Spatial Distribution Of Hosts Research Articles

Spatial effects in parasite-induced marine diseases of immobile hosts.

Emerging marine infectious diseases pose a substantial threat to marine ecosystems and the conservation of their biodiversity. Compartmental models of epidemic transmission in marine sessile organisms, available only recently, are based on non-spatial descriptions in which space is homogenized and parasite mobility is not explicitly accounted for. However, in realistic scenarios epidemic transmission is conditioned by the spatial distribution of hosts and the parasites' mobility patterns, calling for an explicit description of space. In this work, we develop a spatially explicit individual-based model to study disease transmission by waterborne parasites in sessile marine populations. We investigate the impact of spatial disease transmission through extensive numerical simulations and theoretical analysis. Specifically, the effects of parasite mobility into the epidemic threshold and the temporal progression of the epidemic are assessed. We show that larger values of pathogen mobility imply more severe epidemics, as the number of infections increases, and shorter timescales to extinction. An analytical expression for the basic reproduction number of the spatial model, , is derived as a function of the non-spatial counterpart, R0, which characterizes a transition between a disease-free and a propagation phase, in which the disease propagates over a large fraction of the system.

Species distribution patterns and the scale of host interactions quantitatively but not qualitatively affect the diversity–disease relationship

Evidence is accumulating to show that changes in host diversity can affect disease risk, yet the relationship between them is still disputed. Understanding what factors influence this relationship will help to reconcile this debate. In this paper, through spatially-explicit simulations, we investigated the effects of host spatial distribution patterns, the scale of host interactions (i.e. local vs. global interactions), the presence or absence of life history trade-offs, and community assembly rules on the diversity–disease relationship when the pathogen was a generalist. The results showed that intraspecific aggregation enhanced disease risk, while increased global interactions between hosts reduced disease risk. For a given host species richness level, the effect of the scale of host interactions on disease risk increased with intraspecific aggregation. However, neither species distribution patterns nor the scale of host interactions could reverse the diversity-disease relationship. The direction and strength of diversity on disease risk depended on community assembly and life history trade-offs. These results suggest that species distribution patterns and the scale of host interactions do not contribute to the discrepancy in results that lead to the debate over dilution effect, highlighting the complexity of mechanisms underlying the diversity–disease relationship.

Trade-off between local transmission and long-range dispersal drives infectious disease outbreak size in spatially structured populations.

Transmission of infectious diseases between immobile hosts (e.g., plants, farms) is strongly dependent on the spatial distribution of hosts and the distance-dependent probability of transmission. As the interplay between these factors is poorly understood, we use spatial process and transmission modelling to investigate how epidemic size is shaped by host clustering and spatial range of transmission. We find that for a given degree of clustering and individual-level infectivity, the probability that an epidemic occurs after an introduction is generally higher if transmission is predominantly local. However, local transmission also impedes transfer of the infection to new clusters. A consequence is that the total number of infections is maximal if the range of transmission is intermediate. In highly clustered populations, the infection dynamics is strongly determined by the probability of transmission between clusters of hosts, whereby local clusters act as multiplier of infection. We show that in such populations, a metapopulation model sometimes provides a good approximation of the total epidemic size, using probabilities of local extinction, the final size of infections in local clusters, and probabilities of cluster-to-cluster transmission. As a real-world example we analyse the case of avian influenza transmission between poultry farms in the Netherlands.

Gut microbial diversity across a contact zone for California voles: Implications for lineage divergence of hosts and mitonuclear mismatch in the assembly of the mammalian gut microbiome.

