Gypsy moths (Lymantria dispar) in the Niagara Region : population density variation, introduction of an entomopathogenic fungus (Entomophaga maimaiga) and occurrence of nuclear polyhedrosis virus

The gypsy moth, Lymantria dispar, a major defoliator of broad leaf trees, was accidentally introduced into North America in 1869. Much interest has been generated regarding the potential of using natural pathogens for biological control of this insect. One of these pathogens, a highly specific fungus, Entomophaga maimaiga, was accredited with causing major epizootics in populations of gypsy moth across the north-eastern United States in 1989 and 1990 and is thought to be spreading northwards into Canada. This study examined gypsy moth population densities in the Niagara Region. The fungus, .E.. maimaiga, was artificially introduced into one site and the resulting mortality in host populations was noted over two years. The relationship between fungal mortality, host population density and occurrence of another pathogen, the nuclear polyhedrosis virus (NPV), was assessed. Gypsy moth population density was assessed by counting egg masses in 0.01 hectare (ha) study plots in six areas, namely Louth, Queenston, Niagara-on-the-Lake, Shorthills Provincial Park, Chippawa Creek and Willoughby Marsh. High variability in density was seen among sites. Willoughby Marsh and Chippawa Creek, the sites with the greatest variability, were selected for more intensive study. The pathogenicity of E. maimaiga was established in laboratory trials. Fungal-infected gypsy moth larvae were then released into experimental plots of varying host density in Willoughby Marsh in 1992. These larvae served as the inoculum to infect field larvae. Other larvae were injected with culture medium only and released into control plots also of varying host density. Later, field larvae were collected and assessed for the presence of .E.. maimaiga and NPV. A greater proportion of larvae were infected from experimental plots than from control plots indicating that the experimental augmentation had been successful. There was no relationship between host density and the proportion of infected larvae in either experimental or control plots. In 1992, 86% of larvae were positive for NPV. Presence and intensity of NPV infection was independent of fungal presence, plot type or interaction of these two factors. Sampling was carried out in the summer of 1993, the year after the introduction, to evaluate the persistence of the pathogen in the environment. Almost 50% of all larvae were infected with the fungus. There was no difference between control and experimental plots. Data collected from Willoughby Marsh indicated that there was no correlation between the proportion of larvae infected with the fungus and host population density in either experimental or control plots. About 10% of larvae collected from a nearby site, Chippawa Creek, were also positive for .E.. maimaiga suggesting that low levels of .E.. maimaiga probably occurred naturally in the area. In 1993, 9.6% of larvae were positive for NPV. Again, presence or absence of NPV infection was independent of fungal presence plot type or interaction of these two factors. In conclusion, gypsy moth population densities were highly variable between and within sites in the Niagara Region. The introduction of the pathogenic fungus, .E.. maimaiga, into Willoughby Marsh in 1992 was successful and the fungus was again evident in 1993. There was no evidence for existence of a relationship between fungal mortality and gypsy moth density or occurrence of NPV. The results from this study are discussed with respect to the use of .E.. maimaiga in gypsy moth management programs.

Aggregation of parasitism risk in an aphid-parasitoid system: Effects of plant patch size and aphid density

Few studies have linked density dependence of parasitism and the tritrophic environment within which a parasitoid forages. In the non-crop plant-aphid, Centaurea nigra-Uroleucon jaceae system, mixed patterns of density-dependent parasitism by the parasitoids Aphidius funebris and Trioxys centaureae were observed in a survey of a natural population. Breakdown of density-dependent parasitism revealed that density dependence was inverse in smaller colonies but direct in large colonies (>20 aphids), suggesting there is a threshold effect in parasitoid response to aphid density. The CV2 of searching parasitoids was estimated from parasitism data using a hierarchical generalized linear model, and CV2>1 for A. funebris between plant patches, while for T. centaureae CV2>1 within plant patches. In both cases, density independent heterogeneity was more important than density-dependent heterogeneity in parasitism. Parasitism by T. centaureae increased with increasing plant patch size. Manipulation of aphid colony size and plant patch size revealed that parasitism by A. funebris was directly density dependent at the range of colony sizes tested (50-200 initial aphids), and had a strong positive relationship with plant patch size. The effects of plant patch size detected for both species indicate that the tritrophic environment provides a source of host density independent heterogeneity in parasitism, and can modify density-dependent responses. (c) 2007 Gessellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

Quality of Cotesia flavipes (Hymenoptera: Braconidae) reared at different host densities and the estimated cost of its commercial production

