Impact of biodiversity and seasonality on Lyme-pathogen transmission

Lyme disease imposes increasing global public health challenges. To better understand the joint effects of seasonal temperature variation and host community composition on the pathogen transmission, a stage-structured periodic model is proposed by integrating seasonal tick development and activity, multiple host species and complex pathogen transmission routes between ticks and reservoirs. Two thresholds, one for tick population dynamics and the other for Lyme-pathogen transmission dynamics, are identified and shown to fully classify the long-term outcomes of the tick invasion and disease persistence. Seeding with the realistic parameters, the tick reproduction threshold and Lyme disease spread threshold are estimated to illustrate the joint effects of the climate change and host community diversity on the pattern of Lyme disease risk. It is shown that climate warming can amplify the disease risk and slightly change the seasonality of disease risk. Both the “dilution effect” and “amplification effect” are observed by feeding the model with different possible alternative hosts. Therefore, the relationship between the host community biodiversity and disease risk varies, calling for more accurate measurements on the local environment, both biotic and abiotic such as the temperature and the host community composition.     

Does life history mediate changing disease risk when communities disassemble?

Biodiversity loss sometimes increases disease risk or parasite transmission in humans, wildlife and plants. Some have suggested that this pattern can emerge when host species that persist throughout community disassembly show high host competence – the ability to acquire and transmit infections. Here, we briefly assess the current empirical evidence for covariance between host competence and extirpation risk, and evaluate the consequences for disease dynamics in host communities undergoing disassembly. We find evidence for such covariance, but the mechanisms for and variability around this relationship have received limited consideration. This deficit could lead to spurious assumptions about how and why disease dynamics respond to community disassembly. Using a stochastic simulation model, we demonstrate that weak covariance between competence and extirpation risk may account for inconsistent effects of host diversity on disease risk that have been observed empirically. This model highlights the predictive utility of understanding the degree to which host competence relates to extirpation risk, and the need for a better understanding of the mechanisms underlying such relationships.     

Fine-scale variation in vector host use and force of infection drive localized patterns of West Nile virus transmission

The influence of host diversity on multi-host pathogen transmission and persistence can be confounded by the large number of species and biological interactions that can characterize many transmission systems. For vector-borne pathogens, the composition of host communities has been hypothesized to affect transmission; however, the specific characteristics of host communities that affect transmission remain largely unknown. We tested the hypothesis that vector host use and force of infection (i.e., the summed number of infectious mosquitoes resulting from feeding upon each vertebrate host within a community of hosts), and not simply host diversity or richness, determine local infection rates of West Nile virus (WNV) in mosquito vectors. In suburban Chicago, Illinois, USA, we estimated community force of infection for West Nile virus using data on Culex pipiens mosquito host selection and WNV vertebrate reservoir competence for each host species in multiple residential and semi-natural study sites. We found host community force of infection interacted with avian diversity to influence WNV infection in Culex mosquitoes across the study area. Two avian species, the American robin (Turdus migratorius) and the house sparrow (Passer domesticus), produced 95.8% of the infectious Cx. pipiens mosquitoes and showed a significant positive association with WNV infection in Culex spp. mosquitoes. Therefore, indices of community structure, such as species diversity or richness, may not be reliable indicators of transmission risk at fine spatial scales in vector-borne disease systems. Rather, robust assessment of local transmission risk should incorporate heterogeneity in vector host feeding and variation in vertebrate reservoir competence at the spatial scale of vector-host interaction.     

The influence of host competition and predation on tick densities and management implications

Host community composition and biodiversity can limit and regulate tick abundance which can have profound impacts on the incidence and severity of tick-borne diseases. Our understanding of the relationship between host community composition and tick abundance is still very limited. Here, we present a novel mathematical model of a stage-structured tick population to study the influence of host behaviour and competition in the presence of heterospecifics and the influence of host predation on tick densities. We examine the influence of specific changes in biodiversity that modify the competition among and the predation on small and large host populations. We find that increasing biodiversity will not always reduce tick populations, but depends on changes in species composition affecting the degree and type competition among hosts, and the host the predation is acting on. With indirect competition, tick densities are not regulated by increasing biodiversity; however, with direct competition, increased biodiversity will regulate tick densities. Generally, we find that biodiversity will regulate tick densities when it affects tick-host encounter rates. We also find that predation on small hosts have a limited influence on reducing tick populations, but when the predation was on large hosts this increased the magnitude of tick population oscillations. Our results have tick-management implications: while controlling large host populations (e.g. deer) and adult ticks will decrease tick densities, measures that directly control the nymph ticks could also be effective.     

