Dispersal and spatial heterogeneity allow coexistence between enemies and protective mutualists

Protective mutualisms, where a symbiont reduces the negative effects of another species on a shared host, represent a common type of species interaction in natural communities, yet it is still unclear what ecological conditions might favor their emergence. Studies suggest that the initial evolution of protective mutualists might involve closely related pathogenic variants with similar life histories, but different competitive abilities and impacts on host fitness. We derive a model to evaluate this hypothesis and show that, in general, a protective variant cannot spread from rarity or exclude a more pathogenic strain. While the conditions allowing mutualist invasion are more likely with increased environmental productivity, they still depend on initial densities in the invaded patch exceeding a threshold, highlighting the likely importance of spatial structure and demographic stochasticity. Using a numerical simulation approach, we show that regional coexistence is in fact possible in an explicitly spatial system and that, under some circumstances, the mutualist population can exclude the enemy. More broadly, the establishment of protective mutualists may be favored when there are other life‐history differences from more pathogenic symbionts, such as vertical transmission or additional direct benefits to hosts.     

Competitive coexistence in spatially structured environments: a synthesis

Theoretical developments in spatial competitive coexistence are far in advance of empirical investigations. A framework that makes comparative predictions for alternative hypotheses is a crucial element in narrowing this gap. This review attempts to synthesize spatial competition theory into such a framework, with the goal of motivating empirical investigations that adopt the comparative approach. The synthesis presented is based on a major axis, coexistence in spatially homogeneous vs. heterogeneous competitive environments, along which the theory can be organized. The resulting framework integrates such key concepts as niche theory, spatial heterogeneity and spatial scale(s) of coexistence. It yields comparative predictions that can guide empirical investigations.     

Trade-offs in community ecology: linking spatial scales and species coexistence

Trade‐offs in species performances of different ecological functions is one of the most common explanations for coexistence in communities. Despite the potential for species coexistence occurring at local or regional spatial scales, trade‐offs are typically approached at a single scale. In recent years, ecologists have increasingly provided evidence for the importance of community processes at both local and regional spatial scales. This review summarizes the theoretical predictions for the traits associated with trade‐offs under different conditions and at different spatial scales. We provide a spatial framework for understanding trade‐offs, coexistence and the supportive empirical evidence. Predictions are presented that link the patterns of diversity observed to the patterns of trade‐offs that lead to coexistence at different spatial scales. Recent evidence for the evolution of trade‐offs under different conditions is provided which explores both laboratory microcosm studies and phylogenetic tests. Examining trade‐offs within a spatial framework can provide a strong approach to understanding community structure and dynamics, while explaining patterns of species diversity.     

Dispersal and natural enemies interact to drive spatial synchrony and decrease stability in patchy populations

Spatial synchrony is widespread in natural populations but the mechanisms that underpin it are not yet fully understood. Two key biotic drivers of spatial synchrony have been identified: dispersal and trophic interactions (e.g. natural enemies). We used spatially structured, patchy bacterial populations to show that although increased dispersal always enhanced spatial synchrony of fluctuations in bacterial abundance, this effect was far stronger in the presence of a bacteriophage parasite. Bacteriophages drove strong within patch fluctuations in bacterial abundance that became phase locked through dispersal. Furthermore, the way in which stability, measured as constancy, responded to increasing dispersal was qualitatively different depending on whether parasites were present or not. Patch-level constancy decreased with dispersal in the presence of parasites, whereas dispersal increased patch-level constancy in the absence of parasites. Population-level constancy also decreased with dispersal in the presence of parasites, but was unaffected by dispersal in the absence of parasites. These contrasting patterns were likely due to the different role played by dispersal in the presence and absence of parasites, synchronizing dynamics in the former case and averaging stochastic fluctuations in the latter. Taken together, our findings suggest that dispersal and natural enemies can interact to drive spatially synchronous population fluctuations that decrease stability at both the patch and population level.     

