Controlling Echinococcus multilocularis-ecological implications of field trials

Two field trials to reduce the prevalence of Echinococcus multilocularis in foxes have been conducted in recent years. Although both trials reduced prevalence considerably, they failed to eradicate the parasite in the study region. Following the control trial in northern Germany, prevalence recovered unexpectedly and rapidly, reaching pre-control levels five quarters (15 months) after the end of control. To understand the internal dynamics of the parasite-host system's reaction to control, we developed a spatially explicit simulation model, Echi. The simulation model incorporates the information available concerning fox tapeworm population dynamics.Using epidemiological parameters to adjust pre-control prevalence, the model predicts the temporal evolution of the prevalence of E. multilocularis in controlled foxes without departing from the range of uncertainty of the field data. However, the model does not predict the rapid pre-control recovery observed in the field trial.The deviation of the model's prediction from field data indicates the involvement of processes not yet taken into account. We modified the model step by step to mimic processes with the potential to cause the rapid post-control recovery of the prevalence of E. multilocularis in foxes.Neither the longevity of tapeworm eggs nor the migratory behaviour of foxes showed any influence on the post-control reaction of the parasite-host system. However, landscape structures leading to a heterogeneous distribution of infected foxes have the potential to alter the system's reaction to control. If infected foxes are concentrated in multiple clusters in the landscape, the model prediction tallied with the range of uncertainty of the field data. Such spatial distribution of infected foxes may be caused by differential abiotic conditions influencing the survival of tapeworm eggs.The model was found to comply best with field data if the foxes acquire partial immunity by being exposed to the fox tapeworm.Both hypotheses explaining the rapid post-control recovery of the prevalence of E. multilocularis observed in the fox population were supported by field data.Both hypotheses have far-reaching consequences for future control trials. The spatial aggregation of infected foxes would enable control efforts to be concentrated on these highly infected areas. However, the acquisition of immunity acts as a buffer to control, necessitating intensified control measures.     

Resource-use,body condition and parasite load metrics indicate contrasting health of stocked and native game fishes in Canadian prairie lakes

This study evaluated the status of native and stocked fish species in 13 prairie lakes in central Canada over eight years (2007–14) using three metrics: resource-use (benthic versus pelagic carbon via stable isotopes); body condition (relative weight index W_r); and parasite load (cestode gut enumerations). Analyses included game and non-game fishes, like naturally occurring northern pike, Esox lucius L., and yellow perch, Perca flavescens Mitchill, but focused on stocked walleye, Sander vitreus (Mitchill) because it supports a robust recreational fishery. Walleye and northern pike were significantly more reliant on benthic carbon than yellow perch or forage fishes (p < 0.05), but this reliance was not associated with any measured environmental variables for any species. In lakes with game fishes, forage fishes exhibited higher reliance on benthic energy, possibly due to predator avoidance strategy, particularly yellow perch. Walleye body condition index was consistently lower (<95–105) than values exhibited by the other two game fishes (81–139), and parasites were only common in walleye and associated with lake-water salinity (r² = 0.93, p < 0.05) and sex (p < 0.05). Based on the results, the most desirable game fish, walleye, appears less resilient to environmental variability than northern pike and yellow perch, making this species more susceptible to impacts of future climate change.     

Impaired littoral energy pathways cause a shift to pelagic resources by fish in recovering lake food webs

