Increasing Prevalence of Plant‐Fungal Symbiosis Across Two Centuries of Environmental Change

Bacterial assemblages on the skins of amphibians are known to influence pathogen resistance and other important physiological functions in the host. Host-specific factors and the environment play significant roles in structuring skin assemblages. This study used high-throughput 16S rRNA sequencing and multivariate analyses to examine differences in skin-bacterial assemblages from 246 salamanders belonging to three genera in the lungless family Plethodontidae along multiple spatial gradients. Composition and α- and β-diversity of bacterial assemblages were defined, indicator species were identified for each host group, and the relative influences of host- versus environment-specific ecological factors were evaluated. At the broadest spatial scale, host genus, host species, and sampling site were predictive of skin assemblage structure, but host genus and species were more influential after controlling for the marginal effects of site, as well as nestedness of site. Furthermore, assemblage similarity within each host genus did not change with increasing geographic distance. At the smallest spatial scale, site-specific climate analyses revealed different relationships to climatic variables for each of the three genera, and these relationships were determined by host ecomode. Variation in bacterial assemblages of terrestrial hosts correlated with landscape-level climatic variability, and this pattern decayed with increasing water dependence of the host. Results from this study highlight host-specific considerations for researchers studying wildlife diseases in co-occurring, yet ecologically divergent, species.

Summary 1. Broad-scale patterns of above-ground net primary production (ANPP) are closely coupled to climate features, particularly the distribution and magnitude of rainfall. In arid and semi-arid regions, however, the precipitation regime, together with local geomorphology and plant life history traits, combine to regulate soil water availability and patterns of growth, complicating simple correlations with climatic variables. 2. To better understand the drivers of plant growth in desert ecosystems, we characterized the rates and spatial heterogeneity of stem elongation by the dominant perennial shrub, creosote bush (Larrea tridentata) in the northern Sonoran Desert of Arizona (USA). Estimates of stem growth were made over a 5-year period (2006–2010) from 60 plots at 15 sites spanning c. 120 km across the Central Arizona–Phoenix (CAP) Long-term Ecological Research (LTER) area. 3. We observed both the highest and lowest rates of stem growth during summer, and these patterns were related to the amount of monsoon rainfall and local rates of water infiltration. The relationship between growth and precipitation in the summer was nonlinear, with rates increasing more than eightfold at plots receiving more than 100 mm of seasonal rainfall. Conversely, growth during the winter/spring was intermediate in magnitude, similar among years and poorly correlated with seasonal precipitation. 4. The spatial heterogeneity of stem growth also differed markedly between seasons and was greater both within and among sites during summer compared to winter/spring growing periods. At broad scales, spatial heterogeneity of shrub growth was correlated with seasonal changes in the spatial variability of rainfall across the study area. At small spatial scales, greater heterogeneity during the summer appears linked to local soil properties that influence infiltration and water availability following monsoon storms. 5. Overall, the strong, nonlinear growth response by L. tridentata to monsoon rainfall suggests that the recurrence interval of wet summer seasons is an important driver of ANPP for this long-lived shrub. More generally, our results illustrate how regional climate variability and local soil properties can interact to influence the rates and heterogeneity of desert plant growth at multiple scales.

Extensive research has documented the importance of social trust for economic development, yet the origins of trust remain largely unexplored. This paper examines the historical relationship between risk, cooperation and the emergence of social trust. I hypothesize that norms of trust developed in preindustrial times as a result of experiences of collective action and mutual insurance triggered by the need for subsistence farmers to cope with climatic risk. These norms persisted over time, even after climate had become largely unimportant for economic activity. I test this hypothesis in the context of Europe combining high-resolution climate data for the period 1500-2000 with contemporary survey data at the sub-national level. I find that regions characterized by higher year-to-year variability in precipitation and temperature display higher levels of trust. Consistent with a theory of insurance through geographic differentiation, I also find that trust is higher in regions with more spatially heterogeneous precipitation. Furthermore, variation in social trust is driven by weather patterns during the growing season and by historical rather than recent variability. These results are robust to the inclusion of country fixed-effects, a variety of geographical controls, and regional measures of early political and economic development.

