Parasites and invasions: a biogeographic examination of parasites and hosts in native and introduced ranges

Aim To use a comparative approach to understand parasite demographic patterns in native versus introduced populations, evaluating the potential roles of host invasion history and parasite life history. Location North American east and west coasts with a focus on San Francisco Bay (SFB). Methods Species richness and prevalence of trematode parasites were examined in the native and introduced ranges of two gastropod host species, Ilyanassa obsoleta and Littorina saxatilis. We divided the native range into the putative source area for introduction and areas to the north and south; we also sampled the overlapping introduced range in SFB. We dissected 14,781 snails from 103 populations and recorded the prevalence and identity of trematode parasites. We compared trematode species richness and prevalence across the hosts’ introduced and native ranges, and evaluated the influence of host availability on observed patterns. Results Relative to the native range, both I. obsoleta and L. saxatilis have escaped (lost) parasites in SFB, and L. saxatilis demonstrated a greater reduction of trematode diversity and infection prevalence than I. obsoleta. This was not due to sampling inequalities between the hosts. Instead, rarefaction curves suggested complete capture of trematode species in native source and SFB subregions, except for L. saxatilis in SFB, where infection was extremely rare. For I. obsoleta, infection prevalence of trematodes using fish definitive hosts was significantly lower in SFB compared to the native range, unlike those using bird hosts. Host availability partly explained the presence of introduced trematodes in SFB. Main conclusions Differential losses of parasite richness and prevalence for the two gastropod host species in their introduced range is probably the result of several mechanistic factors: time since introduction, propagule pressure, vector of introduction, and host availability. Moreover, the recent occurrence of L. saxatilis’ invasion and its active introduction vector suggest that its parasite diversity and distribution will probably increase over time. Our study suggests that host invasion history and parasite life history play key roles in the extent and diversity of trematodes transferred to introduced populations. Our results also provide vital information for understanding community‐level influences of parasite introductions, as well as for disease ecology in general.     

Ecologically heterogeneous populations of the invasive ant Wasmannia auropunctata within its native and introduced ranges

Abstract 1. The biology of most invasive species in their native geographical areas remains largely unknown. Such studies are, however, crucial in shedding light on the ecological and evolutionary processes underlying biological invasions. 2. The present study focuses on the little fire ant Wasmannia auropunctata, a species native to Central and South America that has been widely introduced and which has become invasive throughout the tropics. We characterise and compare several ecological traits of native populations in French Guiana with those in one of its introduced ranges, New Caledonia. 3. We found ecologically heterogeneous populations of W. auropunctata coexisting in the species’ native geographical area. First, we found populations restricted to naturally perturbed areas (particularly floodplains) within the primary forest, and absent from the surrounding forest areas. These populations were characterised by low nest and worker densities. Second, we found dominant populations in recent anthropogenic areas (e.g. secondary forest or forest edge along road) characterised by high nest and worker densities, and associated with low ant species richness. The local dominance of W. auropunctata in such areas can be due to the displacement of other species (cause) or the filling‐up of empty habitats unsuitable to other ants (effect). With respect to their demographic features and ant species richness, the populations of native anthropogenic habitats were to a large extent similar to the invasive populations introduced into remote areas. 4. The results point to the need for greater research efforts to better understand the ecological and demographic features of invasive species within their native ranges.     

Predicting the ecological impacts of a new freshwater invader: functional responses and prey selectivity of the ‘killer shrimp’, Dikerogammarus villosus,compared to the native Gammarus pulex

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Native and invasive hosts play different roles in host–parasite networks

