshold densities (20 adults or 30 second-instar nymphs per patch) were reached. The maximum number of progeny produced in both host stages over a 24-h period was limited by the daily egg load, which was on average 23. Brood size was consistently higher in adult mealybugs and at higher host densities. The proportion of males decreased with an increase in host density and was higher when emerging from the adult mealybugs. Brood size and sex ratio of Anagyrus sp. nov. nr. sinope were similar at higher host densities, suggesting that there was a critical density after which the parasitoids only deposited a fixed brood size or sex ratio.

Infection dynamics of an oyster parasite in its newly expanded range

Over a 2-year period in 1990 and 1991, coincident with a pronounced warming episode, Dermo disease outbreaks in the oyster, Crassostrea virginica, caused by the parasite Perkinsus marinus, occurred over a 500-km range from Delaware Bay to Cape Cod, in the northeastern United States. The parasite had not previously been recorded or known to cause mortalities in this region. To document infection patterns and levels in this region several years after the initial outbreaks, and to compare them with those in the parasite’s historic southern range, we deployed and sampled oysters from 1996 to 1998 at multiple sites spanning the expanded range. During this 2-year period, the parasite was documented to occur in oysters at high prevalences throughout the new range, in sites varying from small, enclosed embayments to large estuaries, and in both cultured and wild-set oysters. Infection and mortality patterns, and levels were similar to those in southern locations where the parasite has been enzootic for at least decades. The persistence of high P. marinus infection levels in the new range after the initial expansion is probably due to several factors: (1) winter temperatures continued to increase during the 1990s and early 2000s, albeit at a slower rate than in 1990–1991, facilitating overwinter survival of the parasite; (2) many oyster-growing sites in the northeast are in relatively shallow water in which summer temperatures offer ample time for the parasite to proliferate and spread; and (3) the combination of high parasite burdens and high host densities in oyster farms results in an abundance of parasites and high transmission rates. Colder winters and high rainfall after 2002 reduced prevalences in some regions, but P. marinus can survive low temperatures and low salinities, and epizootic conditions are likely to return if temperatures rise again, as predicted by climate-change models.

Spatial patterns of host exploitation in a larval parasitoid of the predatory dusky large blue Maculinea nausithous

Summary The foraging behaviour of the parasitoid wasp Neotypus melanocephalus and factors affecting parasitism at the population level were studied. This specialised parasitoid attacks caterpillars of the butterfly Maculinea nausithous , which sequentially feed on the plant Sanguisorba officinalis and specific red Myrmica ants. Among M. nausithous populations, there is considerable variation in caterpillar densities. At low M. nausithous densities, foraging might be time consuming for N. melanocephalus. High host densities may not always be advantageous to foraging parasitoids due to the caterpillars’ frequent overexploitation of ant resources and subsequent density-dependent mortality. In order to disperse progeny, we hypothesised that N. melanocephalus should search in a non-random way at the level of the micro-habitat, i.e., single flower heads of S. officinalis . Our analysis of 32 natural populations in the Upper Rhine valley in Germany did not show a density-dependent relationship between M. nausithous caterpillars and parasitism. Furthermore, habitat parameters like patch size and density of the host's food plant did not affect the parasitism rate. Foraging N. melanocephalus females preferred to search on large flower heads. They probed host-occupied flower heads only, visiting non-host-exploited flower heads only briefly. Time spent on a flower head was independent of the number of caterpillars per flower head. This study indicates that N. melanocephalus increases its foraging efficiency by preferring large flower heads that were previously shown to contain more host caterpillars than small flower heads. Furthermore, oviposition increases the likelihood of continuing to search on a flower head, which is an adaptive strategy for parasitoids foraging for aggregated hosts. However, many host-occupied flower heads were not probed by N. melanocephalus . We discuss the possibility that temporal host refuges of M. nausithous caterpillars might contribute to heterogeneity of parasitism, and why spreading offspring might constitute a suitable strategy for a parasitoid of an ant-parasitic butterfly.