1. We applied a Markov random fields/conditional random fields (MRF/CRF) modelling to understand whether and how species associations in ectoparasite infracommunities of small mammals vary along environmental gradients across space and time.2. We analysed the variation in species associations in flea and mite infracommunities of seven host species across localities (spatial scale) and time periods (in the same locality; temporal scale).3. The performance of the CRF models differed between ectoparasite taxa, scales, and host species. The model performance for flea and mite infracommunities in the majority of host species was satisfying although the proportions of correctly predicted positive occurrences and model sensitivity were moderate.4. The probability of occurrence of many flea and mite species depended mostly on the occurrence of another flea and mite species (respectively), with many negative interspecific interactions, although occurrences of some species were mainly affected by environmental factors. The patterns of pairwise associations between some ectoparasite species were affected by environmental factors.5. Although many ectoparasite pairs demonstrated stability in the strength or sign of association across space or time, interactions between other species pairs were spatially or temporally variable, with the association between the same pair of species being stable in one host species and variable in another host species.6. We conclude that ectoparasite associations are mediated by the off‐host environment and may depend on the life histories of both host and ectoparasite species.

Vertically transmitted microbes are common in macro‐organisms and can enhance host defense against environmental stress. Because vertical transmission couples host and symbiont lineages, symbionts may become specialized to host species or genotypes. Specialization and contrasting reproductive modes of symbiotic partners could create incompatibilities between inherited symbionts and novel host genotypes when hosts outcross or hybridize. Such incompatibilities could manifest as failed colonization or poor symbiont growth in host offspring that are genetically dissimilar from their maternal host. Moreover, outcrossing between host species could influence both host and symbiont reproductive performance. We tested these hypotheses by manipulating outcrossing between populations and species of two grasses, Elymus virginicus and E. canadensis, that host vertically transmitted fungal endophytes (genus Epichloё). In both greenhouse and field settings, we found that host–symbiont compatibility was robust to variation in host genetic background, spanning within‐population, between‐population and between‐species crosses. Symbiont transmission into the F1 generation was generally high and weakly affected by host outcrossing. Furthermore, endophytes grew equally well in planta regardless of host genetic background and transmitted at high frequencies into the F2 generation. However, outcrossing, especially inter‐specific hybridization, reduced reproductive fitness of the host, and thereby the symbiont. Our results challenge the hypothesis that host genetic recombination, which typically exceeds that of symbionts, is a disruptive force in heritable symbioses. Instead, symbionts may be sufficiently generalized to tolerate ecologically realistic variation in host outcrossing.

Heritable microbes are abundant in nature and influential to their hosts and the communities in which they reside. However, drivers of variability in the prevalence of heritable symbionts and their rates of transmission are poorly resolved, particularly across host populations experiencing variable biotic and abiotic environments. To fill these gaps, we surveyed 25 populations of two native grasses (Elymus virginicus and Elymus canadensis) across the southern Great Plains (USA). Both grass species host heritable endophytic fungi (genus Epichloё) and can hybridize where their ranges overlap. From a subset of hosts, we characterized endophyte genotype using genetic loci that link to bioactive alkaloid production. First, we found mean vertical transmission rates and population-level prevalence were positively correlated, specifically for E. virginicus. However, both endophyte prevalence and transmission varied substantially across populations and did not strongly correlate with abiotic variables, with one exception: endophyte prevalence decreased as drought stress decreased for E. virginicus hosts. Second, we evaluated the potential influence of biotic factors and found that, after accounting for climate, endophyte genotype explained significant variation in symbiont inheritance. We also contrasted populations where host species co-occurred in sympatry vs. allopatry. Sympatry could potentially increase interspecific hybridization, but this variable did not associate with patterns of symbiont prevalence or transmission success. Our results reveal substantial variability in symbiont prevalence and transmission across host populations and identify symbiont genotype, and to a lesser extent, the abiotic environment as sources of this variation.

