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AbstractClimate change will alter interactions between parasites and their hosts. Warming may affect patterns of local adaptation, shifting the environment to favor the parasite or host and thus changing the prevalence of disease. We assessed local adaptation to hosts and temperature in the facultative ciliate parasite Lambornella clarki, which infects the western tree hole mosquito Aedes sierrensis. We conducted laboratory infection experiments with mosquito larvae and parasites collected from across a climate gradient, pairing sympatric or allopatric populations across three temperatures that were either matched or mismatched to the source environment. Lambornella clarki parasites were locally adapted to their hosts, with 2.6 times higher infection rates on sympatric populations compared with allopatric populations, but they were not locally adapted to temperature. Infection peaked at the intermediate temperature of 12.5°C, notably lower than the optimum temperature for free-living L. clarki growth, suggesting that the host's immune response can play a significant role in mediating the outcome of infection. Our results highlight the importance of host selective pressure on parasites, despite the impact of temperature on infection success.
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Aedes , Interacciones Huésped-Parásitos , Larva , Temperatura , Animales , Aedes/parasitología , Larva/parasitología , Larva/crecimiento & desarrollo , Adaptación Fisiológica , Apicomplexa/fisiologíaRESUMEN
How mosquitoes may respond to rapid climate warming remains unknown for most species, but will have major consequences for their future distributions, with cascading impacts on human well-being, biodiversity and ecosystem function. We investigated the adaptive potential of a wide-ranging mosquito species, Aedes sierrensis, across a large climatic gradient by conducting a common garden experiment measuring the thermal limits of mosquito life-history traits. Although field-collected populations originated from vastly different thermal environments that spanned over 1200 km, we found limited variation in upper thermal tolerance between populations. In particular, the upper thermal limits of all life-history traits varied by less than 3°C across the species range and, for most traits, did not differ significantly between populations. For one life-history trait-pupal development rate-we did detect significant variation in upper thermal limits between populations, and this variation was strongly correlated with source temperatures, providing evidence of local thermal adaptation for pupal development. However, we found that maximum environmental temperatures across most of the species' range already regularly exceed the highest upper thermal limits estimated under constant temperatures. This result suggests that strategies for coping with and/or avoiding thermal extremes are likely key components of current and future mosquito thermal tolerance.
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Aedes , Ecosistema , Humanos , Animales , Aclimatación , Biodiversidad , Habilidades de AfrontamientoRESUMEN
Variation in heat tolerance among populations can determine whether a species is able to cope with ongoing climate change. Such variation may be especially important for ectotherms whose body temperatures, and consequently, physiological processes, are regulated by external conditions. Additionally, differences in body size are often associated with latitudinal clines, thought to be driven by climate gradients. While studies have begun to explore variation in body size and heat tolerance within species, our understanding of these patterns across large spatial scales, particularly regarding the roles of plasticity and genetic differences, remains incomplete. Here, we examine body size, as measured by wing length, and thermal tolerance, as measured by the time to immobilisation at high temperatures ("thermal knockdown"), in populations of the mosquito Aedes sierrensis collected from across a large latitudinal climate gradient spanning 1300 km (34-44° N). We find that mosquitoes collected from lower latitudes and warmer climates were more tolerant of high temperatures than those collected from higher latitudes and colder climates. Moreover, body size increased with latitude and decreased with temperature, a pattern consistent with James' rule, which appears to be a result of plasticity rather than genetic variation. Our results suggest that warmer environments produce smaller and more thermally tolerant populations.