Spatial context strongly affects community composition of both passively and actively dispersing pool invertebrates in a highly heterogeneous landscape

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Spatial coherence and the persistence of high diversity in spatially heterogeneous landscapes

Our planet hosts a variety of highly diverse ecosystems. The persistence of high diversity is generally attributed to factors such as the structure of interactions among species and the dispersal of species in metacommunities. Here, we show that large contiguous landscapes—that are characterized by high dispersal—facilitate high species richness due to the spatial heterogeneity in interspecies interactions. We base our analysis on metacommunities under high dispersal where species densities become equal across habitats (spatially coherent). We find that the spatially coherent metacommunity can be represented by an effective species interaction‐web that has a significantly lower complexity than the constituent habitats. Our framework also explains how spatial heterogeneity eliminates differences in the effective interaction‐web, providing a basis for deviations from the area‐heterogeneity tradeoff. These results highlight the often‐overlooked case of high dispersal where spatial coherence provides a novel mechanism for supporting high diversity in large heterogeneous landscapes.     

Habitat fragmentation and species diversity in competitive communities

Habitat loss is one of the key drivers of the ongoing decline of biodiversity. However, ecologists still argue about how fragmentation of habitat (independent of habitat loss) affects species richness. The recently proposed habitat amount hypothesis posits that species richness only depends on the total amount of habitat in a local landscape. In contrast, empirical studies report contrasting patterns: some find positive and others negative effects of fragmentation per se on species richness. To explain this apparent disparity, we devise a stochastic, spatially explicit model of competitive species communities in heterogeneous habitats. The model shows that habitat loss and fragmentation have complex effects on species diversity in competitive communities. When the total amount of habitat is large, fragmentation per se tends to increase species diversity, but if the total amount of habitat is small, the situation is reversed: fragmentation per se decreases species diversity.     

Changes in forest structure, species diversity and spatial pattern following hurricane disturbance in a Piedmont North Carolina forest, USA

Aims: Large hurricanes have profound impacts on temperate forests,but owing to their infrequent nature these effects have rarelybeen examined in detail. In 1996, Hurricane Fran significantlydamaged many long-term tree census plots in the Duke Foreston the North Carolina Piedmont, thereby providing an exceptionalopportunity to examine pre- and post-hurricane forest compositionaltrajectories. Our goal was to examine immediate, short-term(0–4 years) and longer term (5 year) hurricane-inducedstructural, spatial and compositional changes in the tree population(stem d.b.h > 1 cm) in the context of our detailed, long-termknowledge of the dynamics of these forests. Methods: We surveyed stem damage and tree mortality in 34 long-term permanentplots (ca. 70-year record; 404–1 012 m²) and 7 large mappedtree stands (ca. 20-year record; 5 250–65 000 m²) representingboth transition-phase, even-aged pine stands and uneven-agedupland hardwood forests. We employed three types of damage measuresto quantify stand-level damage severity: percentage of stemsdamaged, percentage of basal area lost and a ‘stand-leveldamage index’. Second-order spatial analysis (Ripley'sK-function) was used to investigate patterns in tree mortality. Important findings: Our study found hurricane effects on the structural attributesof Piedmont forests to be variable and patchy. Changes in treespecies composition, however, were modest. Uprooting was themajor damage type for the overstory trees [diameter at breastheight (d.b.h.) >10 cm] apparently due to the exposure ofthe crowns to high wind combined with heavy rainfall prior toand during the storm. Saplings, juvenile trees and small trees(1–10 cm d.b.h.) of the understory and midstory were mainlydamaged by being pinned or bent by their damaged large neighbors.Hurricane-induced tree mortality varied weakly among species,was positively correlated with pre-hurricane tree size and remainedup to 2-fold higher than pre-hurricane background mortality5 years after the hurricane. Spatial point pattern analysisrevealed a patchy distribution of tree mortality during thehurricane sampling interval. Hurricane Fran resulted in a dramaticincrease in average gap size from ca. 400 m² pre-hurricane toca 1100 m² after the hurricane, whereas maximum gap sizes reached18–34 times larger than the pre-hurricane levels.     