The richness and structure of symbiont assemblages are shaped by many factors acting at different spatial and temporal scales. Among them, host phylogeny and geographic distance play essential roles. To explore drivers of richness and structure of symbiont assemblages, feather mites and seabirds are an attractive model due to their peculiar traits. Feather mites are permanent ectosymbionts and considered highly host-specific with limited dispersal abilities. Seabirds harbour species-rich feather mite communities and their colonial breeding provides opportunities for symbionts to exploit several host species. To unravel the richness and test the influence of host phylogeny and geographic distance on mite communities, we collected feather mites from 11 seabird species breeding across the Atlantic Ocean and Mediterranean Sea. Using morphological criteria, we identified 33 mite species, of which 17 were new or recently described species. Based on community similarity analyses, mite communities were clearly structured by host genera, while the effect of geography within host genera or species was weak and sometimes negligible. We found a weak but significant effect of geographic distance on similarity patterns in mite communities for Cory’s shearwaters Calonectris borealis. Feather mite specificity mainly occurred at the host-genus rather than at host-species level, suggesting that previously inferred host species-specificity may have resulted from poorly sampling closely related host species. Overall, our results show that host phylogeny plays a greater role than geography in determining the composition and structure of mite assemblages and pinpoints the importance of sampling mites from closely-related host species before describing mite specificity patterns.

Minimizing Data Waste: Conservation in the Big Data Era

Summary 1. Both evolutionary theory and empirical evidence from agricultural research support the view that asexual, vertically transmitted fungal endophytes are typically plant mutualists that develop high infection frequencies within host grass populations. In contrast, endophyte–grass interactions in natural ecosystems are more variable, spanning the range from mutualism to antagonism and comparatively little is known about their range of response to environmental stress. 2. We examined patterns in endophyte prevalence and endophyte–grass interactions across nutrient and grazing (from Greylag and Canada geese) gradients in 15 sites with different soil moisture levels in 13 island populations of the widespread grass Festuca rubra in a boreal archipelago in Sweden. 3. In the field, endophyte prevalence levels were generally low (range = 10–53%) compared with those reported from agricultural systems. Under mesic‐moist conditions endophyte prevalence was constantly low (mean prevalence = 15%) and was not affected by grazing pressure or nutrient availability. In contrast, under conditions of drought, endophyte prevalence increased from 10% to 53% with increasing nutrient availability and increasing grazing pressure. 4. In the field, we measured the production of flowering culms, as a proxy for host fitness, to determine how endophyte‐infected plants differed from uninfected plants. At dry sites, endophyte infection did not affect flowering culm production. In contrast, at mesic‐moist sites production of flowering culms in endophyte‐infected plants increased with the covarying effects of increasing nutrient availability and grazing pressure, indicating that the interaction switched from antagonistic to mutualistic. 5. A concurrent glasshouse experiment showed that in most situations, the host appears to incur some costs for harbouring endophytes. Uninfected grasses generally outperformed infected grasses (antagonistic interaction), while infected grasses outperformed uninfected grasses (mutualistic interaction) only in dry, nutrient‐rich conditions. Nutrient and water addition affected tiller production, leaf number and leaf length differently, suggesting that tillers responded with different strategies. This emphasizes that several response variables are needed to evaluate the interaction. 6. Synthesis. This study found complex patterns in endophyte prevalence that were not always correlated with culm production. These contrasting patterns suggest that the direction and strength of selection on infected plants is highly variable and depends upon a suite of interacting environmental variables that may fluctuate in the intensity of their impact, during the course of the host life cycle.