Low-level parasitic worm burdens may reduce body condition in free-ranging red deer (Cervus elaphus)

Regulation of ungulate populations by parasites relies on establishing a density-dependent relationship between infection and vital demographic rates which may act through the effect of parasites on body condition. We examine evidence for parasite impacts in 285 red deer (Cervus elaphus) harvested during 1991 and 1992 on the Isle of Rum. In the abomasa, prevalence of nematodes was 100% and the most abundant genus observed were Ostertagia species, however, mean intensity of infection was low (less than 1000) relative to other studies. Additional species, also present in low numbers, included Nematodirus spp., Capillaria spp., Cooperia spp., Monieza expanza, Oesophagostomum venulosum and Trichuris ovis. Lungworm (Dictyocaulus spp.) and tissue worm (Elaphostronygylus cervi) larvae were also observed in faecal samples. There was no evidence for acquired immunity to abomasal nematodes. Despite low levels of infection, both adult male and female deer showed significant negative correlation between indices of condition (kidney fat index, dressed carcass weight and larder weight) and intensity of Ostertagia spp. infection. However, there was no evidence that pregnancy rate in females was related to intensity of infection. For calves, there was no relationship between body condition and intensity of infection. The apparent subclinical effects of low-level parasite infection on red deer performance could alternatively be due to animals in poorer nutritional state being more susceptible to infection. Either way the results suggest that further studies of wild populations are justified, in particular where high local host densities exist or alternative ungulate hosts are present, and, where experimental treatments are tractable.

Importance of host feeding for parasitoids that attack honeydew‐producing hosts

AbstractInsect parasitoids lay their eggs in arthropods. Some parasitoid species not only use their arthropod host for oviposition but also for feeding. Host feeding provides nutrients to the adult female parasitoid. However, in many species, host feeding destroys an opportunity to oviposit. For parasitoids that attack Homoptera, honeydew is a nutrient‐rich alternative that can be directly imbibed from the host anus without injuring the host. A recent study showed that feeding on host‐derived honeydew can be an advantageous alternative in terms of egg quantity and longevity. Here we explore the conditions under which destructive host feeding can provide an advantage over feeding on honeydew. For 5 days, Encarsia formosa Gahan (Hymenoptera: Aphelinidae) parasitoids were allowed daily up to 3 h to oviposit until host feeding was attempted. Host feedings were either prevented or allowed and parasitoids had ad libitum access to honeydew between foraging bouts. Even in the presence of honeydew, parasitoids allowed to host feed laid more eggs per hour of foraging per host‐feeding attempt than parasitoids that were prevented from host feeding. The higher egg‐laying rate was not compromised by survival or by change in egg volume over time. In conclusion, host feeding can provide an advantage over feeding on honeydew. This applies most likely under conditions of high host density or low extrinsic mortality of adult parasitoids, when alternative food sources cannot supply enough nutrients to prevent egg limitation. We discuss how to integrate ecological and physiological studies on host‐feeding behavior

Phonotaxis of the female parasitoid Emblemasoma auditrix (Diptera, Sarcophagidae) in relation to number of larvae and age

The dipteran parasitoid Emblemasoma auditrix locates its host acoustically. Analysis showed that phonotactic female flies usually carry fully developed larvae within their uteri. The mean number of larvae per female at the beginning of the season was 37.9 (range from 10 to 50). The number of larvae decreased rapidly with increasing singing activity of the host cicada ( Okanagana rimosa). In high-density host populations the parasitoid is likely to become egg-limited. A possible selective phonotactic responsiveness depending on the number of larvae or the age of the female was tested with song models. Phonotaxis depended on both the temporal structure and the frequency content, but in the field no correlation was found between the number of larvae and the preferences for the acoustic signal. Experiments in the laboratory showed that flies without host contact broadened their phonotactic stimulus range with age.