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BACKGROUND: Aedes (Stegomyia)-borne diseases are an expanding global threat, but gaps in surveillance make comprehensive and comparable risk assessments challenging. Geostatistical models combine data from multiple locations and use links with environmental and socioeconomic factors to make predictive risk maps. Here we systematically review past approaches to map risk for different Aedes-borne arboviruses from local to global scales, identifying differences and similarities in the data types, covariates, and modelling approaches used. METHODS: We searched on-line databases for predictive risk mapping studies for dengue, Zika, chikungunya, and yellow fever with no geographical or date restrictions. We included studies that needed to parameterise or fit their model to real-world epidemiological data and make predictions to new spatial locations of some measure of population-level risk of viral transmission (e.g. incidence, occurrence, suitability, etc.). RESULTS: We found a growing number of arbovirus risk mapping studies across all endemic regions and arboviral diseases, with a total of 176 papers published 2002-2022 with the largest increases shortly following major epidemics. Three dominant use cases emerged: (i) global maps to identify limits of transmission, estimate burden and assess impacts of future global change, (ii) regional models used to predict the spread of major epidemics between countries and (iii) national and sub-national models that use local datasets to better understand transmission dynamics to improve outbreak detection and response. Temperature and rainfall were the most popular choice of covariates (included in 50% and 40% of studies respectively) but variables such as human mobility are increasingly being included. Surprisingly, few studies (22%, 31/144) robustly tested combinations of covariates from different domains (e.g. climatic, sociodemographic, ecological, etc.) and only 49% of studies assessed predictive performance via out-of-sample validation procedures. CONCLUSIONS: Here we show that approaches to map risk for different arboviruses have diversified in response to changing use cases, epidemiology and data availability. We identify key differences in mapping approaches between different arboviral diseases, discuss future research needs and outline specific recommendations for future arbovirus mapping.
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Aedes , Infecciones por Arbovirus , Arbovirus , Fiebre Chikungunya , Dengue , Fiebre Amarilla , Infección por el Virus Zika , Virus Zika , Animales , Humanos , Infecciones por Arbovirus/epidemiología , Fiebre Amarilla/epidemiología , Mosquitos Vectores , Dengue/epidemiologíaRESUMEN
Parasitism is expected to change in a warmer future, but whether warming leads to substantial increases in parasitism remains unclear. Understanding how warming effects on parasitism in individual hosts (e.g. parasite load) translate to effects on population-level parasitism (e.g. prevalence, R0 ) remains a major knowledge gap. We conducted a literature review and identified 24 host-parasite systems that had information on the temperature dependence of parasitism at both individual host and host population levels: 13 vector-borne systems and 11 environmentally transmitted systems. We found a strong positive correlation between the thermal optima of individual- and population-level parasitism, although several of the environmentally transmitted systems exhibited thermal optima >5°C apart between individual and population levels. Parasitism thermal optima were close to vector performance thermal optima in vector-borne systems but not hosts in environmentally transmitted systems, suggesting these thermal mismatches may be more common in certain types of host-parasite systems. We also adapted and simulated simple models for both types of transmission modes and found the same pattern across the two modes: thermal optima were more strongly correlated across scales when there were more traits linking individual- to population-level processes. Generally, our results suggest that information on the temperature dependence, and specifically the thermal optimum, at either the individual or population level should provide a useful-although not quantitatively exact-baseline for predicting temperature dependence at the other level, especially in vector-borne parasite systems. Environmentally transmitted parasitism may operate by a different set of rules, in which temperature dependence is decoupled in some systems, requiring the need for trait-based studies of temperature dependence at individual and population levels.