Aggregated spatial distributions of species in a subtropical karst forest,southwestern China

Aims Spatial distribution patterns of species reflect not only the ecological processes but also the habitat features that are related to species distribution. In karst topography, species distribution patterns provide more specific information about their environments. The objectives of this study are as follows: (i) to analyse and explain the spatial distribution patterns of conspecific trees in an old-growth subtropical karst forest; (ii) to investigate pattern changes at different spatial scales; (iii) to test the spatial pattern similarity (or dissimilarity) between trees at different abundances, diameter at breast height classes, canopy layers and different functional groups (shade tolerance and seed dispersal mode); (iv) to examine whether habitat heterogeneity has an important effect on the species spatial distribution.Methods The spatial distributions of woody species with ≥20 individuals in a 1-ha subtropical karst forest plot at Maolan in southwestern China were quantified using the relative neighbourhood density Ω based on the average density of conspecific species in a circular neighbourhood around each species.Important findings Aggregated distribution is the dominant pattern in the karst forest, but the ratio of aggregated species in total species number decreases with an increase in spatial scale. Less abundant species are more aggregated than most abundant species. Aggregation is weaker in larger diameter classes, which is consistent with the prediction of self-thinning. Seed dispersal mode influences spatial patterns, with species dispersed by animals being less aggregated than those dispersed by wind and gravity. Other species functional traits (e.g. shade tolerance) also influence the species spatial distributions. Moreover, differences among species habitat associations, e.g. with rocky outcrops, play a significant role in species spatial distributions. These results indicate that habitat heterogeneity, seed dispersal limitation and self-thinning primarily contribute to the species spatial distributions in this subtropical karst forest.     

Disturbance, interspecific interaction and diversity in metapopulations

Metapopulation diversity patterns depend on the relations among the timescales of local biological interactions (predation, competition), the rates of dispersal among local populations and the patterns of disturbance. We investigate these relationships using a family of simple non-linear Markov chain models. We consider three models for interspecific competition; if the species are identified with early and late successional species, the models describe the facilitation, inhibition and tolerance models of ecological succession. By adding a third competing species we also compare transitive competitive hierarchies and intransitive competitive networks. Finally, we examine the effects of predation in mediating coexistence among competing prey species. In each model we find circumstances in which biotic or abiotic disturbance can increase both local and regional diversity, but those circumstances depend on the various timescales in the model in ways that arc neither obvious nor trivial.     

The influence of landscape diversity and heterogeneity on spatial dynamics of agrobiont linyphiid spiders: An individual-based model

Spiders are important generalist predators in natural pest control. However, agricultural fields are highly disturbed and ephemeral habitats, which present a number of challenges to the organisms living there; likewise landscape diversity and heterogeneity are also thought to be important factors in determining spider spatial dynamics. To investigate the interactions between these factors, we present an individual-based simulation model, which integrates life history characteristics of a typical agrobiont linyphiid spider with a dynamic spatially explicit landscape representation. The landscape contains several habitat types of varying quality and varies in time and space. Simulations showed that spatial landscape diversity (number of habitat types available for the spiders) is crucial for the persistence of spiders, but that spatial heterogeneity (spatial arrangement of patches) only had little impact on spider abundance. The necessary landscape diversity could either be provided by a diverse crop rotation or by including refuges in the form of less frequently managed habitats in the landscape. The presence of refuges greatly boosted numbers of spiders in the landscape as a whole. The most important characteristics of refuge were sanctuary from pesticides and extra prey availability, whereas tillage frequency mattered less. The simulations indicated that agrobiont linyphiids combination of high dispersal abilities and high reproductive rate enables it to exploit the transient resources of the different habitats in the agricultural landscape.     

Invasive spread in meta-food-webs depends on landscape structure,fertilization and species characteristics

Land use change and biological invasions collectively threaten biodiversity. Yet, few studies have addressed how altering the landscape structure and nutrient supply can promote biological invasions and particularly invasive spread (the spread of an invader from the place of introduction), or asked whether and how these factors interact with biotic interactions and invader properties. We here bridge this knowledge gap by providing a holistic network-based approach. Our approach combines a trophic network model with a spatial network model allowing us to test which combinations of abiotic and biotic factors can facilitate invasions and in particular invasive spread in food webs. We numerically simulated 6300 single-species invasions in clustered and random landscapes at different levels of nutrient supply. In total, our simulation experiment yielded 69% successful invasions – 71% in clustered landscapes and 66% in random landscapes, with the proportion of successful invasions increasing with nutrient supply. However, invasive spread was generally higher in random than in clustered landscapes. The latter can facilitate invasive spread within a habitat cluster, but prevent invasive spread between clusters. Low nutrient levels generally prevented the establishment of invasive species and their subsequent spread. However, successful invaders could have more severe impacts as they contribute more to total biomass density and species richness under such conditions. Good dispersal abilities drive the broad-scale spread of invasive species in fragmented landscapes. Our approach makes an important contribution towards a better understanding of what combination of landscape and invader properties can facilitate or prevent invasive spread in natural ecosystems. This should allow ecologists to more effectively predict and manage biological invasions.     