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)

Outbreaks of pink wax scale, Ceroplastes rubens Maskell (Hemiptera : Coccidae), on umbrella trees in south-eastern Queensland: Patterns of parasitisation

The degree and distribution of parasitisation in relation to densities of pink wax scale, Ceroplastes rubens Maskell, on umbrella trees, Schefflera actinophylla (Endl.), in south-eastern Queensland were investigated to determine whether scale outbreaks could be attributed, in part, to low levels of parasitisation. Rates of parasitisation were independent of or inversely dependent on host density, and highly variable, especially at low densities. The absence of density dependent parasitisation may occur as a result of: (i) non-aggregation by parasitoids; (ii) aggregation by parasitoids where parasitisation is limited by intrinsic or extrinsic factors; and/or (iii) high rates of hyperparasitisation.

The effect of resource aggregation at different scales: Optimal foraging behavior of Cotesia rubecula

Resources can be aggregated both within and between patches. In this article, we examine how aggregation at these different scales influences the behavior and performance of foragers. We developed an optimal foraging model of the foraging behavior of the parasitoid wasp Cotesia rubecula parasitizing the larvae of the cabbage butterfly Pieris rapae. The optimal behavior was found using stochastic dynamic programming. The most interesting and novel result is that the effect of resource aggregation within and between patches depends on the degree of aggregation both within and between patches as well as on the local host density in the occupied patch, but lifetime reproductive success depends only on aggregation within patches. Our findings have profound implications for the way in which we measure heterogeneity at different scales and model the response of organisms to spatial heterogeneity.

Optimal patch-leaving behaviour: a case study using the parasitoid Cotesia rubecular

1. Parasitoids are predicted to spend longer in patches with more hosts, but previous work on Cotesia rubecula (Marshall) has not upheld this prediction, Tests of theoretical predictions may be affected by the definition of patch leaving behaviour, which is often ambiguous. 2. In this study whole plants were considered as patches and assumed that wasps move within patches by means of walking or flying. Within-patch and between-patch flights were distinguished based on flight distance. The quality of this classification was tested statistically by examination of log-survivor curves of flight times. 3. Wasps remained longer in patches with higher host densities, which is consistent with predictions of the marginal value theorem (Charnov 1976). tinder the assumption that each flight indicates a patch departure, there is no relationship between host density and leaving tendency. 4. Oviposition influences the patch leaving behaviour of wasps in a count down fashion (Driessen et al. 1995), as predicted by an optimal foraging model (Tenhumberg, Keller & Possingham 2001). 5. Wasps spend significantly longer in the first patch encountered following release, resulting in an increased rate of superparasitism.

The population dynamics of disease on short and long time-scales

Observational evidence is scarce concerning the distribution of plant pathogen population sizes or densities as a function of time-scale or spatial scale. For wild pathosystems we can only get indirect evidence from evolutionary patterns and the consequences of biological invasions.We have little or no evidence bearing on extermination of hosts by pathogens, or successful escape of a host from a pathogen. Evidence over the last couple of centuries from crops suggest that the abundance of particular pathogens in the spectrum affecting a given host can vary hugely on decadal timescales. However, this may be an artefact of domestication and intensive cultivation. Host-pathogen dynamics can be formulated mathematically fairly easily–for example as SIR-type differential equation or difference equation models, and this has been the (successful) focus of recent work in crops. “Long-term” is then discussed in terms of the time taken to relax from a perturbation to the asymptotic state. However, both host and pathogen dynamics are driven by environmental factors as well as their mutual interactions, and both host and pathogen co-evolve, and evolve in response to external factors. We have virtually no information about the importance and natural role of higher trophic levels (hyperpathogens) and competitors, but they could also induce long-scale fluctuations in the abundance of pathogens on particular hosts. In wild pathosystems the host distribution cannot be modelled as either a uniform density or even a uniform distribution of fields (which could then be treated as individuals). Patterns of short term density-dependence and the detail of host distribution are therefore critical to long-term dynamics. Host density distributions are not usually scale-free, but are rarely uniform or clearly structured on a single scale. In a (multiply structured) metapopulation with coevolution and external disturbances it could well be the case that the time required to attain equilibrium (if it exists) based on conditions stable over a specified time-scale is longer than that time-scale. Alternatively, local equilibria may be reached fairly rapidly following perturbations but the meta-population equilibrium be attained very slowly. In either case, meta-stability on various time-scales is a more relevant than equilibrium concepts in explaining observed patterns.