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Interacciones Huésped-Parásitos , Parásitos , Animales , Simbiosis , TemperaturaRESUMEN
Deforestation and land use change are among the most pressing anthropogenic environmental impacts. In Brazil, a resurgence of malaria in recent decades paralleled rapid deforestation and settlement in the Amazon basin, yet evidence of a deforestation-driven increase in malaria remains equivocal. We hypothesize an underlying cause of this ambiguity is that deforestation and malaria influence each other in bidirectional causal relationships-deforestation increases malaria through ecological mechanisms and malaria reduces deforestation through socioeconomic mechanisms-and that the strength of these relationships depends on the stage of land use transformation. We test these hypotheses with a large geospatial dataset encompassing 795 municipalities across 13 y (2003 to 2015) and show deforestation has a strong positive effect on malaria incidence. Our results suggest a 10% increase in deforestation leads to a 3.3% increase in malaria incidence (â¼9,980 additional cases associated with 1,567 additional km2 lost in 2008, the study midpoint, Amazon-wide). The effect is larger in the interior and absent in outer Amazonian states where little forest remains. However, this strong effect is only detectable after controlling for a feedback of malaria burden on forest loss, whereby increased malaria burden significantly reduces forest clearing, possibly mediated by human behavior or economic development. We estimate a 1% increase in malaria incidence results in a 1.4% decrease in forest area cleared (â¼219 fewer km2 cleared associated with 3,024 additional cases in 2008). This bidirectional socioecological feedback between deforestation and malaria, which attenuates as land use intensifies, illustrates the intimate ties between environmental change and human health.
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Conservación de los Recursos Naturales , Malaria/transmisión , Brasil , Humanos , Malaria/epidemiología , Bosque LluviosoRESUMEN
Competitive exclusion can be classified as deterministic or as historically contingent. While competitive exclusion is common in nature, it has remained unclear when multispecies communities formed by more than two species should be dominated by deterministic or contingent exclusion. Here, we take a fully parameterised model of an empirical competitive system between invasive annual and native perennial plant species to explain both the emergence and sources of competitive exclusion in multispecies communities. Using a structural approach to understand the range of parameters promoting deterministic and contingent exclusions, we then find heuristic theoretical support for the following three general conclusions. First, we find that the life-history of perennial species increases the probability of observing contingent exclusion by increasing their effective intrinsic growth rates. Second, we find that the probability of observing contingent exclusion increases with weaker intraspecific competition, and not with the level of hierarchical competition. Third, we find a shift from contingent exclusion to deterministic exclusion with increasing numbers of competing species. Our work provides a heuristic framework to increase our understanding about the predictability of species persistence within multispecies communities.
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PlantasRESUMEN
Experiments and models suggest that climate affects mosquito-borne disease transmission. However, disease transmission involves complex nonlinear interactions between climate and population dynamics, which makes detecting climate drivers at the population level challenging. By analysing incidence data, estimated susceptible population size, and climate data with methods based on nonlinear time series analysis (collectively referred to as empirical dynamic modelling), we identified drivers and their interactive effects on dengue dynamics in San Juan, Puerto Rico. Climatic forcing arose only when susceptible availability was high: temperature and rainfall had net positive and negative effects respectively. By capturing mechanistic, nonlinear and context-dependent effects of population susceptibility, temperature and rainfall on dengue transmission empirically, our model improves forecast skill over recent, state-of-the-art models for dengue incidence. Together, these results provide empirical evidence that the interdependence of host population susceptibility and climate drives dengue dynamics in a nonlinear and complex, yet predictable way.
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Dengue , Animales , Dengue/epidemiología , Susceptibilidad a Enfermedades , Dinámica Poblacional , Puerto Rico/epidemiología , TemperaturaRESUMEN
Vector-borne diseases (VBDs) are embedded within complex socio-ecological systems. While research has traditionally focused on the direct effects of VBDs on human morbidity and mortality, it is increasingly clear that their impacts are much more pervasive. VBDs are dynamically linked to feedbacks between environmental conditions, vector ecology, disease burden, and societal responses that drive transmission. As a result, VBDs have had profound influence on human history. Mechanisms include: (1) killing or debilitating large numbers of people, with demographic and population-level impacts; (2) differentially affecting populations based on prior history of disease exposure, immunity, and resistance; (3) being weaponised to promote or justify hierarchies of power, colonialism, racism, classism and sexism; (4) catalysing changes in ideas, institutions, infrastructure, technologies and social practices in efforts to control disease outbreaks; and (5) changing human relationships with the land and environment. We use historical and archaeological evidence interpreted through an ecological lens to illustrate how VBDs have shaped society and culture, focusing on case studies from four pertinent VBDs: plague, malaria, yellow fever and trypanosomiasis. By comparing across diseases, time periods and geographies, we highlight the enormous scope and variety of mechanisms by which VBDs have influenced human history.