Parasites are often key players in biological invasions since they can mediate the impact of host invasions or can themselves become invasive species. However, the nature and extent of parasite-mediated invasions are often difficult to delineate. Here, we used individual-based, weighted bipartite networks to study the roles (degrees of interactions of individuals in a modular network according to their within- and among-module connections) played by native and invasive host individuals to their parasite communities. We studied two phylogenetically and ecologically close fish species, Mugil cephalus s.l. and Planiliza haematocheilus (Teleostei: Mugilidae). Planiliza haematocheilus is native to the Sea of Japan and invasive in the Sea of Azov whereas, M. cephalus s.l. is native to both seas. Based on the common evolutionary history that drives native host–parasite networks, we hypothesised that 1) native networks have higher modularity than invaded ones; and 2) invasive hosts in the invaded area play a peripheral role to structure parasite communities. We analysed the whole parasite community and subsets based on transmission strategy and host specificity of the parasite species to establish whether modularity and host roles are related to these features in the native and invaded areas. All networks were found to be modular. However, modularity tended to be higher in networks of the native area rather than those of the invaded area. Host individuals of both fish species played similar roles in the native area, whereas invasive hosts played a peripheral role in the networks of the invaded area. We propose that long-term monitoring of the roles of invasive hosts in parasite communities can be a useful proxy for estimating the maturity of the establishment of the invasive hosts in an ecosystem.     

Can a breakdown in competition–colonization tradeoffs help explain the success of exotic species in the California flora?

Determining combinations of functional traits that allow a species to colonize new habitats has been central in the development of invasion ecology. Species able to establish in new communities harbor abilities or traits that allow them to use resources or tolerate stress in ways that native species cannot. Tradeoffs among species functional traits along the competition–colonization (CC) continuum, where competitive ability is a decreasing function of dispersal capacity, may allow invasive species to establish themselves in new habitats. The California flora offers a well‐characterized model system to examine whether native and exotic species differ in the distribution of functional traits and to examine whether a breakdown of the CC tradeoff is present. We used a random subset of 1000 plants and examined seed traits and life form characteristics along with their seed size and adult height using the Jepson Manual of the plants of California. To test the hypothesis that active dispersal strategies aid in the success of exotic species, we classified species into four seed types according to the presence/absence of mechanisms associated with efficient dispersal. In addition, for each species we compiled data on seed size and adult plant height. We conducted all comparisons between native and exotic species within the four most speciose families to control for potential taxonomic non‐independence. Exotic species had smaller seed size but greater plant height than natives of the same families. On the other hand, exotic species also displayed significantly greater proportions of functional traits that enhanced dispersal ability. Additionally, certain sets of functional traits were significantly associated with exotic species, such as annual life histories with small seeds and high dispersal capacity. In the random subset of the California flora examined, exotics of the most speciose plant families show functional trait combinations that appear to violate the tradeoff structures observed in their California counterparts. Our results suggest that taxonomically controlled comparisons of the CC tradeoff structure between natives and exotic species may shed light of the capacity of those exotic species invasive ability to colonize new habitats.     

Evolutionary history of the little fire ant Wasmannia auropunctata before global invasion: inferring dispersal patterns,niche requirements and past and present distribution within its native range

The evolutionary history of invasive species within their native range may involve key processes that allow them to colonize new habitats. Therefore, phylogeographic studies of invasive species within their native ranges are useful to understand invasion biology in an evolutionary context. Here we integrated classical and Bayesian phylogeographic methods using mitochondrial and nuclear DNA markers with a palaeodistribution modelling approach, to infer the phylogeographic history of the invasive ant Wasmannia auropunctata across its native distribution in South America. We discuss our results in the context of the recent establishment of this mostly tropical species in the Mediterranean region. Our Bayesian phylogeographic analysis suggests that the common ancestor of the two main clades of W. auropunctata occurred in central Brazil during the Pliocene. Clade A would have differentiated northward and clade B southward, followed by a secondary contact beginning about 380 000 years ago in central South America. There were differences in the most suitable habitats among clades when considering three distinct climatic periods, suggesting that genetic differentiation was accompanied by changes in niche requirements, clade A being a tropical lineage and clade B a subtropical and temperate lineage. Only clade B reached more southern latitudes, with a colder climate than that of northern South America. This is concordant with the adaptation of this originally tropical ant species to temperate climates prior to its successful establishment in the Mediterranean region. This study highlights the usefulness of exploring the evolutionary history of invasive species within their native ranges to better understand biological invasions.     