Gut microbial diversity is thought to reflect the co-evolution of microbes and their hosts as well as current host-specific attributes such as genetic background and environmental setting. To explore interactions among these parameters, we characterized variation in gut microbiome composition of California voles (Microtus californicus) across a contact zone between two recently diverged lineages of this species. Because this contact zone contains individuals with mismatched mitochondrial-nuclear genomes (cybrids), it provides an important opportunity to explore how different components of the genotype contribute to gut microbial diversity. Analyses of bacterial 16S rRNA sequences and joint species distribution modelling revealed that host genotypes and genetic differentiation among host populations together explained more than 50% of microbial community variation across our sampling transect. The ranked importance (most to least) of factors contributing to gut microbial diversity in our study populations were: genome-wide population differentiation, local environmental conditions, and host genotypes. However, differences in microbial communities among vole populations (β-diversity) did not follow patterns of lineage divergence (i.e., phylosymbiosis). Instead, among-population variation was best explained by the spatial distribution of hosts, as expected if the environment is a primary source of gut microbial diversity (i.e., dispersal limitation hypothesis). Across the contact zone, several bacterial taxa differed in relative abundance between the two parental lineages as well as among individuals with mismatched mitochondrial and nuclear genomes. Thus, genetic divergence among host lineages and mitonuclear genomic mismatches may also contribute to microbial diversity by altering interactions between host genomes and gut microbiota (i.e., hologenome speciation hypothesis).

Genetic variation of avian malaria in the tropical Andes: a relationship with the spatial distribution of hosts

BackgroundAvian haemosporidia are obligate blood parasites with an ample range of hosts worldwide. To understand how host communities may influence the diversity of parasites of the neotropics, the spatial genetic variation of avian Plasmodium, Haemoproteus, and Leucocytozoon was examined between areas of host endemism and along the elevational gradient in the tropical Andes.MethodsA total of 1686 accessions of the cytochrome b gene of avian haemosporidia were selected from 43 publications, that further provides additional information on 14.2% of bird species in the Neotropics. Haplotype groups were identified using a similarity-based clustering of sequences using a cut-off level ≥ 99.3% of sequence identity. Phylogenetic-based analyses were implemented to examine the spatial genetic structure of avian haemosporidia among areas of host endemism and the elevation gradient in the tropical Andes.ResultsThe areas of avian endemism, including the tropical Andes, can explain the differential distribution of the haemosporidia cytochrome b gene variation. In the tropical Andes region, the total number of avian haemosporidia haplotypes follows a unimodal pattern that peaks at mid-elevation between 2000 and 2500 m above sea level. Furthermore, the haplotype assemblages of obligate blood parasites tend to overlap towards mid-elevation, where avian host diversity tends to be maximized.ConclusionsSpatial analyses revealed that richness and turnover in haemosporidia suggest an association with montane host diversity, according to elevation in the tropical Andes. In addition, the spatial distribution of haemosporidia diversity is closely associated with patterns of host assemblages over large geographical scale in the tropical Andes and areas of avian endemism nearby.

Host density predicts the probability of parasitism by avian brood parasites.

The spatial distribution of hosts can be a determining factor in the reproductive success of parasites. Highly aggregated hosts may offer more opportunities for reproduction but can have better defences than isolated hosts. Here we connect macro- and micro-evolutionary processes to understand the link between host density and parasitism, using avian brood parasites as a model system. We analyse data across more than 200 host species using phylogenetic comparative analyses and quantify parasitism rate and host reproductive success in relation to spatial distribution using field data collected on one host species over 6 years. Our comparative analysis reveals that hosts occurring at intermediate densities are more likely to be parasitized than colonial or widely dispersed hosts. Correspondingly, our intraspecific field data show that individuals living at moderate densities experience higher parasitism rates than individuals at either low or high densities. Moreover, we show for the first time that the effect of host density on host reproductive success varies according to the intensity of parasitism; hosts have greater reproductive success when living at high densities if parasitism rates are high, but fare better at low densities when parasitism rates are low. We provide the first evidence of the trade-off between host density and parasitism at both macro- and micro-evolutionary scales in brood parasites. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

When parasitoids deal with the spatial distribution of their hosts: consequences for both partners.

Insect parasitoids developing inside hosts face a true challenge: hosts are scattered in the field and their localization and selection require the use of complex and sometime confusing information. It was assumed for a long time that small-brained organisms like parasitoids have evolved simple and efficient behavioral mechanisms, leading them to be adapted to a given ecological situation, for example, the spatial distribution of hosts in the habitat. However, hosts are not static and their distribution may also vary through generations and within the life of parasitoid individuals. We investigated if and how parasitoids deal with such a spatial complexity in a mesocosm experiment. We used the Aphidius rhopalosiphi/Sitobion avenae parasitoid/host system to investigate if parasitoid females experiencing different host aggregation levels exhibit different foraging behaviors independently of the number of hosts in the environment. We showed that A. rhopalosiphi females exploited hosts more intensively both within and among patches at higher host aggregation levels. We discussed the adaptiveness of such behaviors in the light of evolution and biological control.