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Malaria , Enfermedades Transmitidas por Vectores , Vectores de Enfermedades , HumanosRESUMEN
Mathematical models of epidemics are important tools for predicting epidemic dynamics and evaluating interventions. Yet, because early models are built on limited information, it is unclear how long they will accurately capture epidemic dynamics. Using a stochastic SEIR model of COVID-19 fitted to reported deaths, we estimated transmission parameters at different time points during the first wave of the epidemic (March-June, 2020) in Santa Clara County, California. Although our estimated basic reproduction number ([Formula: see text]) remained stable from early April to late June (with an overall median of 3.76), our estimated effective reproduction number ([Formula: see text]) varied from 0.18 to 1.02 in April before stabilizing at 0.64 on 27 May. Between 22 April and 27 May, our model accurately predicted dynamics through June; however, the model did not predict rising summer cases after shelter-in-place orders were relaxed in June, which, in early July, was reflected in cases but not yet in deaths. While models are critical for informing intervention policy early in an epidemic, their performance will be limited as epidemic dynamics evolve. This paper is one of the first to evaluate the accuracy of an early epidemiological compartment model over time to understand the value and limitations of models during unfolding epidemics.
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COVID-19 , Epidemias , Número Básico de Reproducción , Humanos , Modelos Teóricos , SARS-CoV-2RESUMEN
Lyme disease is the most common vector-borne disease in temperate zones and a growing public health threat in the United States (US). The life cycles of the tick vectors and spirochete pathogen are highly sensitive to climate, but determining the impact of climate change on Lyme disease burden has been challenging due to the complex ecology of the disease and the presence of multiple, interacting drivers of transmission. Here we incorporated 18 years of annual, county-level Lyme disease case data in a panel data statistical model to investigate prior effects of climate variation on disease incidence while controlling for other putative drivers. We then used these climate-disease relationships to project Lyme disease cases using CMIP5 global climate models and two potential climate scenarios (RCP4.5 and RCP8.5). We find that interannual variation in Lyme disease incidence is associated with climate variation in all US regions encompassing the range of the primary vector species. In all regions, the climate predictors explained less of the variation in Lyme disease incidence than unobserved county-level heterogeneity, but the strongest climate-disease association detected was between warming annual temperatures and increasing incidence in the Northeast. Lyme disease projections indicate that cases in the Northeast will increase significantly by 2050 (23,619 ± 21,607 additional cases), but only under RCP8.5, and with large uncertainty around this projected increase. Significant case changes are not projected for any other region under either climate scenario. The results demonstrate a regionally variable and nuanced relationship between climate change and Lyme disease, indicating possible nonlinear responses of vector ticks and transmission dynamics to projected climate change. Moreover, our results highlight the need for improved preparedness and public health interventions in endemic regions to minimize the impact of further climate change-induced increases in Lyme disease burden.
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Ixodes , Enfermedad de Lyme , Animales , Cambio Climático , Predicción , Incidencia , Enfermedad de Lyme/epidemiología , Estados Unidos/epidemiologíaRESUMEN
In the aftermath of the 2015 pandemic of Zika virus (ZIKV), concerns over links between climate change and emerging arboviruses have become more pressing. Given the potential that much of the world might remain at risk from the virus, we used a previously established temperature-dependent transmission model for ZIKV to project climate change impacts on transmission suitability risk by mid-century (a generation into the future). Based on these model predictions, in the worst-case scenario, over 1.3 billion new people could face suitable transmission temperatures for ZIKV by 2050. The next generation will face substantially increased ZIKV transmission temperature suitability in North America and Europe, where naïve populations might be particularly vulnerable. Mitigating climate change even to moderate emissions scenarios could significantly reduce global expansion of climates suitable for ZIKV transmission, potentially protecting around 200 million people. Given these suitability risk projections, we suggest an increased priority on research establishing the immune history of vulnerable populations, modeling when and where the next ZIKV outbreak might occur, evaluating the efficacy of conventional and novel intervention measures, and increasing surveillance efforts to prevent further expansion of ZIKV.