Effects of eutrophication and snails on Eurasian watermilfoil (Myriophyllum spicatum) invasion

Exotic species can invade and establish new habitats both as a result of their own traits, and as a result of the characteristics of the environment they invade. Here, we show that the abundance of the invasive submerged aquatic plant, Myriophyllum spicatum (Eurasian watermilfoil) is highly dependent on the conditions of the environment in a mesocosm experiment. M. spicatum is allelopathic towards epiphytic algae, and in the absence of algivorous snails, we found that the abundance of both algae and M. spicatum significantly increased with experimentally increased nutrient loading, while the abundance of native submerged macrophytes declined. However, when snails were present, snail biomass increased with increasing nutrient loading, and M. spicatum biomass was consistently low while native submerged macrophyte biomass was consistently high. Our results stress the importance of the interaction between species traits and environmental conditions when considering the invasiveness of certain exotic species and the invasibility of certain environments.     

Differences in population dynamics and potential impacts of a freshwater invader driven by temporal habitat stability

Understanding population dynamics and population regulation of invasive species is critical for predicting their effects on native ecosystems as well as for control strategies. Many species of gastropod in the genus Pomacea are successful aquatic invaders that have caused economic and ecological impacts in Southeastern Asia where their large fecundity and broad reproductive window helps them to colonize and take advantages of ephemeral agricultural habitats. We followed the population dynamics of P. insularum in permanent, stable freshwater systems (ponds and streams), and in ephemeral agricultural habitats in the upper Texas Gulf Coast region, USA. We found that although P. insularum has a large reproductive potential, its density, biomass and size structure in stable permanent systems did not change significantly from March to November, and densities averaged <2 m⁻². This same species, however, displayed very different population dynamics in ephemeral agricultural environments. We found high densities (>130 m⁻²), and no stable size structure through time. Differences in the stability and persistence of these two types of environments appear to drive these patterns. Stability and persistence of habitats can result in different predator communities and the risk of predation for snails. We suggest that such factors may cause the differences in population dynamics and structure observed. The ability of snails to escape population control and explode in ephemeral habitats could drive the types of impacts seen on agricultural crops.     

Phenotypic traits of the Mediterranean <Emphasis Type="Italic">Phragmites australis</Emphasis> M1 lineage: differences between the native and introduced ranges

The environmental conditions in the new ranges of introduced plant species are often different from the conditions in their native ranges, and invasive plant species have been assumed to adapt to different environmental conditions by rapid ecological evolution in the invasive range after the introduction. Another interpretation of the change in plant traits after their introduction, however, is ecological fitting, which is based on the inherently high phenotypic plasticity of the species rather than on evolution. The Mediterranean haplotype M1 lineage of the wetland grass Phragmites australis was introduced to the coastal wetlands along the Gulf Coast of North America, where it is exposed to a different climate compared to its original range. The climate in the native range is arid or temperate with dry and hot summers, whereas the climate in the introduced range is warmer and has a higher and more uniform precipitation than that in the native range. This warmer and more humid environment is likely to pose different selection pressures to the plants in the introduced range and thus cause rapid evolutionary change and phenotypic differentiation in the introduced range. Here, we compared phenotypic traits of the M1 lineage from the native and introduced ranges in a common garden experiment to study the processes assisting the successful spread in the introduced range. Overall, the native and introduced groups were similar, but we detected a few phenotypic traits that diverged. Ecological fitting could be the fundamental mechanism by which the P. australis M1 lineage survives and spreads in the introduced Gulf Coast region. However, further research is needed to assess how the diverging traits observed in our study in Denmark (lower photosynthetic rates, lower chlorophylls concentration and higher leaf K concentration for the introduced than for the native genotypes) are expressed in the two ranges.     

Fruit type,life form and origin determine the success of woody plant invaders in an urban landscape

The spread of alien plant species is a critical ecological event worldwide, but the forces that control this spread are not well documented. Alien plant species are well known to disrupt ecological services of native ecosystems, change the composition of native habitats, and often lead to the extirpation of native flora and fauna. Here, we report on life history patterns of plant species with rapidly spreading and declining ranges in North America’s major urban region. We tested for differences in life history traits between the 466 native and alien woody flora of the New York metropolitan area. We also examined the relationship between life history traits and change in distribution in the New York metropolitan area between 1900 and 2000. Native and alien species of the New York metropolitan area differ with respect to pollination vector and breeding system. However, pollination vector and breeding system are not associated with success, defined here as increasing range spread in the urban environment; instead, fruit type (dispersal), life form and origin are important determinants of success. Alien species that are deciduous trees, shrubs or vines with fleshy fruit are the most successful in increasing their distribution in this urban landscape. Newly introduced species with these characteristics are expected to have a better chance at establishing in similar urban landscapes and should be targets for intensive management. The ability to predict which alien species will become invasive is also a valuable tool for the prevention of invasions by newly introduced plant species.     

