Abstract
The Dilution Effect Hypothesis (DEH) argues that greater biodiversity lowers the risk of disease and reduces the rates of pathogen transmission since more diverse communities harbour fewer competent hosts for any given pathogen, thereby reducing host exposure to the pathogen. DEH is expected to operate most intensely in vector-borne pathogens and when species-rich communities are not associated with increased host density. Overall, dilution will occur if greater species diversity leads to a lower contact rate between infected vectors and susceptible hosts, and between infected hosts and susceptible vectors. Field-based tests simultaneously analysing the prevalence of several multi-host pathogens in relation to host and vector diversity are required to validate DEH. We tested the relationship between the prevalence in house sparrows (Passer domesticus) of four vector-borne pathogens-three avian haemosporidians (including the avian malaria parasite Plasmodium and the malaria-like parasites Haemoproteus and Leucocytozoon) and West Nile virus (WNV)-and vertebrate diversity. Birds were sampled at 45 localities in SW Spain for which extensive data on vector (mosquitoes) and vertebrate communities exist. Vertebrate censuses were conducted to quantify avian and mammal density, species richness and evenness. Contrary to the predictions of DEH, WNV seroprevalence and haemosporidian prevalence were not negatively associated with either vertebrate species richness or evenness. Indeed, the opposite pattern was found, with positive relationships between avian species richness and WNV seroprevalence, and Leucocytozoon prevalence being detected. When vector (mosquito) richness and evenness were incorporated into the models, all the previous associations between WNV prevalence and the vertebrate community variables remained unchanged. No significant association was found for Plasmodium prevalence and vertebrate community variables in any of the models tested. Despite the studied system having several characteristics that should favour the dilution effect (i.e., vector-borne pathogens, an area where vector and host densities are unrelated, and where host richness is not associated with an increase in host density), none of the relationships between host species diversity and species richness, and pathogen prevalence supported DEH and, in fact, amplification was found for three of the four pathogens tested. Consequently, the range of pathogens and communities studied needs to be broadened if we are to understand the ecological factors that favour dilution and how often these conditions occur in nature.
Highlights
The number of emerging infectious diseases affecting humans is currently increasing [1] and approximately 75% of such diseases are known to be of zoonotic origin [2]
The Dilution Effect Hypothesis (DEH) argues that biodiversity is related to reduced pathogen prevalence because species-rich communities harbour a lower fraction of competent hosts, which reduces pathogen transmission success and, pathogen prevalence [5,6]
The dilution effect was reported by Swaddle et al [10], who noted a lower incidence of West Nile virus (WNV) in humans in US counties with richer avian communities
Summary
The number of emerging infectious diseases affecting humans is currently increasing [1] and approximately 75% of such diseases are known to be of zoonotic origin [2]. Various authors have suggested that more diverse host communities may harbour a higher number of host individuals, which could help vectors proliferate and, eventually, could increase pathogen transmission, thereby neutralizing any potential dilution effect [12]. Rohr et al [25] evaluated the conditions that facilitate a negative relationship between biodiversity and pathogen transmission These authors concluded that the dilution effect is more likely to occur in vector-borne pathogens and will be largely influenced by community assembly rules. Dilution effects may be expected to occur more often when the community assembly is substitutive as opposed to additive In the latter case, the increase in the number of species is associated with increases in host densities since the individuals of the new species are added to those of the species that are already present. It is not possible to analyse how community competence varies within the assemblage, it is possible to analyse whether or not vector and vertebrate community richness and density fit better in additive or substitutive assemblage models
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