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Aedes , Infección por el Virus Zika , Virus Zika , Animales , Europa (Continente) , Humanos , Mosquitos Vectores , América del Norte , Temperatura , Infección por el Virus Zika/epidemiologíaRESUMEN
Mosquito-borne diseases cause a major burden of disease worldwide. The vital rates of these ectothermic vectors and parasites respond strongly and nonlinearly to temperature and therefore to climate change. Here, we review how trait-based approaches can synthesise and mechanistically predict the temperature dependence of transmission across vectors, pathogens, and environments. We present 11 pathogens transmitted by 15 different mosquito species - including globally important diseases like malaria, dengue, and Zika - synthesised from previously published studies. Transmission varied strongly and unimodally with temperature, peaking at 23-29ºC and declining to zero below 9-23ºC and above 32-38ºC. Different traits restricted transmission at low versus high temperatures, and temperature effects on transmission varied by both mosquito and parasite species. Temperate pathogens exhibit broader thermal ranges and cooler thermal minima and optima than tropical pathogens. Among tropical pathogens, malaria and Ross River virus had lower thermal optima (25-26ºC) while dengue and Zika viruses had the highest (29ºC) thermal optima. We expect warming to increase transmission below thermal optima but decrease transmission above optima. Key directions for future work include linking mechanistic models to field transmission, combining temperature effects with control measures, incorporating trait variation and temperature variation, and investigating climate adaptation and migration.
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Aedes/parasitología , Aedes/virología , Temperatura , Animales , Cambio Climático , Virus del Dengue , Malaria/transmisión , Mosquitos Vectores/parasitología , Mosquitos Vectores/virología , Plasmodium , Virus del Río Ross , Virosis/transmisión , Virus ZikaRESUMEN
Niche and fitness differences control the outcome of competition, but determining their relative importance in invaded communities-which may be far from equilibrium-remains a pressing concern. Moreover, it is unclear whether classic approaches for studying competition, which were developed predominantly for pairs of interacting species, will fully capture dynamics in complex species assemblages. We parameterized a population-dynamic model using competition experiments of two native and three exotic species from a grassland community. We found evidence for minimal fitness differences or niche differences between the native species, leading to slow replacement dynamics and priority effects, but large fitness advantages allowed exotics to unconditionally invade natives. Priority effects driven by strong interspecific competition between exotic species drove single-species dominance by one of two exotic species in 80% of model outcomes, while a complex mixture of nonhierarchical competition and coexistence between native and exotic species occurred in the remaining 20%. Fungal infection, a commonly hypothesized coexistence mechanism, had weak fitness effects and is unlikely to substantially affect coexistence. In contrast to previous work on pairwise outcomes in largely native-dominated communities, our work supports a role for nearly neutral dynamics and priority effects as drivers of species composition in invaded communities.
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Pradera , Especies Introducidas , Modelos Biológicos , Poaceae/crecimiento & desarrollo , Poaceae/microbiologíaRESUMEN
Between 2015 and 2017, Zika virus spread rapidly through populations in the Americas with no prior exposure to the disease. Although climate is a known determinant of many Aedes-transmitted diseases, it is currently unclear whether climate was a major driver of the Zika epidemic and how climate might have differentially impacted outbreak intensity across locations within Latin America. Here, we estimated force of infection for Zika over time and across provinces in Latin America using a time-varying susceptible-infectious-recovered model. Climate factors explained less than 5% of the variation in weekly transmission intensity in a spatio-temporal model of force of infection by province over time, suggesting that week to week transmission within provinces may be too stochastic to predict. By contrast, climate and population factors were highly predictive of spatial variation in the presence and intensity of Zika transmission among provinces, with pseudo-R2 values between 0.33 and 0.60. Temperature, temperature range, rainfall and population size were the most important predictors of where Zika transmission occurred, while rainfall, relative humidity and a nonlinear effect of temperature were the best predictors of Zika intensity and burden. Surprisingly, force of infection was greatest in locations with temperatures near 24°C, much lower than previous estimates from mechanistic models, potentially suggesting that existing vector control programmes and/or prior exposure to other mosquito-borne diseases may have limited transmission in locations most suitable for Aedes aegypti, the main vector of Zika, dengue and chikungunya viruses in Latin America.