Effects of sex and locality on the abundance of lice on the wild rodent Oligoryzomys nigripes

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

Spatial pattern of parasitism in Diatraea saccharalis Fab. (Lep., Crambidae) populations at two different spatial scales in sugarcane fields in Brazil

The biological control of Diatraea saccharalis is regarded as one of the best examples of successful classical biological control in Brazil. Since the introduction of the exotic parasitoid, Cotesia flavipes, the decrease in D. saccharalis infestation in sugarcane fields has been attributed to the effectiveness of this agent. Native Tachinidae fly parasitoids (Lydella minense and Paratheresia claripalpis) have also been implicated in the success. Quantitative data confirming the actual contribution of these agents to the control of D. saccharalis are, however, rather scant. The purpose of this study was to investigate the spatial pattern of parasitism of these parasitoids in D. saccharalis populations at two large spatial scales (fields and zones). To investigate this subject, a large data set comprising information collected from a sugarcane mill located in the state of São Paulo, Brazil (São João sugarcane mill) was analysed. When regressions between the proportion parasitism against host density were computed, the percentage of significant regressions with either a positive or a negative slope was very small at both spatial scales for both parasitoid species. Regressing the densities of tachinid-parasitized hosts against host densities per field showed that these parasitoids presented a 'moderate aggregative' response to host densities, as 53.33% of the regressions were positively significant. Cotesia flavipes was 'weakly aggregated' on host densities at the field level, because only 33.33% of the regressions were positively significant. At the zone level, neither aggregative nor spatial proportion parasitism responses were evident for either parasitoid species due to the small percentage of significant regressions computed.

Valor adaptativo do parasitóide Apanteles galleriae (Hymenoptera: Braconidae) de população natural e da população mantida em laboratório

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES)

An Ecological Study of Helminths of Some Wyoming Voles (Microtus spp.) with a Description of a New Species of Nematospiroides (Heligmosomidae: Nematoda)

An ecological and taxonomic study of the helminth parasites of voles (Microtus spp.) in the Jackson Hole region of Wyoming is reported. Nematospiroides microti n. sp. from Microtus montanus nanus and M. richardsoni macropus is described and figured. A cestode, Paranoplocephala infrequens, and a nematode, Syphacia obvelata, were generally distributed throughout the region in all habitats except the sage flats. A trematode, Quinqueserialis hassalli, was recovered only from voles collected near streams at low altitudes. This was presumably due to the localized distribution of the molluscan intermediate host. Four helminths, viz., Hymenolepis horrida, Heligmosomum costellatum, Nematospiroides microti and Trichuris opaca, were restricted in their distribution to the alpine and sub-alpine meadows. Of these parasites, H. horrida and H. costellatum are reported for the first time from North America. Most of the other host and locality records are new. Available data indicate that host specificity was not a factor in restricting the distribution of parasites. Although the greatest numbers of parasites, both qualitative and quantitative, occurred in habitats where host density was greatest, it seems unlikely that host density is the only factor involved.

