Dynamics of a host-parasitoid interaction clarified by modelling and DNA sequencing.

It has been hypothesised that the 2-year oscillations in abundance of Xestia moths are mediated by interactions with 1-year Ophion parasitoid wasps. We tested this hypothesis by modelling a 35-year time series of Xestia and Ophion from Northern Finland. Additionally, we used DNA barcoding to ascertain the species diversity of Ophion and targeted amplicon sequencing of their gut contents to confirm their larval hosts. Modelling of the time-series data strongly supported the hypothesised host-parasitoid dynamics and that periodic occurrence of Xestia moths is mediated by Ophion. DNA barcodes revealed that Ophion included five species rather than just one while targeted amplicon sequencing verified that Ophion does parasitise Xestia. At least one Ophion species employs 1-year Syngrapha interrogationis as an alternate host, but it did not detectably affect Xestia-Ophion dynamics. We also demonstrate the previously unrecognised complexity of this system due to cryptic parasitoid diversity.

A review of Tersilochinae (Hymenoptera: Ichneumonidae) of Finland. Part 1: taxonomy.

The Tersilochinae fauna of Finland is critically revised (except subgenera Tersilochus Holmgren and Euporizon Horstmann). Tersilochinae in two principal Finnish ichneumonid collections, the Finnish Natural History Museum LUOMUS (University of Helsinki) and the private collection of R. Jussila (Turku), have been examined. Thirteen genera and 61 species are found to occur in Finland, including one new species, Barycnemis finnora Khalaim, sp. nov. The Nearctic genus Ctenophion Horstmann is recorded from Finland, as well as from the Palaearctic region, for the first time. Nineteen species are newly recorded from Finland and four species are excluded from the Finnish fauna. Barycnemis agilis (Holmgren) is transferred to the genus Spinolochus Horstmann (comb. nov.). Tersilochus sulcatus Hellén, 1958, a junior homonym of T. sulcatus Smith van Burgst, 1913, is synonymised with Phradis brevicornis Horstmann (syn. nov.). New data on distribution of 25 species in northwestern parts of Russia are also provided.

An annotated and illustrated checklist of Microgastrinae wasps (Hymenoptera, Braconidae) from the Canadian Arctic Archipelago and Greenland.

The Microgastrinae (Hymenoptera: Braconidae) from ten islands of the Canadian Arctic Archipelago (CAA) and Greenland were studied based on 2,183 specimens deposited in collections. We report a total of 33 species in six genera, more than doubling the totals previously known. Most of the species (75.7%) have a distribution restricted to the Nearctic, with nine of those (27.3%) confirmed to be High Arctic endemics and another 10 species considered very likely to be High Arctic endemics as well - accounting for all of those, more than half of all species found are endemic to the region. The most diverse genera were Cotesia (10 species), Glyptapanteles (9 species), and Microplitis (7 species), representing 78.8% of the overall species diversity in the region. The six most frequently collected species comprised 84.7% of all examined specimens. The flight period for Microgastrinae in the High Arctic encompasses only two months, with activity peaking during the first half of July, when almost 40% of all available specimens were collected, and then plummeting in the first half to the end of August. Microgastrinae wasps from the High Arctic are currently known to parasitize eight species within four families of Lepidoptera: three species of Noctuidae, two each of Lymantridae and Nymphalidae, and one species of Pterophoridae. However, that information is very preliminary, as only six of the 33 species of microgastrines currently have associated host data. An annotated checklist, including photographs for 24 of the 33 species, is provided, as well as a key to all Microgastrinae genera present in the region.

Discovery of long-distance gamete dispersal in a lichen-forming ascomycete.

Accurate estimates of gamete and offspring dispersal range are required for the understanding and prediction of spatial population dynamics and species persistence. Little is known about gamete dispersal in fungi, especially in lichen-forming ascomycetes. Here, we estimate the dispersal functions of clonal propagules, gametes and ascospores of the epiphytic lichen Lobaria pulmonaria. We use hierarchical Bayesian parentage analysis, which integrates genetic and ecological information from multiannual colonization and dispersal source data collected in a large, old-growth forest landscape. The effective dispersal range of gametes is several hundred metres to kilometres from potential paternal individuals. By contrast, clonal propagules disperse only tens of metres, and ascospores disperse over several thousand metres. Our study reveals the dispersal distances of individual reproductive units; clonal propagules, gametes and ascospores, which is of great importance for a thorough understanding of the spatial dynamics of ascomycetes. Sexual reproduction occurs between distant individuals. However, whereas gametes and ascospores disperse over long distances, the overall rate of colonization of trees is low. Hence, establishment is the limiting factor for the colonization of new host trees by the lichen in old-growth landscapes.

Local epiphyte establishment and future metapopulation dynamics in landscapes with different spatiotemporal properties.