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Clima , Infección por el Virus Zika/epidemiología , Virus Zika , Animales , Brotes de Enfermedades , Vectores de Enfermedades , Exposición a Riesgos Ambientales/estadística & datos numéricos , Epidemias , Humanos , América Latina/epidemiología , Mosquitos VectoresRESUMEN
Temperature is a strong driver of vector-borne disease transmission. Yet, for emerging arboviruses we lack fundamental knowledge on the relationship between transmission and temperature. Current models rely on the untested assumption that Zika virus responds similarly to dengue virus, potentially limiting our ability to accurately predict the spread of Zika. We conducted experiments to estimate the thermal performance of Zika virus (ZIKV) in field-derived Aedes aegypti across eight constant temperatures. We observed strong, unimodal effects of temperature on vector competence, extrinsic incubation period and mosquito survival. We used thermal responses of these traits to update an existing temperature-dependent model to infer temperature effects on ZIKV transmission. ZIKV transmission was optimized at 29°C, and had a thermal range of 22.7°C-34.7°C. Thus, as temperatures move towards the predicted thermal optimum (29°C) owing to climate change, urbanization or seasonality, Zika could expand north and into longer seasons. By contrast, areas that are near the thermal optimum were predicted to experience a decrease in overall environmental suitability. We also demonstrate that the predicted thermal minimum for Zika transmission is 5°C warmer than that of dengue, and current global estimates on the environmental suitability for Zika are greatly over-predicting its possible range.
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Aedes/fisiología , Cambio Climático , Mosquitos Vectores/fisiología , Temperatura , Infección por el Virus Zika/transmisión , Virus Zika/fisiología , Aedes/virología , Animales , Modelos Biológicos , Mosquitos Vectores/virología , Estaciones del Año , UrbanizaciónRESUMEN
Pathogen infection is common in wild plants and animals, and may regulate their populations. If pathogens have narrow host ranges and increase with the density of their favored hosts, they may promote host species diversity by suppressing common species to the benefit of rare species. Yet, because many pathogens infect multiple co-occurring hosts, they may not strongly respond to the relative abundance of a single host species. Are natural communities dominated by specialized pathogens that respond to the relative abundance of a specific host or by pathogens with broad host ranges and limited responses to the relative abundance of single host? The answer determines the potential for pathogens to promote host coexistence, as often hypothesized, or to have negligible or even negative effects on host coexistence. We lack a systematic understanding of the impacts, identities, and host ranges of pathogens in natural communities. Here we characterize a community of foliar fungal pathogens and evaluate their host specificity and fitness impacts in a California grassland community of native and exotic species. We found that most of the commonly isolated fungal pathogens were multi-host, with intermediate to low specialization. The amount of pathogen damage each host experienced was independent of host species local relative abundance. Despite pathogen sharing among the host species, fungal communities slightly differed in composition across host species. Plants with high pathogen damage tended to have lower seed production but the relationship was weak, suggesting limited fitness impacts. Moreover, seed production was not dependent on the local relative abundance of each plant species, suggesting that stabilizing coexistence mechanisms may operate at larger spatial scales in this community. Because foliar pathogens in this grassland community are multi-host and have small fitness impacts, they are unlikely to promote negative frequency dependence or plant species coexistence in this system. Still, given that pathogen community composition differentiates across host species, some more subtle feedbacks between host relative abundance and pathogen community composition, damage, and fitness impacts are possible, which could, in turn, promote either coexistence or competitive exclusion.