Epidemix—An interactive multi-model application for teaching and visualizing infectious disease transmission

Mathematical models of disease transmission are used to improve our understanding of patterns of infection and to identify factors influencing them. During recent public and animal health crises, such as pandemic influenza, Ebola, Zika, foot-and-mouth disease, models have made important contributions in addressing policy questions, especially through the assessment of the trajectory and scale of outbreaks, and the evaluation of control interventions. However, their mathematical formulation means that they may appear as a “black box” to those without the appropriate mathematical background. This may lead to a negative perception of their utility for guiding policy, and generate expectations, which are not in line with what these models can deliver. It is therefore important for policymakers, as well as public health and animal health professionals and researchers who collaborate with modelers and use results generated by these models for policy development or research purpose, to understand the key concepts and assumptions underlying these models.The software application epidemix (http://shinyapps.rvc.ac.uk) presented here aims to make mathematical models of disease transmission accessible to a wider audience of users. By developing a visual interface for a suite of eight models, users can develop an understanding of the impact of various modelling assumptions – especially mixing patterns – on the trajectory of an epidemic and the impact of control interventions, without having to directly deal with the complexity of mathematical equations and programming languages. Models are compartmental or individual-based, deterministic or stochastic, and assume homogeneous or heterogeneous-mixing patterns (with the probability of transmission depending on the underlying structure of contact networks, or the spatial distribution of hosts). This application is intended to be used by scientists teaching mathematical modelling short courses to non-specialists – including policy makers, public and animal health professionals and students – and wishing to develop hands-on practicals illustrating key concepts of disease dynamics and control.

Linking parasite populations in hosts to parasite populations in space through Taylor's law and the negative binomial distribution

The spatial distribution of individuals of any species is a basic concern of ecology. The spatial distribution of parasites matters to control and conservation of parasites that affect human and nonhuman populations. This paper develops a quantitative theory to predict the spatial distribution of parasites based on the distribution of parasites in hosts and the spatial distribution of hosts. Four models are tested against observations of metazoan hosts and their parasites in littoral zones of four lakes in Otago, New Zealand. These models differ in two dichotomous assumptions, constituting a 2 × 2 theoretical design. One assumption specifies whether the variance function of the number of parasites per host individual is described by Taylor's law (TL) or the negative binomial distribution (NBD). The other assumption specifies whether the numbers of parasite individuals within each host in a square meter of habitat are independent or perfectly correlated among host individuals. We find empirically that the variance-mean relationship of the numbers of parasites per square meter is very well described by TL but is not well described by NBD. Two models that posit perfect correlation of the parasite loads of hosts in a square meter of habitat approximate observations much better than two models that posit independence of parasite loads of hosts in a square meter, regardless of whether the variance-mean relationship of parasites per host individual obeys TL or NBD. We infer that high local interhost correlations in parasite load strongly influence the spatial distribution of parasites. Local hotspots could influence control and conservation of parasites.

Biotic factors associated with the spatial distribution of the mistletoe Psittacanthus calyculatus in a tropical deciduous forest of central Mexico

Psittacanthus calyculatus (DC.) G. Don (Loranthaceae) is a neotropical hemiparasitic mistletoe and one of the most abundant and widely distributed species of the genus. Like most mistletoes, the interaction with its hosts and dispersers are directly related with its distribution and community dynamics. For the last decade, the infested areas in central México have increased, becoming a potential plant health problem. The aim of this study was to investigate three main biotic factors associated with the distribution of P. calyculatus in a tropical deciduous forest: (1) the plant community structure, (2) the host spatial distribution and level of association with P. calyculatus , and (3) bird consumers. Modified Whittaker plots were used to analyze the structure and composition of the plant community, comparing a highly infected mistletoe sites with non-infected sites. The spatial distribution and association of P. calyculatus with its most frequent hosts were calculated through a spatial analysis by distance indices. Study sites differed in plant diversity and species composition, showing a positive relation of mistletoe abundance with low diversity sites. The distance indices analysis presented a clumped distribution of P. calyculatus and Acacia hosts, as well as a high association with leguminous species such as A. farnesiana (X = 0.31, P = 0.0041) and A. schaffneri (X = 0.3977, P = 0.001). Three species consumed P. calyculatus’ berries: the Northern mockingbird, the Silky-Flycatcher and the Scott’s Oriole.