Understanding the relative importance of different ecological processes on the metapopulation dynamics of species is the basis for accurately forecasting metapopulation size in fragmented landscapes. Successful local colonization depends on both species dispersal range and how local habitat conditions affect establishment success. Moreover, there is limited understanding of the effects of different spatiotemporal landscape properties on future metapopulation size. We investigate which factors drive the future metapopulation size of the epiphytic model lichen species Lobaria pulmonaria in a managed forest landscape. First, we test the importance of dispersal and local conditions on the colonization-extinction dynamics of the species using Bayesian state-space modelling of a large-scale data set collected over a 10-yr period. Second, we test the importance of dispersal and establishment limitation in explaining establishment probability and subsequent local population growth, based on a 10-yr propagule sowing experiment. Third, we test how future metapopulation size is affected by different metapopulation and spatiotemporal landscape dynamics, using simulations with the metapopulation models fitted to the empirical data. The colonization probability increased with tree inclination and connectivity, with a mean dispersal distance of 97 m (95% credible intervals, 5-530 m). Local extinctions were mainly deterministic set by tree mortality, but also by tree cutting by forestry. No experimental establishments took place on clearcuts, and in closed forest the establishment probability was higher on trees growing on moist than on dry-mesic soils. The subsequent local population growth rate increased with increasing bark roughness. The simulations showed that the restricted dispersal range estimated (compared to non-restricted dispersal range), and short tree rotation length (65 yr instead of 120) had approximately the same negative effects on future metapopulation size, while regeneration of trees creating a random tree pattern instead of an aggregated one had only some negative effect. However, using the colonization rate obtained with the experimentally added diaspores led to a considerable increase in metapopulation size, making the dispersal limitation of the species clear. The future metapopulation size is thus set by the number of host trees located in shady conditions, not isolated from occupied trees, and by the rotation length of these host trees.

Determinants of parasitoid communities of willow-galling sawflies: habitat overrides physiology, host plant and space.

Studies on the determinants of plant-herbivore and herbivore-parasitoid associations provide important insights into the origin and maintenance of global and local species richness. If parasitoids are specialists on herbivore niches rather than on herbivore taxa, then alternating escape of herbivores into novel niches and delayed resource tracking by parasitoids could fuel diversification at both trophic levels. We used DNA barcoding to identify parasitoids that attack larvae of seven Pontania sawfly species that induce leaf galls on eight willow species growing in subarctic and arctic-alpine habitats in three geographic locations in northern Fennoscandia, and then applied distance- and model-based multivariate analyses and phylogenetic regression methods to evaluate the hierarchical importance of location, phylogeny and different galler niche dimensions on parasitoid host use. We found statistically significant variation in parasitoid communities across geographic locations and willow host species, but the differences were mainly quantitative due to extensive sharing of enemies among gallers within habitat types. By contrast, the divide between habitats defined two qualitatively different network compartments, because many common parasitoids exhibited strong habitat preference. Galler and parasitoid phylogenies did not explain associations, because distantly related arctic-alpine gallers were attacked by a species-poor enemy community dominated by two parasitoid species that most likely have independently tracked the gallers' evolutionary shifts into the novel habitat. Our results indicate that barcode- and phylogeny-based analyses of food webs that span forested vs. tundra or grassland environments could improve our understanding of vertical diversification effects in complex plant-herbivore-parasitoid networks.

Complementary molecular information changes our perception of food web structure.

How networks of ecological interactions are structured has a major impact on their functioning. However, accurately resolving both the nodes of the webs and the links between them is fraught with difficulties. We ask whether the new resolution conferred by molecular information changes perceptions of network structure. To probe a network of antagonistic interactions in the High Arctic, we use two complementary sources of molecular data: parasitoid DNA sequenced from the tissues of their hosts and host DNA sequenced from the gut of adult parasitoids. The information added by molecular analysis radically changes the properties of interaction structure. Overall, three times as many interaction types were revealed by combining molecular information from parasitoids and hosts with rearing data, versus rearing data alone. At the species level, our results alter the perceived host specificity of parasitoids, the parasitoid load of host species, and the web-wide role of predators with a cryptic lifestyle. As the northernmost network of host-parasitoid interactions quantified, our data point exerts high leverage on global comparisons of food web structure. However, how we view its structure will depend on what information we use: compared with variation among networks quantified at other sites, the properties of our web vary as much or much more depending on the techniques used to reconstruct it. We thus urge ecologists to combine multiple pieces of evidence in assessing the structure of interaction webs, and suggest that current perceptions of interaction structure may be strongly affected by the methods used to construct them.

Indirect interactions in the High Arctic.