Populationsdynamik des Cydia pomonella Granulovirus

Das Cydia pomonella Granulovirus (CpGV, Fam. Baculoviridae) ist ein sehr virulentes und hoch spezifisches Pathogen des Apfelwicklers (Cydia pomonella), das seit mehreren Jahren in der Bundesrepublik Deutschland und anderen Ländern der EU als Insektizid zugelassen ist. Wie andere Baculoviren auch befällt es die Larven der Insekten und ist aufgrund seiner Selektivität für Nicht-Zielorganismen unbedenklich. In der Vergangenheit konzentrierte sich die Erforschung des CpGV auf Bereiche, die für die Anwendung im Pflanzenschutz relevant waren, wobei nach fast 20 Jahren nach der ersten Zulassung noch immer nicht bekannt ist, ob und wie sich das CpGV in der Umwelt etablieren kann. Im Rahmen der vorliegenden Arbeit wurden verschiedene Parameter, mit deren Hilfe die Populationsdynamik des CpGV beschrieben werden kann, analysiert und quantitativ bestimmt. Neben den biologischen Eigenschaften wie Virulenz, DNA-Charakterisierung und Quantifizierung der Virusnachkommenschaft wurden insbesondere die horizontale sowie die vertikale Transmission, die Inaktivierung und die Infektion später Larvenstadien untersucht. Letztlich wurden die ermittelten Parameter zusammen mit Daten aus der Literatur in ein mathematisches Modell integriert. Um die Wahrscheinlichkeit der horizontalen Transmission zu quantifizieren, wurde ein Modellsystem mit losen Äpfeln etabliert, in dem verschiedene Szenarien möglicher horizontaler Transmission unter definierten Laborbedingungen getestet wurden. In Versuchsserien, in denen ein Virusfleck, entsprechend der produzierten Virusmenge einer Eilarve, auf einen Apfel appliziert worden war, war unter den aufgesetzten Apfelwicklerlarven lediglich eine sehr geringe Mortalität von 3 - 6% zu beobachten. Wurde jedoch ein an einer Virusinfektion gestorbener Larvenkadaver als Inokulum verwendet, lag die Mortalitätsrate aufgesetzter Larven bei über 40%. Diese beobachtete hohe horizontale Transmissionsrate konnte mit dem Verhalten der Larven erklärt werden. Die Larven zeigten eine deutliche Einbohrpräferenz für den Stielansatz bzw. den Kelch, wodurch die Wahrscheinlichkeit des Zusammentreffens einer an der Infektion verendeten Larve mit einer gesunden Larve um ein Vielfaches zunahm. In einem ähnlich angelegten Freilandversuch konnte eine horizontale Transmission nicht belegt werden. Der Unterschied zur Kontrollgruppe fiel aufgrund einer hohen natürlichen Mortalität und einer damit einhergehenden niedrigen Dichte der Larven zu gering aus. Parallel hierzu wurde außerdem eine Halbwertszeit von 52 Sonnenstunden für das CpGV ermittelt. Weiterhin konnte festgestellt werden, dass die Mortalität von späteren Larvenstadien, die 14 Tage Zeit hatten sich in die Äpfel einzubohren, bevor eine CpGV-Applikation durchgeführt wurde, ebenso hoch war wie bei Larven, die sich im L1-Stadium auf der Apfeloberfläche infizierten. Aufgrund des höheren Alters jener Larven war der Fraßschaden an befallenen Äpfeln jedoch wesentlich größer und vergleichbar mit dem Fraßschaden einer unbehandelten Kontrolle. Der Versuch zur vertikalen Transmission zeigte dass, obwohl die verwendete Apfelwicklerzucht nicht frei von CpGV war, die Mortalitätsrate der Nachkommen subletal infizierter Weibchen (44%) jedoch deutlich höher war als die der Nachkommen subletal infizierter Männchen (28%) und der unbehandelten Kontrolle (27%). Auch in den PCR-Analysen konnte eine größere Menge an CpGV-Trägern bei den Nachkommen subletal infizierter Weibchen (67%) als bei den Nachkommen subletal infizierter Männchen (49%) und bei der Kontrolle (42%) nachgewiesen werden. Die Ergebnisse deuten darauf hin, dass eine Infektion durch subletal infizierte Weibchen vertikal in die nächste Generation übertragen werden kann. Dies lässt erkennen, dass in der Folgegeneration des Apfelwicklers eine zusätzliche Wirkung des CpGV durch vertikale Transmission auftreten kann. Hierin wäre auch ein potentieller Mechanismus für eine dauerhafte Etablierung des Virus zu sehen. Letztlich wurden alle Parameter, die die CpGV-Apfelwickler-Beziehung beschreiben, in ein mathematisches Modell GRANULO integriert. Nach einer Sensitivitätsanalyse wurde GRANULO teilweise mit Daten aus den Freilandversuchen verifiziert. Durch Modifikation der Virusparameter im Modell konnte anschließend der Einfluss veränderter biologischer Eigenschaften (UV-Stabilität und Transmissionsraten) der Viren in Simulationen theoretisch erprobt werden. Das beschriebene Modell, das allerdings noch einer weitergehenden Verifizierung und Validierung bedarf, ist eine erste Annäherung an die quantitative Erfassung und Modellierung der Populationsdynamik des Systems CpGV-Apfelwickler. Die im Zusammenhang mit der Populationsdynamik des Apfelwicklers erhobenen Daten können einen wertvollen Beitrag zur Optimierung von Kontrollstrategien des Apfelwicklers mittels CpGV leisten. Außerdem geben sie Aufschluss über die Etablierungsmöglichkeiten dieses Bioinsektizids.

Epidemiologia delle strongilosi dell'asino: quali applicazioni per il controllo delle infezioni da elminti?