Host traits associated with species roles in parasite sharing networks

The community of host species that a parasite infects is often explained by functional traits and phylogeny, predicting that closely related hosts or those with particular traits share more parasites with other hosts. Previous research has examined parasite community similarity by regressing pairwise parasite community dissimilarity between two host species against host phylogenetic distance. However, pairwise approaches cannot target specific host species responsible for disproportionate levels of parasite sharing. To better identify why some host species contribute differentially to parasite diversity patterns, we represent parasite sharing using ecological networks consisting of host species connected by instances of shared parasitism. These networks can help identify host species and traits associated with high levels of parasite sharing that may subsequently identify important hosts for parasite maintenance and transmission within communities. We used global‐scale parasite sharing networks of ungulates, carnivores, and primates to determine if host importance – encapsulated by the network measures degree, closeness, betweenness, and eigenvector centrality – was predictable based on host traits. Our findings suggest that host centrality in parasite sharing networks is a function of host population density and range size, with range size reflecting both species geographic range and the home range of those species. In the full network, host taxonomic family became an important predictor of centrality, suggesting a role for evolutionary relationships between host and parasite species. More broadly, these findings show that trait data predict key properties of ecological networks, thus highlighting a role for species traits in understanding network assembly, stability, and structure.     

A phylogenetic comparative study of preadaptation for invasiveness in the genus <Emphasis Type="Italic">Silene</Emphasis> (Caryophyllaceae)

The role of preadaptation in ecology and evolution is determined by how the traits evolved by a species in one environment allow it to be successful in novel environments. This concept bears directly on modern biological invasions, as species are introduced to new locations beyond their historical borders. In this study, we used a phylogenetically-controlled analysis of the flowering plant genus Silene (Caryophyllaceae) to show that native geographic range size, along with a suite of life history traits affecting plant growth and reproduction, have preadapted some species for the invasion of new ranges. Using a path analytic approach, we further show that some of the covariance between life history traits and invasiveness is indirect, caused by mutual associations with native range size. Specifically, we found that reproductive traits such as the number of flowers per inflorescence and length of the flowering season directly preadapt species for invasion, while plant height is indirectly associated with invasion through a correlation with native range size. Other traits such as ovule number and leaf size are both directly and indirectly associated with invasion success. Our results reveal the importance of accounting for correlations among plant traits and geographic range size when predicting preadaptation for invasiveness. We also highlight that the traits predictive of invasion success among species of Silene are often those found to be rapidly evolving within introduced populations, suggesting common forces of selection operating at these different biological scales of organization during invasion.     

温度对福寿螺生物学特性的影响及高低温适应机制研究进展

在全球气候变化背景下, 福寿螺在我国及全球进一步扩散。极强的生态耐受力及快速适应力是福寿螺能够在入侵地区迅速扩散的重要原因。其中, 环境温度对福寿螺的生存、生长、发育及繁殖至关重要, 是影响福寿螺分布、扩散及暴发的重要因素之一。文章在综述温度耐受范围的基础上, 总结了福寿螺高低温适应的生理生化及分子机制, 并对从温度适应性角度揭示入侵机制的研究前景进行展望。当前, 福寿螺温度适应的生理生化机制研究主要针对化合物以及相关酶活性变化开展, 分子机制研究主要集中在HSP基因的表达差异上。在染色体水平基因组完成测序的基础上, 福寿螺快速适应性进化的生理生态耐受性机制和表型可塑性机制有待深入开展。     