Penetration of the Amur form of Lernaea elegans (Crustacea: Lernaeidae) into the Mongolian part of the Selenga River basin and its host-spatial distribution

For the first time, the parasitic copepods Lernaea elegans are registered in the fish of the Mongolian part of the Selenga River basin. They are detected at the cyclopoid-copepodid and adult stages of development. The main hosts of copepods are Cyprinus carpio haematopterus, Esox lucius, Leuciscus leuciscus, and Perca fluviatilis. The highest levels of infection with L. elegans are marked in these fish in the lower reach of the Orkhon River. The probable donor of the alien species of L. elegans is the Amur carp, which self-spread to Mongolia from the Russian part of the Selenga River.

Predators, host abundance, and host spatial distribution affect the movement of wingless non-colonizing vector Rhopalosiphum padi (L.) and PVY prevalence in an oat/potato system

In the study of insect-vectored plant viruses, colonizing vector species remain the focus. However, non-colonizing vector species, those that do not settle and reproduce on the viral plant host, are often the most abundant in the field and may be the largest contributors to disease spread. While non-colonists may have a substantial effect on disease prevalence, the factors influencing their movement and transmission on non-host plants have been little studied. Here we evaluated how a common biological control agent (Hippodamia convergens), host and non-host plant abundance, and plant spatial distribution impact the movement and density of a wingless non-colonizing vector [Rhopalosiphum padi (L.)] and transmission of potato virus Y (PVY) in potatoes in experimental arenas. The results of this work illustrate the importance of plant species function (host or non-host) and distribution to vector behavior and disease spread. Predation, host plant abundance, and plant spatial distribution interactively affected viral prevalence within infected arenas. Increasing the number of vector non-host plants increased the distance and frequency of aphid movement, and the effect was influenced by plant spatial distribution, the arrangement of plant species in the experimental arena. Increasing the number of vector host plants increased the density of aphids. Although the interaction of the plant and predator treatments affected the proportion of potato plants infected in arenas where infection occurred, and host abundance and spatial distribution impacted vector movement and viral prevalence, aphid movement did not appear to mediate the effect of plant and predator treatments on PVY prevalence. This work demonstrates that both wingless non-colonizing vector behavior and transmission are aggregated responses to multiple environmental drivers.

Effectiveness of Erythrina gall wasp biocontrol and implications for the recovery of threatened Wiliwili trees (Fabaceae: Erythrina sandwicensis)1

Abstract Wiliwili (Erythrina sandwicensis), an endemic Hawaiian dry forest tree species, is threatened by an invasive gall-forming wasp (Quadrastichus erythrinae) first detected in Hawai‘i in 2005. Eurytoma erythrinae, a predator of Q. erythrinae from Tanzania, was selected as a biological control agent and was released at sites throughout the Hawaiian Islands. To assess the effectiveness of this biocontrol release, we measured the extent of damage attributable to Q. erythrinae wasps in 124 E. sandwicensis trees at the Waikoloa Village Dry Forest Recovery Project before (January, 2009) and after (January, 2011) the release of the parasitoid wasp at this site. Because host spatial distribution has important consequences for the establishment and success of natural enemies such as parasitoids, we took a spatially explicit approach to account for the potential spatial heterogeneity in E. sandwicensis. We observed an increase in galling damage across the population and 21 trees became completely defoliated be...