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Pradera , Plantas/microbiología , Animales , California , Hongos , Especificidad del HuéspedRESUMEN
Pathogens live in diverse, competitive communities, yet the processes that maintain pathogen diversity remain elusive. Here, we use a species-rich, well-studied plant virus system, the barley yellow dwarf viruses, to examine the mechanisms that regulate pathogen diversity. We empirically parameterized models of three viruses, their two aphid vectors, and one perennial grass host. We found that high densities of both aphids maximized virus diversity and that competition limited the coexistence of two closely related viruses. Even limited ability to simultaneously infect (coinfect) host individuals strongly promoted virus coexistence; preventing coinfection led to priority effects. Coinfection generated stabilizing niche differences by allowing viruses to share hosts. However, coexistence also required trade-offs between vector generalist and specialist life-history strategies. Our predicted outcomes broadly concur with previous field observations. These results show how competition within individual hosts and vectors may lead to unexpected population-level outcomes between pathogens, including coexistence, competitive exclusion, and priority effects, and how contemporary coexistence theory can help to predict these outcomes.
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Áfidos/virología , Luteovirus/fisiología , Modelos Biológicos , Enfermedades de las Plantas/virología , Poaceae/virología , Animales , Biodiversidad , Interacciones Huésped-Patógeno , Insectos Vectores/virología , Densidad de PoblaciónRESUMEN
Competition - colonization tradeoffs occur in many systems, and theory predicts that they can strongly promote species coexistence. However, there is little empirical evidence that observed competition- colonization tradeoffs are strong enough to maintain diversity in natural systems. This is due in part to a mismatch between theoretical assumptions and biological reality in some systems. We tested whether a competition - colonization tradeoff explains how a diverse trematode guild coexists in California horn snail populations, a system that meets the requisite criteria for the tradeoff to promote coexistence. A field experiment showed that subordinate trematode species tended to have higher colonization rates than dominant species. This tradeoff promoted coexistence in parameterized models but did not fully explain trematode diversity and abundance, suggesting a role of additional diversity maintenance mechanisms. Spatial heterogeneity is an alternative way to promote coexistence if it isolates competing species. We used scale transition theory to expand the competition - colonization tradeoff model to include spatial variation. The parameterized model showed that spatial variation in trematode prevalence did not isolate most species sufficiently to explain the overall high diversity, but could benefit some rare species. Together, the results suggest that several mechanisms combine to maintain diversity, even when a competition - colonization tradeoff occurs.
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Caracoles/fisiología , Trematodos/fisiología , Animales , California , Conducta Competitiva , Ecosistema , Modelos BiológicosRESUMEN
Competition - colonization tradeoffs occur in many systems, and theory predicts that they can strongly promote species coexistence. However, there is little empirical evidence that observed competition - colonization tradeoffs are strong enough to maintain diversity in natural systems. This is due in part to a mismatch between theoretical assumptions and biological reality in some systems. We tested whether a competition - colonization tradeoff explains how a diverse trematode guild coexists in California horn snail populations, a system that meets the requisite criteria for the tradeoff to promote coexistence. A field experiment showed that subordinate trematode species tended to have higher colonization rates than dominant species. This tradeoff promoted coexistence in parameterized models but did not fully explain trematode diversity and abundance, suggesting a role of additional diversity maintenance mechanisms. Spatial heterogeneity is an alternative way to promote coexistence if it isolates competing species. We used scale transition theory to expand the competition - colonization tradeoff model to include spatial variation. The parameterized model showed that spatial variation in trematode prevalence did not isolate most species sufficiently to explain the overall high diversity, but could benefit some rare species. Together, the results suggest that several mechanisms combine to maintain diversity, even when a competition - colonization tradeoff occurs.