Indirect interactions as mediated by higher and lower trophic levels have been advanced as key forces structuring herbivorous arthropod communities around the globe. Here, we present a first quantification of the interaction structure of a herbivore-centered food web from the High Arctic. Targeting the Lepidoptera of Northeast Greenland, we introduce generalized overlap indices as a novel tool for comparing different types of indirect interactions. First, we quantify the scope for top-down-up interactions as the probability that a herbivore attacking plant species i itself fed as a larva on species j. Second, we gauge this herbivore overlap against the potential for bottom-up-down interactions, quantified as the probability that a parasitoid attacking herbivore species i itself developed as a larva on species j. Third, we assess the impact of interactions with other food web modules, by extending the core web around the key herbivore Sympistis nigrita to other predator guilds (birds and spiders). We find the host specificity of both herbivores and parasitoids to be variable, with broad generalists occurring in both trophic layers. Indirect links through shared resources and through shared natural enemies both emerge as forces with a potential for shaping the herbivore community. The structure of the host-parasitoid submodule of the food web suggests scope for classic apparent competition. Yet, based on predation experiments, we estimate that birds kill as many (8%) larvae of S. nigrita as do parasitoids (8%), and that spiders kill many more (38%). Interactions between these predator guilds may result in further complexities. Our results caution against broad generalizations from studies of limited food web modules, and show the potential for interactions within and between guilds of extended webs. They also add a data point from the northernmost insect communities on Earth, and describe the baseline structure of a food web facing imminent climate change.

Geographical variation in host use of a blood-feeding ectoparasitic fly: implications for population invasiveness.

Invasive generalist ectoparasites provide a tool to study factors affecting expansion rates. An increase in the number of host species may facilitate geographic range expansion by increasing the number of suitable habitats and by affecting local extinction and colonization rates. A geographic perspective on parasite host specificity and its implications on range expansion are, however, insufficiently understood. We conducted a field study to explore if divergent host specificity could explain the observed variation in expansion rates between Fennoscandian populations of the deer ked (Lipoptena cervi), which is a blood-feeding ectoparasitic fly of cervids. We found that the rapidly expanding eastern population in Finland appears to specialize on moose, whereas the slowly expanding western population in Norway breeds successfully on both moose and roe deer. The eastern population was also found to utilize the wild forest reindeer as an auxiliary host, but this species is apparently of low value for L. cervi in terms of adult maintenance, reproductive output and offspring quality. Abundant numbers of roe deer and white-tailed deer were observed to be apparently uninfected in Finland, suggesting that host use is not a plastic response to host availability, but rather a consequence of population-level evolutionary changes. Locally compatible hosts were found to be the ones sharing a long history with the deer ked in the area. Cervids that sustained adult deer keds also allowed successful reproduction. Thus, host use is probably determined by the ability of the adult to exploit particular host species. We conclude that a wide host range alone does not account for the high expansion rate or wide geographic distribution of the deer ked, although loose ecological requirements would increase habitat availability.

Use of forest corridors by boreal Xestia moths.

We studied movements of the noctuid moths Xestia speciosa and X. fennica in a fragmented forest landscape including old-growth spruce forest corridors in eastern Finland. We individually marked and released 1,796 adults of the habitat generalist X. speciosa and 683 individuals of X. fennica, which prefers old-growth spruce forests. Moths were recaptured with 60 sugar-bait traps that were placed within habitat corridors, in the non-habitat matrix and in surrounding old-growth forest patches. Both species favoured the corridors during their movements and tended to avoid entering the matrix. We analysed recapture rates and movements of the recaptured moths. Results suggest that X. speciosa moves longer distances than X. fennica. Though corridors clearly canalized movements by both species, the impact of old-growth forest corridors at the larger landscape scale was less clear in X. fennica. Movement behaviour of the two sexes differed more than that of the two species.

Genetic differentiation between alternate-year cohorts of Xestia tecta (Lepidoptera, Noctuidae) in Finnish Lapland.

Several species in the noctuid genus Xestia exhibit periodic dynamics, with two coexisting cohorts flying in alternate years. The population dynamics and two-year life cycle of Xestia moths suggest that the two cohorts are more or less isolated in time. Typically one cohort is abundant and the other one is rare. Knowledge of the extent of isolation between the two cohorts is important to fully understand the population dynamics and the evolution of alternate-year flight in these species. We applied allozyme electrophoresis and mitochondrial genome sequencing to infer the extent of genetic differentiation among different cohorts of Xestia tecta (Hübner) within the same geographical area as well as between cohorts with opposite-phase flight pattern in different geographical regions. We found no evidence for substantial genetic differentiation and isolation between the even- and odd-year cohorts in eastern Lapland, nor between the cohorts in eastern and western Lapland. The most informative markers were the most polymorphic allozyme loci (Pgm and Mpi) and the AT-rich region in the mtDNA. However, owing to the generally low levels of genetic variation it was not possible to establish conclusively the degree of genetic isolation between the different cohorts. We discuss the implications of our results in relation to two different hypotheses which could account for this pattern: ongoing gene flow between different cohorts and recent common ancestry.