Manipulative parasites may not alter intermediate host distribution but still enhance their transmission: field evidence for increased vulnerability to definitive hosts and non-host predator avoidance

Behavioural alterations induced by parasites in their intermediate hosts can spatially structure host populations, possibly resulting in enhanced trophic transmission to definitive hosts. However, such alterations may also increase intermediate host vulnerability to non-host predators. Parasite-induced behavioural alterations may thus vary between parasite species and depend on each parasite definitive host species. We studied the influence of infection with 2 acanthocephalan parasites (Echinorhynchus truttae and Polymorphus minutus) on the distribution of the amphipod Gammarus pulex in the field. Predator presence or absence and predator species, whether suitable definitive host or dead-end predator, had no effect on the micro-distribution of infected or uninfected G. pulex amphipods. Although neither parasite species seem to influence intermediate host distribution, E. truttae infected G. pulex were still significantly more vulnerable to predation by fish (Cottus gobio), the parasite's definitive hosts. In contrast, G. pulex infected with P. minutus, a bird acanthocephalan, did not suffer from increased predation by C. gobio, a predator unsuitable as host for P. minutus. These results suggest that effects of behavioural changes associated with parasite infections might not be detectable until intermediate hosts actually come in contact with predators. However, parasite-induced changes in host spatial distribution may still be adaptive if they drive hosts into areas of high transmission probabilities.

Effects of mixed host spatial distribution on the efficiency of blue phosphorescent organic light-emitting diodes

We present a systematic study of how graded-host profile in the emissive layer (EML) affects the efficiency of blue phosphorescent organic light-emitting diodes. Three sets of devices with uniformly mixed and linearly graded mixed (LGM) hosts are fabricated with iridium-(III)-bis(4,6-difluorophenyl-pyridinato-N,C2) picolinate as dopant. By varying the position of the dopant within the emissive layer, we show that the LGM architecture improves device efficiency by moving the recombination zone further from the interface thereby reducing quenching due to interactions with charges in the transport layers.

How temperature and habitat quality affect parasitoid lifetime reproductive success—A simulation study

Adult parasitoid females lay their eggs in or on host insects. Most species are incapable of de novo lipogenesis as adults, and lipids accumulated during the larval stage are allocated either to egg production or to adult survival. Lipid consumption increases with distance covered by the parasitoids and thus with the distance between available hosts within a habitat. Temperature should affect parasitoid fitness because it changes the constraint imposed by a limited reserve of lipids and because it influences behaviour. Climate change involves both an increase in average temperature and an increased frequency of extreme weather such as heat waves. We investigated how the predicted increase of temperature will affect parasitoid fitness and how this depends on habitat parameters (spatial distribution of hosts and lipid cost of habitat exploitation). We studied optimal behaviour and calculated fitness at different temperatures and in different habitats using a stochastic dynamic programming model for pro-ovigenic parasitoids (which mature all their eggs before becoming adult). We show that an increase in temperature decreases fitness of parasitoids adapted to lower temperatures. This decrease in fitness depends on habitat quality. In field conditions (assuming small costs of intra-patch foraging), the loss of fitness should be larger in habitats with high inter-patch distance and in habitats with a more aggregated distribution of hosts. The foraging behaviour of parasitoids is also affected; at higher temperature we show that intra-patch foraging becomes less efficient, and patch residence times are longer.

Egg dispersion is more important than competition type for herbivores attacked by a parasitoid

Herbivore fitness can be altered by a combination of interacting organisms, such as its food plant, conspecifics, and predators/parasitoids. Here, we tested relative effects of plant species, herbivore intraspecific competition type, and spatial distribution of the herbivore among plant units on herbivore survival and whether parasitoids modified these effects. We used an endophagous bruchine seed predator Callosobruchusmaculatus for the herbivore, and a braconid wasp Heterospilusprosopidis for the parasitoid. The survival rate of C. maculatus was measured for each of 16 combinations of two plants (bean species, Vigna unguiculata and V. radiata), two competition types of C. maculatus larvae (contest and scramble), two spatial distributions of hosts [sparse (1 C. maculatus larva per seed over 20 seeds) and dense (2 C. maculatus larvae per seed over ten seeds)], and with/without a parasitoid pair. In the absence of the parasitoid, C. maculatus survival rate was lower with V. radiata and in the contest type. With the parasitoid, the proportion parasitized hosts was independent of total host density. Neither the proportion of parasitized hosts nor host survival rate was affected by plant species or host strain, but they were affected by host spatial distribution. When host distribution was dense, a higher proportion of hosts were parasitized, and C. maculatus survival rate was lower. Here we discuss parasitoid potential as a selective agent for the sparse within-pod distribution of its hosts in the field.