How do similarities in spatial distributions and interspecific associations affect the coexistence of Quercus species in the Baotianman National Nature Reserve,Henan, China

Congeneric species often have similar ecological characteristics and use similar resources. These similarities may make it easier for them to co‐occur in a similar habitat but may also lead to strong competitions that limit their coexistence. Hence, how do similarities in congeneric species affect their coexistence exactly? This study mainly used spatial point pattern analysis in two 1 hm² plots in the Baotianman National Nature Reserve, Henan, China, to compare the similarities in spatial distributions and interspecific associations of Quercus species. Results revealed that Quercus species were all aggregated under the complete spatial randomness null model, and aggregations were weaker under the heterogeneous Poisson process null model in each plot. The interspecific associations of Quercus species to non‐Quercus species were very similar in Plot 1. However, they can be either positive or negative in different plots between the co‐occurring Quercus species. The spatial distributions of congeneric species, interspecific associations with non‐Quercus species, neighborhood richness around species, and species diversity were all different between the two plots. We found that congeneric species did have some similarities, and the closely related congeneric species can positive or negative associate with each other in different plots. The co‐occurring congeneric species may have different survival strategies in different habitats. On the one hand, competition among congenerics may lead to differentiation in resource utilization. On the other hand, their similar interspecific associations can strengthen their competitive ability and promote local exclusion to noncongeneric species to obtain more living space. Our results provide new knowledge for us to better understand the coexistence mechanisms of species.     

Aspects of the ecology of the prickly sculpin,Cottus asper Richardson,a persistent native species in Clear Lake,Lake County,California

Synopsis The biology of the prickly sculpin was investigated in Clear Lake, Lake County, California in order to determine how it has persisted in the face of introductions of numerous exotic species when most other native species have declined in abundance or have become extinct. Sculpins over 15 mm SL inhabited all types of benthic habitats in the lake, while post larval sculpins were pelagic when the postlarvae of exotic species were absent. The feeding ecology of sculpins was distinct from the other fishes in the lake in that they fed largely on amphipods and chironomid midge larvae regardless of the time of year, time of day, or habitat. Sculpins were uncommon in the stomachs of piscivorous fishes, except juvenile largemouth bass (Micropterus salmoides). It is concluded that prickly sculpins have persisted in Clear Lake in part because they are ecologically distinct from the exotic species and are not preyed upon by them to any great extent, and in part because they have managed to survive other man-related perturbations of this ecosystem.     

Tree heterogeneity,resource availability,and larger scale processes regulating arboreal ant species richness

Abstract Processes acting on different spatial and temporal scales may influence local species richness. Ant communities are usually described as interactive and therefore determined by local processes. In this paper we tested two hypotheses linked to the question of why there is local variation in arboreal ant species richness in the Brazilian savanna (‘cerrado’). The hypotheses are: (i) there is a positive relationship between ant species richness and tree species richness, used as a surrogate of heterogeneity; and (ii) there is a positive relationship between ant species richness and tree density, used as a surrogate of resource availability. Arboreal ants were sampled in two cerrado sites in Brazil using baited pitfall traps and manual sampling, in quadrats of 20 m × 50 m. Ant species richness in each quadrat was used as the response variable in regression tests, using tree species richness and tree density as explanatory variables. Ant species richness responded positively to tree species richness and density. Sampling site also influenced ant species richness, and the relationship between tree density and tree species richness was also positive and significant. Tree species richness may have influenced ant species richness through three processes: (i) increasing the variety of resources and allowing the existence of a higher number of specialist species; (ii) increasing the amount of resources to generalist species; and (iii) some other unmeasured factor may have influenced both ant and tree species richness. Tree density may also have influenced ant species richness through three processes: (i) increasing the amount of resources and allowing a higher ant species richness; (ii) changing habitat conditions and dominance hierarchies in ant communities; and (iii) increasing the area and causing a species–area pattern. Processes acting on larger scales, such as disturbance, altitude and evolutionary histories, as well as sampling effect may have caused the difference between sites.