Frontiers in research on biodiversity and disease

Global losses of biodiversity have galvanised efforts to understand how changes to communities affect ecological processes, including transmission of infectious pathogens. Here, we review recent research on diversity–disease relationships and identify future priorities. Growing evidence from experimental, observational and modelling studies indicates that biodiversity changes alter infection for a range of pathogens and through diverse mechanisms. Drawing upon lessons from the community ecology of free‐living organisms, we illustrate how recent advances from biodiversity research generally can provide necessary theoretical foundations, inform experimental designs, and guide future research at the interface between infectious disease risk and changing ecological communities. Dilution effects are expected when ecological communities are nested and interactions between the pathogen and the most competent host group(s) persist or increase as biodiversity declines. To move beyond polarising debates about the generality of diversity effects and develop a predictive framework, we emphasise the need to identify how the effects of diversity vary with temporal and spatial scale, to explore how realistic patterns of community assembly affect transmission, and to use experimental studies to consider mechanisms beyond simple changes in host richness, including shifts in trophic structure, functional diversity and symbiont composition.     

Spatiotemporal Model of Barley and Cereal Yellow Dwarf Virus Transmission Dynamics with Seasonality and Plant Competition

Many generalist pathogens are influenced by the spatial distributions and relative abundances of susceptible host species. The spatial structure of host populations can influence patterns of infection incidence (or disease outbreaks), and the effects of a generalist pathogen on host community dynamics in a spatially heterogeneous community may differ from predictions derived via simple models. In this paper, we model the transmission of a generalist pathogen within a patch framework that incorporates the movement of vectors between discrete host patches to investigate the effects of local host community composition and vector movement rates on disease dynamics.     

Traits explain community disassembly and trophic contraction following experimental environmental change

Many ecosystems are currently undergoing dramatic changes in biodiversity due to habitat loss and climate change. Responses to global change at the community level are poorly understood, as are the impacts of community disassembly on ecosystem‐level processes. Uncertainties remain regarding the patterns of extirpation and persistence under single vs. multiple forms of environmental change. Here, we use a trait‐based and food web approach to examine the effects of experimentally changing moisture, temperature and habitat ‘openness’ on a functionally important group of microarthropods associated with a boreal forest floor bryosphere (detrital moss) system. Overall, the outcome of community disassembly was mediated by the correlation between our environmental factors and species traits, particularly body size. Minor increases in summer temperatures maintained greater species richness, whereas drought stress had a significant negative effect on community‐level abundance and richness. These effects were reflected in modifications to the community‐wide body‐size spectra. Habitat openness alleviated biodiversity loss in the larger‐bodied species of the most abundant taxonomic group, but did not fully mitigate the effects of drought. The most striking result of this experiment was an overall contraction of the food web among persistent species under drought stress (i.e. those not extirpated by environmental change). These results suggest that major changes in boreal microarthropod community structure are likely to occur in response to common forms of global change. Moreover, the contraction in trophic structure even amongst tolerant species suggests that ecosystem function within the bryosphere can be altered by environmental change.     

The Effects of Host Diversity on Vector-Borne Disease: The Conditions under Which Diversity Will Amplify or Dilute the Disease Risk

Multihost vector-borne infectious diseases form a significant fraction of the global infectious disease burden. In this study we explore the relationship between host diversity, vector behavior, and disease risk. To this end, we have developed a new dynamic model which includes two distinct host species and one vector species with variable preferences. With the aid of the model we were able to compute the basic reproductive rate, R ₀, a well-established measure of disease risk that serves as a threshold parameter for disease outbreak. The model analysis reveals that the system has two different qualitative behaviors: (i) the well-known dilution effect, where the maximal R₀ is obtained in a community which consists a single host (ii) a new amplification effect, denoted by us as diversity amplification, where the maximal R₀ is attained in a community which consists both hosts. The model analysis extends on previous results by underlining the mechanism of both, diversity amplification and the dilution, and specifies the exact conditions for their occurrence. We have found that diversity amplification occurs where the vector prefers the host with the highest transmission ability, and dilution is obtained when the vector does not show any preference, or it prefers to bite the host with the lower transmission ability. The mechanisms of dilution and diversity amplification are able to account for the different and contradictory patterns often observed in nature (i.e., in some cases disease risk is increased while in other is decreased when the diversity is increased). Implication of the diversity amplification mechanism also challenges current premises about the interaction between biodiversity, climate change, and disease risk and calls for retrospective thinking in planning intervention policies aimed at protecting the preferred host species.     