Influence of foraging behavior and host spatial distribution on the localized spread of the emerald ash borer, Agrilus planipennis

AbstractManagement programs for invasive species are often developed at a regional or national level, but physical intervention generally takes place over relatively small areas occupied by newly founded, isolated populations. The ability to predict how local habitat variation affects the expansion of such newly founded populations is essential for efficiently targeting resources to slow the spread of an invasive species. We assembled a coupled map lattice model that simulates the local spread of newly founded colonies of the emerald ash borer (Agrilus planipennis Fairmaire), a devastating forest insect pest of ash (Fraxinus spp.) trees. Using this model, we investigated the spread of A. planipennis in environments with different Fraxinus spp. distributions, and explored the consequences of ovipositional foraging behavior on the local spread of A. planipennis. Simulations indicate that increased larval density, resulting from lower host tree density or higher initial population sizes, can increase the spread rate during the first few years after colonization by increasing a density‐dependent developmental rate and via host resource depletion. Both the radial spread rate and population size were greatly influenced by ovipositional foraging behavior. Two known behaviors of ovipositing A. planipennis females, attraction towards areas with high ash tree density and attraction to stressed trees, had opposing effects on spread. Results from this model illustrate the significant influence of resource distribution and foraging behavior on localized spread, and the importance of these factors when formulating strategies to monitor and manage invasive pests.

Estimation of dispersal distances of the obligately plant-associated ant Crematogaster decamera

1. In obligate symbioses with horizontal transmission, the population dynamics of the partner organisms are highly interdependent. Host population size limits symbiont number, and distribution of partners is restricted by the presence and thus dispersal abilities of their respective partner. The Crematogaster decamera–Macaranga hypoleuca ant–plant symbiosis is obligate for both partners. Host survival depends on colonisation by its ant partner while foundress queens require hosts for colony establishment. 2. An experimental approach and population genetic analyses were combined to estimate dispersal distances of foundresses in their natural habitat in a Bornean primary rainforest. 3. Colonisation frequency was significantly negatively correlated with distance to potential reproductive colonies. Results were similar for seedlings at natural densities as well as for seedlings brought out in the area experimentally. Population genetic analysis revealed significant population differentiation with an FST of 0.041 among foundresses (n = 157) located at maximum 2280 m apart. In genetic spatial autocorrelation, genotypes of foundresses were significantly more similar than expected at random below 550 m and less similar above 620 m. Direct estimation of dispersal distances by pedigree analysis yielded an average dispersal distance of 468 m (maximum 1103 m). 4. For ants that disperse on the wing, genetic differentiation at such small spatial scales is unusual. The specific nesting requirements of the queens and the necessity for queens to find a host quickly could lead to colonisation of the first suitable seedling encountered, promoting short dispersal distances. Nonetheless, dispersal distances of C. decamera queens may vary with habitat or host spatial distribution.

Spatio‐temporal population genetic structure of the parasitic mite Spinturnix bechsteini is shaped by its own demography and the social system of its bat host

Information about the population genetic structures of parasites is important for an understanding of parasite transmission pathways and ultimately the co-evolution with their hosts. If parasites cannot disperse independently of their hosts, a parasite's population structure will depend upon the host's spatial distribution. Geographical barriers affecting host dispersal can therefore lead to structured parasite populations. However, how the host's social system affects the genetic structure of parasite populations is largely unknown. We used mitochondrial DNA (mtDNA) to describe the spatio-temporal population structure of a contact-transmitted parasitic wing mite (Spinturnix bechsteini) and compared it to that of its social host, the Bechstein's bat (Myotis bechsteinii). We observed no genetic differentiation between mites living on different bats within a colony. This suggests that mites can move freely among bats of the same colony. As expected in case of restricted inter-colony dispersal, we observed a strong genetic differentiation of mites among demographically isolated bat colonies. In contrast, we found a strong genetic turnover between years when we investigated the temporal variation of mite haplotypes within colonies. This can be explained with mite dispersal occuring between colonies and bottlenecks of mite populations within colonies. The observed absence of isolation by distance could be the result from genetic drift and/or from mites dispersing even between remote bat colonies, whose members may meet at mating sites in autumn or in hibernacula in winter. Our data show that the population structure of this parasitic wing mite is influenced by its own demography and the peculiar social system of its bat host.