Differences in trait compositions between rocky natural and artificial habitats

Question: What are the differences in trait compositions that enable native plants to colonise comparable natural and man‐made habitats? Are these traits independent of phylogenetic relationships between species? Location: Czech Republic. Methods: The relative importance of biological, ecological and distributional traits of native species was studied, using a dataset of 75 species growing in rock and wall habitats in the Czech Republic. Species preferences for individual habitats due to climatic conditions and proportions of different vegetation types in their surroundings were partialled out using partial canonical correspondence analysis. The pattern of plant traits along a gradient from natural rock habitats to secondary wall habitats was analysed using regression trees and generalized linear models with and without phylogenetic correction. Results: The most common native species colonising rock habitats are phanerophytes, mostly woody juveniles, with a CSR life strategy and most are adapted to epizoochory. Summer green leaves, annual life span, CR life strategy, reproduction mostly by seeds and dispersal by ants are all traits positively associated with the ability of species to colonise wall habitats. These species are also characterised by their high demand for nutrients, temperature, base‐rich substrates and light. Biological and ecological traits are more important for colonising new habitats than traits related to species dispersal ability or phylogenetic relationships between species. Biological and ecological traits alone explained 29.3% of variability in the species dataset, while dispersal characteristics and phylogeny alone explained 9.1% and 4.8%, respectively. Conclusions: We outline how the process of environmental filtering determines native species assemblages and identify a set of species traits that enable them to persist in particular habitats. We conclude that although urbanisation generally results in loss of natural habitats, there are new, man‐made habitats potentially suitable for native species.     

Invasive parasites are detectable by their abundance-occupancy relationships: the case of helminths from Liza haematocheilus (Teleostei: Mugilidae)

The biogeographic patterns of abundance and prevalence of helminths from Liza haematocheilus were studied across its native (Sea of Japan) and introduced (Sea of Azov) distribution ranges. Abundance-occupancy relationships (AORs) were tested for the core-satellite and enemy release (ERH) species hypotheses in eight and 14 host samples from the native and introduced host ranges, respectively. The AOR model fitted parasite data extremely well, irrespective of whether the host or the parasite species were native or invasive. Except for co-introduced monogeneans, species were less abundant and prevalent in the introduced host population than in the native one, which agrees well with the ERH. Two occupancy patterns were observed. A unimodal, right-skewed distribution of prevalence frequency was common for the acquired groups of helminth parasites in the introduced range, whereas a bimodal distribution was more common in the native range. Core species in the native range were monogeneans, adult and larval digeneans, whereas host-specific, co-introduced monogeneans were the only core species in the introduced range. Acquired grey-mullet specialists and host generalists infected only a small portion of the introduced host population with low mean abundance. These results indicate that strict host specificity, together with a direct life cycle, are the traits that enabled helminth species to entirely occupy the invasive host population. The AORs showed that parasite individuals tend to accumulate in a relatively small fraction of susceptible introduced hosts, probably as an adaptation to enhance mating opportunities, thereby providing a mechanistic explanation of the ERH. All this evidence suggests that co-introduced and acquired species use the introduced host population in very different ways. Therefore, we posit that the examination of AORs can be instrumental in understanding the role of co-introduced parasites in invasion theory.     

Ecological implications of parasites in natural <Emphasis Type="Italic">Daphnia</Emphasis> populations

In natural host populations, parasitism is considered to be omnipresent and to play an important role in shaping host life history and population dynamics. Here, we study parasitism in natural populations of the zooplankton host Daphnia magna investigating their individual and population level effects during a 2-year field study. Our results revealed a rich and highly prevalent community of parasites, with eight endoparasite species (four microsporidia, one amoeba, two bacteria and one nematode) and six epibionts (belonging to five different taxa: Chlorophyta, Bacillariophyceae, Ciliata, Fungi and Rotifera). Several of the endoparasites were associated with a severe overall fecundity reduction of the hosts, while such effects were not seen for epibionts. In particular, infections by Pasteuria ramosa, White Fat Cell Disease and Flabelliforma magnivora were strongly associated with a reduction in overall D. magna fecundity. Across the sampling period, average population fecundity of D. magna was negatively associated with overall infection intensity and total endoparasite richness. Population density of D. magna was negatively correlated to overall endoparasite prevalence and positively correlated with epibiont richness. Finally, the reduction in host fecundity caused by different parasite species was negatively correlated to both parasite prevalence and the length of the time period during which the parasite persisted in the host population. Consistent with epidemiological models, these results indicate that parasite mediated host damages influence the population dynamics of both hosts and parasites.     