Effects of species diversity on disease risk

The transmission of infectious diseases is an inherently ecological process involving interactions among at least two, and often many, species. Not surprisingly, then, the species diversity of ecological communities can potentially affect the prevalence of infectious diseases. Although a number of studies have now identified effects of diversity on disease prevalence, the mechanisms underlying these effects remain unclear in many cases. Starting with simple epidemiological models, we describe a suite of mechanisms through which diversity could increase or decrease disease risk, and illustrate the potential applicability of these mechanisms for both vector-borne and non-vector-borne diseases, and for both specialist and generalist pathogens. We review examples of how these mechanisms may operate in specific disease systems. Because the effects of diversity on multi-host disease systems have been the subject of much recent research and controversy, we describe several recent efforts to delineate under what general conditions host diversity should increase or decrease disease prevalence, and illustrate these with examples. Both models and literature reviews suggest that high host diversity is more likely to decrease than increase disease risk. Reduced disease risk with increasing host diversity is especially likely when pathogen transmission is frequency-dependent, and when pathogen transmission is greater within species than between species, particularly when the most competent hosts are also relatively abundant and widespread. We conclude by identifying focal areas for future research, including (1) describing patterns of change in disease risk with changing diversity; (2) identifying the mechanisms responsible for observed changes in risk; (3) clarifying additional mechanisms in a wider range of epidemiological models; and (4) experimentally manipulating disease systems to assess the impact of proposed mechanisms.     

Eutrophication,biodiversity loss,and species invasions modify the relationship between host and parasite richness during host community assembly

Host and parasite richness are generally positively correlated, but the stability of this relationship in response to global change remains poorly understood. Rapidly changing biotic and abiotic conditions can alter host community assembly, which in turn, can alter parasite transmission. Consequently, if the relationship between host and parasite richness is sensitive to parasite transmission, then changes in host composition under various global change scenarios could strengthen or weaken the relationship between host and parasite richness. To test the hypothesis that host community assembly can alter the relationship between host and parasite richness in response to global change, we experimentally crossed host diversity (biodiversity loss) and resource supply to hosts (eutrophication), then allowed communities to assemble. As previously shown, initial host diversity and resource supply determined the trajectory of host community assembly, altering post‐assembly host species richness, richness‐independent host phylogenetic diversity, and colonization by exotic host species. Overall, host richness predicted parasite richness, and as predicted, this effect was moderated by exotic abundance—communities dominated by exotic species exhibited a stronger positive relationship between post‐assembly host and parasite richness. Ultimately, these results suggest that, by modulating parasite transmission, community assembly can modify the relationship between host and parasite richness. These results thus provide a novel mechanism to explain how global environmental change can generate contingencies in a fundamental ecological relationship—the positive relationship between host and parasite richness.     

Mesocosms Reveal Ecological Surprises from Climate Change

Understanding, predicting, and mitigating the impacts of climate change on biodiversity poses one of the most crucial challenges this century. Currently, we know more about how future climates are likely to shift across the globe than about how species will respond to these changes. Two recent studies show how mesocosm experiments can hasten understanding of the ecological consequences of climate change on species’ extinction risk, community structure, and ecosystem functions. Using a large-scale terrestrial warming experiment, Bestion et al. provide the first direct evidence that future global warming can increase extinction risk for temperate ectotherms. Using aquatic mesocosms, Yvon-Durocher et al. show that human-induced climate change could, in some cases, actually enhance the diversity of local communities, increasing productivity. Blending these theoretical and empirical results with computational models will improve forecasts of biodiversity loss and altered ecosystem processes due to climate change.     

Local diversity reduces infection risk across multiple freshwater host‐parasite associations

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Linking community and disease ecology: the impact of biodiversity on pathogen transmission

The increasing number of zoonotic diseases spilling over from a range of wild animal species represents a particular concern for public health, especially in light of the current dramatic trend of biodiversity loss. To understand the ecology of these multi-host pathogens and their response to environmental degradation and species extinctions, it is necessary to develop a theoretical framework that takes into account realistic community assemblages. Here, we present a multi-host species epidemiological model that includes empirically determined patterns of diversity and composition derived from community ecology studies. We use this framework to study the interaction between wildlife diversity and directly transmitted pathogen dynamics. First, we demonstrate that variability in community composition does not affect significantly the intensity of pathogen transmission. We also show that the consequences of community diversity can differentially impact the prevalence of pathogens and the number of infectious individuals. Finally, we show that ecological interactions among host species have a weaker influence on pathogen circulation than inter-species transmission rates. We conclude that integration of a community perspective to study wildlife pathogens is crucial, especially in the context of understanding and predicting infectious disease emergence events.     