Transmission dynamics of an iridescent virus in an experimental mosquito population: the role of host density

Abstract. 1. The transmission of insect pathogens cannot be adequately described by direct linear functions of host and pathogen density due to heterogeneity generated from behavioural or physiological traits, or from the spatial distribution of pathogen particles. Invertebrate iridescent viruses (IIVs) can cause patent and lethal infection or a covert sub‐lethal infection in insects.Aedes aegyptilarvae were exposed to suspensions of IIV type 6 at two densities. High larval density increased the prevalence of aggression resulting in potentially fatal wounding.2. The overall prevalence of infection (patent + covert) was positively influenced by host density and increased with exposure time in both densities. The survival time of patently infected insects was extended by ≈ 5 days compared with non‐infected insects.3. Maximum likelihood models based on the binomial distribution were fitted to empirical results. A model incorporating heterogeneity in host susceptibility by inclusion of a pathogen‐free refuge was a significantly better fit to data than an all‐susceptible model, indicating that transmission is non‐linear. The transmission coefficient (υ) did not differ with host density whereas the faction of the population that occupied the pathogen‐free refuge (ΠR) was significantly reduced at high host density compared with the low density treatment.4. The transmission of free‐living infective stages of an IIV inAe. aegyptilarvae is non‐linear, probably because of density‐related changes in the frequency of aggressive encounters between hosts. This alters host susceptibility to infection and effectively reduces the proportion of hosts that occupy the pathogen‐free refuge.

Spatial distributions of Cephus cinctus Norton (Hymenoptera: Cephidae) and its braconid parasitoids in Montana wheat fields

Bracon cephi and Bracon lissogaster are native parasitoids of the wheat stem sawfly, Cephus cinctus , an important pest of dryland wheat production. This spatial distribution study, using survey data from seven dryland wheat fields at four locations in north-central Montana over two years, examined: (1) the spatial distribution and spatial association of C. cinctus infestation and parasitism, and (2) parasitism in response to the level of C. cinctus infestation. Spatial distribution analyses revealed that C. cinctus infestations were significantly aggregated along field edges in three of five fields, while parasitism followed a random spatial distribution in all five fields. We demonstrate that the level of parasitism is highest for lower levels of C. cinctus infestation, in fields where parasitoids are either scarce or abundant. The combination of (1) a spatial aggregation of C. cinctus infestations along field edges, and (2) a potential negative interaction between parasitism and C. cinctus infestation may lower the level of parasitism at high host densities. This outcome can be viewed as beneficial to the insect pest, because parasitized C. cinctus larvae are at a greater risk of being cannibalized by conspecifics. An equally plausible hypothesis for this negative interaction is a limitation on the successful location of unparasitized hosts at higher host densities, which is also compelling because of the cryptic location of hosts within wheat stems that belie little visible evidence of infestation. Supporting data from other heavily infested fields that were sampled in 1999–2000 are used to illustrate the recurrence of this pattern. These findings are discussed in the context of developing redistribution strategies for introducing these parasitoids at sawfly infested locations where they are currently present at negligible levels.

The life history responses of the abalone pest, Terebrasabella heterouncinata, under natural and aquaculture conditions

The sabellid, Terebrasabella heterouncinata, is a small (<5 mm) intratubular brooder that lives in burrows within the host’s shell matrix. It is a semi-continuous breeder and despite producing small numbers of large eggs, infestation by this animal has reached epidemic proportions on local abalone farms. The present study compared the morphometrics and reproductive characteristics of worms from farmed and wild abalone, in the Walker Bay area of the south Western Cape Province of South Africa, to gain insights into why this animal has become so successful under aquaculture conditions. The farms designated farm A and farm B each had one “on-farm” site, and two wild sites, while farm C had two on-farm sites and two wild sites. The wild sites were natural abalone habitats located within 2.5 km of the farms. Our results conclusively showed that environmental conditions prevalent on the farms enhanced the reproductive success of these worms relative to that observed in its natural environment. At farms B and C, worms occurred in significantly higher densities at the on-farm sites than in the corresponding wild samples, but at farm A, density was equally low at the three sites. At all three farms, a greater proportion of the population was reproductively active in the on-farm samples than in the wild samples. Worms on farmed abalone had a higher instantaneous fecundity, brooded more clutches simultaneously and were larger than their conspecifics from the wild. There was a positive correlation between adult size and brood size and the number of clutches brooded simultaneously. Within the three on-farm sites there was a negative correlation between egg volume and brood size, indicating a trade-off between these traits. However, such a trade-off was not apparent between sites, with brood size being higher at the on-farm sites than at the wild sites, irrespective of egg size. This suggests that the stable nutrient-enriched environment on the farm led to an increase in fecundity without compromising the size (and implicitly the quality) of the eggs. Worm density did not have a significant effect on body size or any other reproductive traits at most sites, and the density of T. heterouncinata was unaffected by the density of other shell-infesting polychaetes. The results suggest that the farm environment has selected for larger, more fecund worms that breed rapidly with high recruitment success as a consequence of abundant nutrients, high host density, habitat stability and a possible lack of predation and interspecific competition.