Strongylosis in equids, despite being very common, have never been studied from a strictly ecological point of view. Mathematical models are important ecological tools used to study the temporal dynamics of parasite populations, and are useful to study the effect of different biological parameters, as well as to analyse the outcome produced by perturbations such as anthelmintic treatments. This work describes the study of the temporal dynamics of strongyles infection in an organic donkey population, performed using coprological quantitative analysis and donkeys’ age as a proxy of the time of infection. Force of infection was then estimated for Strongylus vulgaris and small strongyles and the results used as the basis for the development of mathematical models. In particular, the comparison of models output and field data made it possible to estimate the transmission coefficient  and to consequently calculate the basic reproduction number R0 and the threshold host density. Small strongyles model includes hypobiosis and, more interestingly as never found in literature, a density-dependent development rate of hypobiotic larvae in adult parasites in order to simulate a negative feedback between larvae emergence from hypobiosis and adult parasite abundance. Simulations of pharmacological and environmental treatments showed that parasite eradication was possible for S. vulgaris only, while small strongyles, due to hypobiosis and density-dependent development rate of their hypobiotic larvae, are very difficult to control and impossible to eradicate. In addition, density-dependence in larval development has been demonstrated to act as a key factor in improving parasite population survival and abundance even in absence of human intervention.

Effects of Habitat Fragmentation on Abundance, Larval Food and Parasitism of a Spider-Hunting Wasp.

Habitat fragmentation strongly affects species distribution and abundance. However, mechanisms underlying fragmentation effects often remain unresolved. Potential mechanisms are (1) reduced dispersal of a species or (2) altered species interactions in fragmented landscapes. We studied if abundance of the spider-hunting and cavity-nesting wasp Trypoxylon figulus Linnaeus (Hymenoptera: Crabronidae) is affected by fragmentation, and then tested for any effect of larval food (bottom up regulation) and parasitism (top down regulation). Trap nests of T. figulus were studied in 30 agricultural landscapes of the Swiss Plateau. The sites varied in the level of isolation from forest (adjacent, in the open landscape but connected, isolated) and in the amount of woody habitat (from 4 % to 74 %). We recorded wasp abundance (number of occupied reed tubes), determined parasitism of brood cells and analysed the diversity and abundance of spiders that were deposited as larval food. Abundances of T. figulus were negatively related to forest cover in the landscape. In addition, T. figulus abundances were highest at forest edges, reduced by 33.1% in connected sites and by 79.4% in isolated sites. The mean number of spiders per brood cell was lowest in isolated sites. Nevertheless, structural equation modelling revealed that this did not directly determine wasp abundance. Parasitism was neither related to the amount of woody habitat nor to isolation and did not change with host density. Therefore, our study showed that the abundance of T. figulus cannot be fully explained by the studied trophic interactions. Further factors, such as dispersal and habitat preference, seem to play a role in the population dynamics of this widespread secondary carnivore in agricultural landscapes.

Higher plant diversity promotes higher diversity of fungal pathogens, while it decreases pathogen infection per plant

Fungal plant pathogens are common in natural communities where they affect plant physiology, plant survival, and biomass production. Conversely, pathogen transmission and infection may be regulated by plant community characteristics such as plant species diversity and functional composition that favor pathogen diversity through increases in host diversity while simultaneously reducing pathogen infection via increased variability in host density and spatial heterogeneity. Therefore, a comprehensive understanding of multi-host multi-pathogen interactions is of high significance in the context of biodiversity-ecosystem functioning. We investigated the relationship between plant diversity and aboveground obligate parasitic fungal pathogen (''pathogens'' hereafter) diversity and infection in grasslands of a long-term, large-scale, biodiversity experiment with varying plant species (1-60 species) and plant functional group diversity (1-4 groups). To estimate pathogen infection of the plant communities, we visually assessed pathogen-group presence (i.e., rusts, powdery mildews, downy mildews, smuts, and leaf-spot diseases) and overall infection levels (combining incidence and severity of each pathogen group) in 82 experimental plots on all aboveground organs of all plant species per plot during four surveys in 2006. Pathogen diversity, assessed as the cumulative number of pathogen groups on all plant species per plot, increased log-linearly with plant species diversity. However, pathogen incidence and severity, and hence overall infection, decreased with increasing plant species diversity. In addition, co-infection of plant individuals by two or more pathogen groups was less likely with increasing plant community diversity. We conclude that plant community diversity promotes pathogen-community diversity while at the same time reducing pathogen infection levels of plant individuals.