Fish communities may increase in biomass and productivity due to energy subsidies from the littoral invertebrate community. In lakes recovering from acidification and metal contamination, such as those in Sudbury, Ontario, Canada, impaired benthic invertebrate communities (i.e., low diversity with higher abundance of small‐bodied taxa) allowed a critical test of the role of these littoral pathways on fish diet. We compared fish abundance, diversity, diet and biomass in eight recovering and eight reference lakes and related availability of the main littoral and pelagic invertebrate groups to fish diet regime using stable isotope analysis. A Bayesian mixing model (MixSIR) was used to estimate diet likelihood, and convex hull analysis was used to estimate trophic niche space of fish communities. Fish biomass did not differ between impaired and reference lakes despite substantial differences in potential diet. Fish depended strongly on littoral benthos in the reference lakes but consumed more pelagic food in the impaired lakes. The trophic niche of the focal, most common fish species (i.e., yellow perch, smallmouth bass, pumpkinseed and brown bullhead) was larger in the impaired lakes. We attributed these differences to low diversity at the highest trophic levels of fish communities in the impaired lakes as well as to depauperate benthic invertebrate communities. In contrast to the food webs of most temperate lakes, fish in impaired lakes preyed less on littoral invertebrates yet still managed to maintain a reference lake level of biomass standing crop by relying more on pelagic resources – macro zooplankton such as Chaoborus.     

The Viability of Metapopulations: Individual Dispersal Behaviour Matters

Metapopulation models are frequently used for analysing species–landscape interactions and their effect on structure and dynamic of populations in fragmented landscapes. They especially support a better understanding of the viability of metapopulations. In such models, the processes determining metapopulation viability are often modelled in a simple way. Animals’ dispersal between habitat fragments is mostly taken into account by using a simple dispersal function that assumes the underlying process of dispersal to be random movement. Species-specific dispersal behaviour such as a systematic search for habitat patches is likely to influence the viability of a metapopulation. Using a model for metapopulation viability analysis, we investigate whether such specific dispersal behaviour affects the predictions of ranking orders among alternative landscape configurations rated regarding their ability to carry viable metapopulations. To incorporate dispersal behaviour in the model, we use a submodel for the colonisation rates which allows different movement patterns to be considered (uncorrelated random walk, correlated random walk with various degrees of correlation, and loops). For each movement pattern, the landscape order is determined by comparing the resulting mean metapopulation lifetime T_m of different landscape configurations. Results show that landscape orders can change considerably between different movement patterns. We analyse whether and under what circumstances dispersal behaviour influences the ranking orders of landscapes. We find that the ‘competition between patches for migrants’ – i.e. the fact that dispersers immigrating into one patch are not longer available as colonisers for other patches – is an important factor driving the change in landscape ranks. The implications of our results for metapopulation modelling, planning and conservation are discussed.     

Nitrogen fertilization and liming increased CO2 and N2O emissions from tropical ferralsols,but not from a vertisol

The application of nitrogen (N) fertilizers and liming (CaCO₃) to improve soil quality and crop productivity are regarded as effective and important agricultural practices. However, they may increase greenhouse gas (GHG) emissions. There is limited information on the GHG emissions of tropical soils, specifically when liming is combined with N fertilization. We therefore conducted a full factorial laboratory incubation experiment to investigate how N fertilizer (0 kg N ha⁻¹, 12.5 kg N ha⁻¹ and 50 kg N ha⁻¹) and liming (target pH = 6.5) affect GHG emissions and soil N availability. We focussed on three common acidic soils (two ferralsols and one vertisol) from Lake Victoria (Kenya). After 8 weeks, the most significant increase in cumulative carbon dioxide (CO₂) and nitrous oxide (N₂O) fluxes compared with the unfertilized control was found for the two ferralsols in the N + lime treatment, with five to six times higher CO₂ fluxes than the control. The δ¹³C signature of soil-emitted CO₂ revealed that for the ferralsols, liming (i.e. the addition of CaCO₃) was the dominant source of CO₂, followed by urea (N fertilization), whereas no significant effect of liming or of N fertilization on CO₂ flux was found for the vertisol. In addition, the N₂O fluxes were most significantly increased by the high N + lime treatment in the two ferralsols, with four times and 13 times greater N₂O flux than that of the control. No treatment effects on N₂O fluxes were observed for the vertisol. Liming in combination with N fertilization significantly increased the final nitrate content by 14.5%–39% compared with N fertilization alone in all treatment combinations and soils. We conclude that consideration should be given to the GHG budgets of agricultural ferralsols since liming is associated with high liming-induced CO₂ and N₂O emissions. Therefore, nature-based and sustainable sources should be explored as an alternative to liming in order to manage the pH and the associated fertility of acidic tropical soils.     