Interspecific Relationships Between the Solitary Ectoparasitoid, Eupelmus vuilleti (Crw.) (Eupelmidae), and Its Sympatric Species, Dinarmus basalis (Rond.) (Pteromalidae), in the Presence of Their Host, Callosobruchus maculatus Pic (Coleoptera Bruchidae)

The eupelmid Eupelmus vuilleti CRW. and the pteromalid Dinarmus basalis Rond. are sympatric in West Africa. The reproduction of E. vuilleti and D. basalis females was analyzed in 10, 20, and 40 host patches when present alone or together. In 10- and 20-hosts patches, the presence of E. vuilleti affected the reproduction and offspring survival chances of D. basalis. In 40-hosts patch, only the offspring survival of D. basalis was affected by the presence of E. vuilleti. In contrast, the presence of D. basalis did not influence the reproduction of E. vuilleti and the survival chances of its offspring at all host densities tested. Multiparasitism did not occur at random. In a choice test, E. vuilleti showed a preference for hosts previously parasitized by D. basalis on healthy hosts. This attraction is mediated by chemical markers deposited by D. basalis females on the surface of the seed. At high host density, the likelihood of encountering a seed containing a host previously parasitized by D. basalis was lower for E. vuilleti, affecting moderately D. basalis reproductive success.

Role of Diadegma semiclausum (Hymenoptera: Ichneumonidae) in controlling Plutella xylostella (Lepidoptera: Plutellidae): Cage exclusion experiments and direct observation

We evaluated the role of the larval parasitoid, Diadegma semiclausum Hellén (Hymenoptera: Ichneumonidae), in controlling Plutella xylostella (L.) (Lepidoptera: Plutellidae) by cage exclusion experiments and direct field observation during the winter season in southern Queensland, Australia. The cage exclusion experiment involved uncaged, open cage and closed cage treatments. A higher percentage (54–83%) of P. xylostella larvae on sentinel plants were lost in the uncaged treatment than the closed (4–9%) or open cage treatments (11–29%). Of the larvae that remained in the uncaged treatment, 72–94% were parasitized by D. semiclausum, much higher than that in the open cage treatment (8–37% in first trial, and 38–63% in second trial). Direct observations showed a significant aggregation response of the field D. semiclausum populations to high host density plants in an experimental plot and to high host density plots that were artificially set-up near to the parasitoid source fields. The degree of aggregation varied in response to habitat quality of the parasitoid source field and scales of the manipulated host patches. As a result, density-dependence in the pattern of parasitism may depend on the relative degree of aggregation of the parasitoid population at a particular scale. A high degree of aggregation seems to be necessary to generate density-dependent parasitism by D. semiclausum. Integration of the cage exclusion experiment and direct observation demonstrated the active and dominant role of this parasitoid in controlling P. xylostella in the winter season. A biologically based IPM strategy, which incorporates the use of D. semiclausum with Bt, is suggested for the management of P. xylostella in seasons or regions with a mild temperature.

Experimental examination of bacteriophage latent-period evolution as a response to bacterial availability.