Climate change and community disassembly: impacts of warming on tropical and temperate montane community structure

Both tropical and temperate species are responding to global warming through range shifts, but our understanding of the consequences of these shifts for whole communities is limited. Here, we use current elevational range data for six taxonomic groups spanning 90° in latitude to examine the potential impacts of climate-driven range shifts on community change, or 'disassembly', across latitude. Elevational ranges are smaller at low latitudes for most groups and, as a consequence, tropical communities appear to be more sensitive to temperature increases compared with temperate communities. Under site-specific temperature projections, we generally found greater community disassembly in tropical compared with temperate communities, although this varied by dispersal assumptions. Mountain height can impact the amount of community disassembly, with greater change occurring on smaller mountains. Finally, projected community disassembly was higher for ectotherms than endotherms, although the variation among ectotherms was greater than the variation separating endotherms and ectotherms.     

Effect of Biodiversity Changes in Disease Risk: Exploring Disease Emergence in a Plant-Virus System

The effect of biodiversity on the ability of parasites to infect their host and cause disease (i.e. disease risk) is a major question in pathology, which is central to understand the emergence of infectious diseases, and to develop strategies for their management. Two hypotheses, which can be considered as extremes of a continuum, relate biodiversity to disease risk: One states that biodiversity is positively correlated with disease risk (Amplification Effect), and the second predicts a negative correlation between biodiversity and disease risk (Dilution Effect). Which of them applies better to different host-parasite systems is still a source of debate, due to limited experimental or empirical data. This is especially the case for viral diseases of plants. To address this subject, we have monitored for three years the prevalence of several viruses, and virus-associated symptoms, in populations of wild pepper (chiltepin) under different levels of human management. For each population, we also measured the habitat species diversity, host plant genetic diversity and host plant density. Results indicate that disease and infection risk increased with the level of human management, which was associated with decreased species diversity and host genetic diversity, and with increased host plant density. Importantly, species diversity of the habitat was the primary predictor of disease risk for wild chiltepin populations. This changed in managed populations where host genetic diversity was the primary predictor. Host density was generally a poorer predictor of disease and infection risk. These results support the dilution effect hypothesis, and underline the relevance of different ecological factors in determining disease/infection risk in host plant populations under different levels of anthropic influence. These results are relevant for managing plant diseases and for establishing conservation policies for endangered plant species.     

Predicting impacts of climate change on biodiversity: a role for semi‐mechanistic community‐level modelling

Aim Robust and reliable predictions of the effects of climate change on biodiversity are required in formulating conservation and management strategies that best retain biodiversity into the future. Significant challenges in modelling climate change impacts arise from limitations in our current knowledge of biodiversity. Community‐level modelling can complement species‐level approaches in overcoming these limitations and predicting climate change impacts on biodiversity as a whole. However, the community‐level approaches applied to date have been largely correlative, ignoring the key processes that influence change in biodiversity over space and time. Here, we suggest that the development of new ‘semi‐mechanistic’ community‐level models would substantially increase our capacity to predict climate change impacts on biodiversity. Location Global. Methods Drawing on an expansive review of biodiversity modelling approaches and recent advances in semi‐mechanistic modelling at the species level, we outline the main elements of a new semi‐mechanistic community‐level modelling approach. Results Our quantitative review revealed a sharp divide between mechanistic and non‐mechanistic biodiversity modelling approaches, with very few semi‐mechanistic models developed to date. Main conclusions We suggest that the conceptual framework presented here for combining mechanistic and non‐mechanistic community‐level approaches offers a promising means of incorporating key processes into predictions of climate change impacts on biodiversity whilst working within the limits of our current knowledge.     

The influence of host demography, pathogen virulence, and relationships with pathogen virulence on the evolution of pathogen transmission in a spatial context

A major challenge in evolutionary ecology is to explain extensive natural variation in transmission rates and virulence across pathogens. Host and pathogen ecology is a potentially important source of that variation. Theory of its effects has been developed through the study of non-spatial models, but host population spatial structure has been shown to influence evolutionary outcomes. To date, the effects of basic host and pathogen demography on pathogen evolution have not been thoroughly explored in a spatial context. Here we use simulations to show that space produces novel predictions of the influence of the shape of the pathogen’s transmission–virulence tradeoff, as well as host reproduction and mortality, on the pathogen’s evolutionary stable transmission rate. Importantly, non-spatial models predict that neither the slope of linear transmission–virulence relationships, nor the host reproduction rate will influence pathogen evolution, and that host mortality will only influence it when there is a transmission–virulence tradeoff. We show that this is not the case in a spatial context, and identify the ecological conditions under which spatial effects are most influential. Thus, these results may help explain observed natural variation among pathogens unexplainable by non-spatial models, and provide guidance about when space should be considered. We additionally evaluate the ability of existing analytical approaches to predict the influence of ecology, namely spatial moment equations closed with an improved pair approximation (IPA). The IPA is known to have limited accuracy, but here we show that in the context of pathogens the limitations are substantial: in many cases, IPA incorrectly predicts evolution to pathogen-driven extinction. Despite these limitations, we suggest that the impact of ecology can still be understood within the conceptual framework arising from spatial moment equations, that of “self-shading’’, whereby the spread of highly transmissible pathogens is impeded by local depletion of susceptible hosts.     