Dispersal behaviour in fragmented landscapes: Deriving a practical formula for patch accessibility

Dispersal has been shown to be a key determinant of spatially structured populations. One crucial aspect is predicting patch accessibility: the probability r_ij of a certain patch j being reached by individuals starting at another patch i. Patch accessibility r_ij depends on both the landscape structure and the individuals’ dispersal behaviour. To investigate the effects of these factors on r_ij, we developed a simulation model focusing on animal dispersal. Our model analyses show that there is an important intrinsic effect of the interplay between landscape structure and dispersal behaviour on patch accessibility: the competition between patches for migrants. We derive a formula for patch accessibility. This formula is very simple because it just takes distances into account: not only the distance between start patch and target patch, but also between the start patch and all the other patches in the landscape. Despite its simplicity, the formula is able to cover effects such as the competition for migrants. The formula was found to have high predictive power for a variety of movement behaviours (random walk with various degrees of correlation, Archimedean spirals and loops) in any given landscape. The formula can be interpreted as a generic function for patch accessibility for further population dynamics analyses. It also delivers insights into the consequences of dispersal in fragmented landscapes.     

Spatial aspects of metapopulation survival – from model results to rules of thumb for landscape management

The role of spatial configuration for metapopulation survival is analyzed by using a stochastic metapopulation model. This model reveals conditions which must be satisfied by the species' ecology and the landscape settings before a metapopulation can persist over a long term. Taking this as a basis, initial rules of thumb for landscape management are deduced. The following results are highlighted: (1) the critical correlation length d_c of the extinction processes determines a spatial scale of the metapopulation dynamics. (2) Only species with a dispersal range d_r above the correlation length d_c are able to benefit from landscape management at all. (3) A certain metapopulation can only persist over a long term if no patch is inside the range of correlation of another one. (4) There is a hierarchy of importance in the characteristics of a spatial configuration (scale and type) and, hence, in the scopes of landscape management. To conclude, some general consequences for supporting species survival by management are discussed. Some prospects concerning the use of models for decision support in landscape planning are discussed.     

The spatio-temporal dynamics of a post-vaccination resurgence of rabies in foxes and emergency vaccination planning

We used a simulation model to study the spatio-temporal dynamics of a potential rabies outbreak in an immunized fox population after the termination of a long-term, large-scale vaccination program with two campaigns per year one in spring and one in autumn. The 'worst-case' scenario of rabies resurgence occurs if rabies has persisted at a low prevalence despite control and has remained undetected by a customary surveillance program or if infected individuals invade to the control area. Even if the termination of a vaccination program entails such a risk of a subsequent new outbreak, prolonged vaccination of a wild host population is expensive and the declining cost-benefit ratio over time eventually makes it uneconomic. Based on the knowledge of the spatio-temporal dynamics of a potential new outbreak gained from our modelling study, we suggest "terminating but observing" to be an appropriate strategy. Simulating the decline of population immunity without revaccination, we found that a new outbreak of rabies should be detected by customary surveillance programs within two years after the termination of the control. The time until detection does not depend on whether vaccination was terminated within the fourth, fifth or sixth years of repeated biannual campaigns. But it is faster if the program was completed with an autumn campaign (because next-year dispersal then occurs after a noticeable decrease in population immunity). Finally, if a rabid fox is detected after terminating vaccination, we determine a rule for defining a circular hazard area based on the simulated spatial spread of rabies. The radius of this area should be increased with the time since the last vaccination campaign. The trade-off between the number of foxes potentially missed by the emergency treatment and the cost for the emergency measures in an enlarged hazard area was found.