The combined influence of trematode parasites and predatory salamanders on wood frog (<Emphasis Type="Italic">Rana sylvatica</Emphasis>) tadpoles

Predators can have important impacts on host–parasite dynamics. For many directly transmitted parasites, predators can reduce transmission by removing the most heavily infected individuals from the population. Less is known about how predators might influence parasite dynamics in systems where the parasite relies on vectors or multiple host species to complete their life cycles. Digenetic trematodes are parasitic flatworms with complex life cycles typically involving three host species. They are common parasites in freshwater systems containing aquatic snails, which serve as obligate first intermediate hosts, and multiple trematode species use amphibians as second intermediate hosts. We experimentally examined the impact of predatory salamanders (Ambystoma jeffersonianum) and trematode parasites (Echinostoma trivolvis and Ribeiroia ondatrae) on short-term survival of wood frog tadpoles (Rana sylvatica) in 150-L outdoor pools. Two trematode species were used in experiments because field surveys indicated the presence of both species at our primary study site. Parasites and predators both significantly reduced tadpole survival in outdoor pools; after 6 days, tadpole survival was reduced from 100% in control pools to a mean of 46% in pools containing just parasites and a mean of 49% in pools containing just predators. In pools containing both infected snails and predators, tadpole survival was further reduced to a mean of 5%, a clear risk-enhancement or synergism. These dramatic results suggest that predators may alter transmission dynamics of trematodes in natural systems, and that a complete understanding of host–parasite interactions requires studying these interactions within the ecological framework of community interactions.     

Distribution of non-native invasive species and soil properties in proximity to paved roads and unpaved roads in a quartzitic mountainous grassland of southeastern Brazil (rupestrian fields)

One of the most important disturbances of roads is the facilitation of the increase of non-native invasive species into adjacent plant communities. The rupestrian fields of Serra do Cipó, a montane grassland ecosystem in southeastern Brazil, are recognized for their enormous richness of species and endemism rates. The presence of non-native invasive species in this ecosystem could threaten the existence of the native flora and its associated organisms. The aim of this study is to understand how non-native invasive species and native species are distributed along paved and unpaved roads, in a montaneous grassland ecosystem such as the Brazilian rupestrian fields. The two road surfaces provide differing gradients from their edges with respect to nutrients, soil chemical aspects and plant species diversity. High content of calcium at the roadside in the paved road resulted from the paving process, in which limestone gravel is used in one of the several paving phases. In these newly created habitats the toxicity of aluminum is drastically reduced and nutrient enriched, hence representing favorable sites from where non-native invasive species are capable to colonize and grow for undetermined period waiting the chance to invade the adjacent pristine habitats. Disturbances provoked by any natural or human-caused event can provide the opportunity for the non-native invasive species to colonize new plant communities.     

Ecomorphology of a generalist freshwater gastropod: complex relations of shell morphology,habitat, and fecundity

Evolutionary and ecological situations in a species’ native and invasive ranges can be drastically different. This is the case for Potamopyrgus antipodarum Gray (1843) a morphologically highly variable freshwater snail native to New Zealand, where sexual and asexual individuals coexist and experience selective pressure by sterilizing endoparasites. By contrast, only a few asexual lineages have been established in invaded regions around the globe, where parasite infection is extremely rare. We analyzed the ecomorphology of 996 native P. antipodarum in a geometric morphometric framework, using brood size as proxy for fecundity, and mtDNA and nuclear SNPs to account for relatedness and identify reproductive mode. As expected, we found genetic and morphological diversity to be higher in native than in invasive snails investigated previously, but surprisingly no higher morphological diversity in sexual versus asexual individuals. The relationships between shell morphology, habitat, and fecundity were complex. Shape variation was primarily linked to genetic relatedness but specific environmental factors including flow rate induced similar shell shapes. By contrast, shell size was largely explained by environmental factors. Fecundity was correlated with size but showed trade-offs with shape in increasingly extreme conditions. With increasing flow and toward small springs, the trend of shell shape becoming wider was reversed, i.e., snails with narrower shells were brooding more embryos. We concluded that both genetic and environmental contributions to variation in shell morphology in P. antipodarum likely play an important role in the ability of this species to adapt to a wide spectrum of habitats.