For obligately lytic bacteriophage (phage) a trade-off exists between fecundity (burst size) and latent period (a component of generation time). This trade-off occurs because release of phage progeny from infected bacteria coincides with destruction of the machinery necessary to produce more phage progeny. Here we employ phage mutants to explore issues of phage latent-period evolution as a function of the density of phage-susceptible bacteria. Theory suggests that higher bacterial densities should select for shorter phage latent periods. Consistently, we have found that higher host densities (>/== approximately 10(7) bacteria/ml) can enrich stocks of phage RB69 for variants that display shorter latent periods than the wild type. One such variant, dubbed sta5, displays a latent period that is approximately 70 to 80% of that of the wild type-which is nearly as short as the RB69 eclipse period-and which has a corresponding burst size that is approximately 30% of that of the wild type. We show that at higher host densities (>/== approximately 10(7) bacteria/ml) the sta5 phage can outcompete the RB69 wild type, though only under conditions of direct (same-culture) competition. We interpret this advantage as corresponding to slightly faster sta5 population growth, resulting in multifold increases in mutant frequency during same-culture growth. The sta5 advantage is lost, however, given indirect (different-culture) competition between the wild type and mutant or given same-culture competition but at lower densities of phage-susceptible bacteria (</= approximately 10(6) bacteria/ml). From these observations we suggest that phage displaying very short latent periods may be viewed as specialists for propagation when bacteria within cultures are highly prevalent and transmission between cultures is easily accomplished.

Habitat of pre-hibernating larvae of the endangered butterfly Euphydryas aurinia (Lepidoptera: Nymphalidae): What can be learned from vegetation composition and architecture?

Habitats of pre-hibernating gregarious larvae of the endangered Marsh Fritillary butterfly (Euphydryas aurinia) were studied in field in Western Bohemia, Czech Republic. The species inhabits moist seminatural meadows managed by light grazing and haymaking; the only local host plant is Succisa pratensis. The redundancy analyses of the vegetation composition (around 166 nest-bearing and 381 unoccupied host plants) showed that nest presence was positively associated with short cushion-forming grasses (esp. Nardus stricta). It was negatively associated with competitively superior tall grasses (e.g., Deschampsia caespitosa) and tall herbs. Comparison of Ellenberg's indicator values of vegetation in occupied vs. unoccupied plots revealed that the nests were more often found in drier, nitrogen-poorer and more acidic conditions than unoccupied plants. Multiple regressions of nest presence against architecture of the host plants (170 occupied, 1280 unoccupied) revealed that the nests were associated with densely clumped host plants, low to medium height of sward and mechanical disturbance. The patterns agree with our knowledge of the biology of the larvae: short sward (related to low nitrogen, humidity, and low pH) facilitates larval basking; high host density reduces the likelihood of starvation. Sod disturbance facilitates host plant germination. The conditions favourable for the nests were also favourable for the host plant, but the plants grew in broader range of conditions than that occupied by the nests. The abandoning of a site results into a situation when the conditions first become intolerable for the butterfly, and ultimately for the plant. Since grazing and mowing have a different impact on vegetation composition and architecture, we propose that the studied populations have persisted in a semi-dynamic state in the two modes of management. Conservation management should mimic the dynamics of traditional land use on the smaller scales of extant colonies.

Density-dependent foraging behaviors in a parasitoid lead to density-dependent parasitism of its host.

Empirical studies of spatial heterogeneity in parasitism by insect parasitoids have focused largely on patterns, while the many possible underlying mechanisms have been little studied in the field. We conducted experimental and observational studies on Tachinomyia similis (Diptera: Tachinidae) attacking western tussock moths (Orgyia vetusta; Lepidoptera: Lymantriidae) on lupine bushes at Bodega Bay, Calif., USA. We examined several foraging behaviors that have been hypothesized to create density-dependent variation in parasitism rates, including spatial aggregation of parasitoids to high host density, mutual interference among searching parasitoids and decelerating functional responses of the parasitoid. At the spatial scale of individual bushes, we detected both aggregation to a high density and a decelerating functional response. The resulting spatial pattern of parasitism was best fit by two models; one included an effect of parasitoid aggregation and the other included an effect of aggregation and a decelerating functional response. Most of the variation in parasitism was not correlated with density of O. vetusta.