Dilution effect in bovine tuberculosis: risk factors for regional disease occurrence in Africa

Changes in host diversity have been postulated to influence the risk of infectious diseases, including both dilution and amplification effects. The dilution effect refers to a negative relationship between biodiversity and disease risk, whereas the amplification effect occurs when biodiversity increases disease risk. We tested these effects with an influential disease, bovine tuberculosis (BTB), which is widespread in many countries, causing severe economic losses. Based on the BTB outbreak data in cattle from 2005 to 2010, we also tested, using generalized linear mixed models, which other factors were associated with the regional BTB presence in cattle in Africa. The interdependencies of predictors and their correlations with BTB presence were examined using path analysis. Our results suggested a dilution effect, where increased mammal species richness was associated with reduced probability of BTB presence after adjustment for cattle density. In addition, our results also suggested that areas with BTB infection in the preceding year, higher cattle density and larger percentage of area occupied by African buffalo were more likely to report BTB outbreaks. Climatic variables only indirectly influenced the risk of BTB presence through their effects on cattle density and wildlife distribution. Since most studies investigating the role of wildlife species on BTB transmission only involve single-species analysis, more efforts are needed to better understand the effect of the structure of wildlife communities on BTB dynamics.     

Population dynamics of a polychaete during three El Niño events: disentangling biotic and abiotic factors

The macrobenthic community in shallow soft-bottom areas in the Bay of Ancón, Peru, is characterised by low biodiversity due to low oxygen concentrations. During El Niño events, higher temperature and higher concentrations of dissolved oxygen induce a temporary increase in biodiversity. However, the structure and dynamics of the emerging communities vary strongly among events. The reasons for this variation are poorly understood, in particular the relative influence of abiotic vs biotic factors. To disentangle the roles of abiotic and biotic factors, population models based on detailed monitoring data of three El Niño events were developed focused on the population dynamics of one species in the community, the polychaete Sigambra bassi , which showed different responses in different El Niño events. Calculated and observed population dynamics are compared using root mean square deviation (RMSD). The results show that S. bassi abundance can be determined by abiotic environmental conditions. Besides, three biotic factors improved model performance in different El Niño events: negative density dependence in larval settlement, lower carrying capacity in the presence of other species and positive density dependence in adult survival. The results demonstrate how population models can be used to decode information hidden in long-term monitoring data of abiotic and biotic community parameters about factors driving population dynamics.     

Patterns of parasite community dissimilarity: the significant role of land use and lack of distance‐decay in a bat–helminth system

Increasing community dissimilarity across geographic distance has been described for a wide variety of organisms and understanding its underlying causes is key to understanding mechanisms driving patterns of biodiversity. Both niche‐based and neutral processes may produce a distance decay relationship; however, disentangling their relative influence requires simultaneous examination of multiple potential drivers. Parasites represent a unique opportunity in which to study distance decay because community dissimilarity may depend on environmental requirements and dispersal capability of parasites as well also those of their hosts. We used big brown bats Eptesicus fuscus and their intestinal helminths to investigate: 1) independent contributions of geographic and environmental distances on dissimilarity of intestinal helminth component communities between populations of big brown bats; 2) which environmental variables best explained variation in community dissimilarity; and 3) whether similar patterns of decay with geographic or environmental distance were observed for within‐host population and within‐individual host parasite communities. We used compositional measures of community dissimilarity to examine how parasite communities may change with geographic distance and varying environmental conditions. Non‐spatial variables strongly influenced compositional parasite community dissimilarity over multiple community scales, and we observed little evidence for spatial processes such as distance decay. Environment surrounding roost sites better predicted helminth community dissimilarity than any other class of variables and landcover classes representing anthropogenic modification consistently explained variation in community structure. Our results indicate that human disturbance drives significant patterns of parasite community dissimilarity, most likely by changing the presence or abundance of intermediate hosts in an area.