ABSTRACTPredictions that Brexit will precipitate the disintegration of the European Union (EU) have greatly overstated the prospect of such an event. Britain's centuries-long conflicted relationship with Europe and contemporary survey data – exhibiting marked differences between Britain and the rest of Europe with regard to Euroskepticism – suggest that Brexit is a sui generis event, unique to Britain, and is not a harbinger of things to come. Conversely, by triggering the departure of the EU's biggest obstacle to an ever deeper union, Brexit's most likely effect is to strengthen European integration and foster greater consolidation in the EU.

Sessile marine invertebrate (biofouling) communities have served as an important model in ecology for evaluating fundamental patterns and processes, including invasion dynamics, which vary at broad spatial and temporal scales. Here, we tested for differences in biofouling community development among three biogeographically distinct bays in North America (Chesapeake Bay, Tampa Bay, and San Francisco Bay), exploring possible explanations for differential non-native species success. In particular, we aimed to examine if San Francisco Bay (a global hotspot for non-native species richness) differed in composition and space occupation, especially since open space can facilitate colonization. Additionally, we explored how biogeography, assembly and succession dynamics over the short and long term, and space availability affect marine communities across broad spatial scales. Patterns of community assembly differed among bays, with more bare space and less secondary cover (species settling on species) in San Francisco Bay. San Francisco Bay was also distinguished by a higher percent cover of Tunicata (almost all of which are non-native and historically absent) over multiple time scales. Cirripedia recruited on bare panels in all three Bays but cover increased only in Tampa Bay, as soak time increased. Tube-dwelling Polychaeta distinguished Chesapeake Bay from Tampa Bay and San Francisco Bay. Low-salinity events temporarily restructured the communities in all three bays. Whether differences among bays reflect coastal versus bay-specific patterns remains to be tested.

Understanding how species assemble into communities is a central issue in community ecology. So far, most studies have focused on the assembly mechanisms of vascular plant communities, while the role of deterministic (environmental filtering and biotic interactions) and stochastic (e.g. dispersal limitation) processes structuring bryophyte assemblages remains poorly understood. To evaluate how different assembly processes shape bryophyte communities in mountain streams, we examined functional trait patterns across spatial scales and along environmental gradients. To do so, we sampled 754 microhabitat plots (0.25 m2) nested within 165 sites (100‐m long stream segments) and 13 sub‐basins, located in the northwest and central‐west of Portugal. At each spatial scale, observed functional diversity indices (Functional Richness and Rao's quadratic entropy) were compared to random expectations derived from null models, followed by the analysis of changes in functional trait patterns along environmental gradients by fitting a series of generalized additive mixed models. At local scales (site and microhabitat plot), coexisting species tended to be more functionally similar than expected, suggesting the prevalence of environmental filtering effects. In contrast, no significant deviations from random expectations were detected at the broadest spatial scale (sub‐basin), indicating the prevalence of stochastic processes. We found contrasting assembly processes along environmental gradients: environmental filtering prevailed in stressful environments, while competitive interactions were more important in favourable conditions. Synthesis. Our results highlight the role of environmental filtering in bryophyte community assembly at fine spatial scales, emphasizing the importance of measuring environmental conditions at the same spatial scales where biotic interactions take place. In line with the stress‐dominance hypothesis, the relative importance of environmental filtering increased with abiotic stress. Thus, analysing functional trait patterns across different spatial scales and environmental gradients may contribute to a better understanding of the mechanisms underlying community assembly.

Patterns of spatial genetic variation can be generated by a variety of ecological processes, including individual preferences based on habitat. These ecological processes act at multiple spatial and temporal scales, generating scale-dependent effects on gene flow. In this study, we focused on bobcats (Lynx rufus), a highly mobile, generalist felid that exhibits ecological and behavioral plasticity, high abundance, and broad connectivity across much of their range. However, bobcats also show genetic differentiation along habitat breaks, a pattern typically observed in cases of isolation-by-ecology (IBE). The IBE observed in bobcats is hypothesized to occur due to habitat-biased dispersal, but it is unknown if this occurs at other habitat breaks across their range or at what spatial scale IBE becomes most apparent. Thus, we used a multiscale approach to examine isolation by ecology (IBE) patterns in bobcats (Lynx rufus) at both fine and broad spatial scales in western Texas. We genotyped 102 individuals at nine microsatellite loci and used partial redundancy analysis (pRDA) to test if a suite of landscape variables influenced genetic variation in bobcats. Bobcats exhibited a latitudinal cline in population structure with a spatial signature of male-biased dispersal, and no clear barriers to gene flow. Our pRDA tests revealed high genetic similarity in similar habitats, and results differed by spatial scale. At the fine spatial scale, herbaceous rangeland was an important influence on gene flow whereas mixed rangeland and agriculture were significant at the broad spatial scale. Taken together, our results suggests that complex interactions between spatial-use behavior and landscape heterogeneity can create non-random gene flow in highly mobile species like bobcats. Furthermore, our results add to the growing body of data highlighting the importance of multiscale study designs when assessing spatial genetic structure.

The number of species coexisting in a community may be regulated by local factors (e.g., competitive interactions), or by regional processes (e.g., dispersal from a regional species pool). The relative importance of local and regional processes can be inferred from the shape of the relationship between local and regional species richness. We investigated this relationship in communities of fleas parasitic on small mammals at two spatial scales: between the richness of fleas on individual hosts (infracommunities) and that of fleas on host populations (component communities), and between the richness of component communities and that of the entire regional species pool. We tested linearity (proportional sampling) versus curvilinearity with an asymptote (species saturation) by plotting "local" against "regional" species richness of fleas either among host species or within host species among populations. At the two spatial scales, we found consistent curvilinear relationships between species richness of the more "local" communities and richness of the more "regional" communities. This was true across all host species in the data set and for geographic subsets, even after controlling for the influence of sampling effort on estimates of species richness, and that of host phylogeny in interspecific analyses. We also tested for density compensation in species-poor communities. There was no strong evidence for density compensation at the infracommunity level, although its existence at the component community level appeared likely. Our results suggest that identical patterns in local-versus-regional species richness observed on two different spatial scales arise via different mechanisms: infracommunities appear saturated with flea species most likely because of local processes, such as host immune defenses, whereas component communities are saturated with species through interspecific competition, possibly among larval stages.

Biodiversity can influence disease risk. One example of a diversity-disease relationship is the dilution effect, which suggests higher host species diversity (often indexed by species richness) reduces disease risk. While numerous studies support the dilution effect, its generality remains controversial. Most studies of diversity-disease relationships have overlooked the potential importance of phylogenetic diversity. Furthermore, most studies have tested diversity-disease relationships at one spatial scale, even though such relationships are likely scale dependent. Using Lyme disease as a model system, we investigated the effects of host species richness and phylogenetic relatedness on the number of reported Lyme disease cases in humans in the U.S.A. at two spatial scales (the county level and the state level) using piecewise structural equation modelling. We also accounted for relevant climatic and habitat-related factors and tested their correlations with the number of Lyme disease cases. We found that species assemblages with more related species (i.e., host species in the order Rodentia) were associated with more Lyme disease cases in humans. Host species richness correlated negatively with the number of Lyme disease cases at the state level (i.e., a dilution effect), a pattern that might be explained by the higher number of reservoir-incompetent species at high levels of species richness at this larger spatial scale. In contrast, a positive correlation was found between species richness and the number of Lyme disease cases at the county level, where a higher proportion of rodent species was associated with higher levels of species richness, potentially amplifying the disease risk. Our results highlight that analyse at a single spatial scale can miss some impacts of biodiversity on human health. Thus, multi-scale analyses with consideration of host phylogenetic diversity are critical for improving our understanding of diversity-disease relationships.