Spider dynamics under vertical vibration and its implications for biological vibration sensing.

Often overlooked, vibration transmission through the entire body of an animal is an important factor in understanding vibration sensing in animals. To investigate the role of dynamic properties and vibration transmission through the body, we used a modal test and lumped parameter modelling for a spider. The modal test used laser vibrometry data on a tarantula, and revealed five modes of the spider in the frequency range of 20-200 Hz. Our developed and calibrated model took into account the bounce, pitch and roll of the spider body and bounce of all the eight legs. We then performed a parametric study using this calibrated model, varying factors such as mass, inertia, leg stiffness, damping, angle and span to study what effect they had on vibration transmission. The results support that some biomechanical parameters can act as physical constraints on vibration sensing. But also, that the spider may actively control some biomechanical parameters to change the signal intensity it can sense. Furthermore, our analysis shows that the parameter changes in front and back legs have a greater influence on whole system dynamics, so may be of particular importance for active control mechanisms to facilitate biological sensing functions.

Effects of Community Richness and Competitive Asymmetry on Protozoa Evolution in Sarracenia purpurea Leaves.

AbstractPredicting evolution in natural systems will require understanding how selection operates in multispecies communities. We predicted that the amount that traits evolve in multispecies mixtures would be less than the amount that would be predicted from the additive contributions of the pairwise interactions and that subordinate species will be more likely to evolve in competitive systems than dominant species. We conducted an experimental test of these predictions using a guild of protozoans found in the water-filled leaves of the pitcher plant Sarracenia purpurea. The response to selection did not significantly change as we increased richness from monocultures to two- and four-species mixtures. In accordance with our second prediction, subordinate species demonstrated greater growth in competition after selection than before, while dominant species generally showed no response to selection. Monod-type experiments to determine minimum resource levels found that the dominant species had much higher resource requirements than the subordinate species and that the minimum resource requirements evolved to be higher in the subordinate species. Importantly, these results suggest that subordinate species evolve to become more similar to dominant species, which may involve resource use convergence. Our findings and other recent works suggest that community diversity can affect evolution in surprising ways that warrant further investigation.

Slit sense organ distribution on the legs of two species of orb-weaving spider (Araneae: Araneidae).

Biotic and abiotic mechanical stimuli are ubiquitous in the environment, and are a widely used source of sensory information in arthropods. Spiders sense mechanical stimuli using hundreds of slit sense organs (small isolated slits, large isolated slits, groups of slits and lyriform organs) distributed across their bodies and appendages. These slit sense organs are embedded in the exoskeleton and detect cuticular strain. Therefore, the spatial pattern of these sensors can give clues into how mechanical stimuli from different sources might be processed and filtered as they are transmitted through the body. Here, we map the distribution of slit sense organs on the legs in two species of orb-weaving spider, A. diadematus and T. edulis, in which slit sense organ distribution has not previously been investigated. We image the spiders' legs using scanning electron microscopy, and trace the position and orientation of slits on these images to describe the distribution and external morphology of the slit sense organs. We show that both species have a similar distribution of slit sense organs, with small isolated slits occurring in consistent lines parallel to the long axis of the legs, whilst large isolated slits, groups of slits and lyriform organs appear in fixed positions near the leg joints. Our findings support what has been described in the literature for several other species of spider, which indicates that slit organ arrangement is conserved across spiders in different evolutionary lineages and with disparate hunting strategies. The dispersed distribution of small isolated slits along the whole length of the leg may be used to detect large-scale strain of the leg segment as a result of muscle activity or internal changes in haemolymph pressure.

Correction: Host and environmental determinants of microbial community structure in the marine phyllosphere.

[This corrects the article DOI: 10.1371/journal.pone.0235441.].

Composition of seagrass phyllosphere microbial communities suggests rapid environmental regulation of community structure.

Recent studies have revealed that seagrass blade surfaces, also known as the phyllosphere, are rich habitats for microbes; however, the primary drivers of composition and structure in these microbial communities are largely unknown. This study utilized a reciprocal transplant approach between two sites with different environmental conditions combined with 16S rRNA gene sequencing (iTag) to examine the relative influence of environmental conditions and host plant on phyllosphere community composition of the seagrass Thalassia testudinum. After 30 days, identity of phyllosphere microbial community members was more similar within the transplant sites than between despite differences in the source of host plant. Additionally, the diversity and evenness of these communities was significantly different between the two sites. These results indicated that local environmental conditions can be a primary driver in structuring seagrass phyllosphere microbial communities over relatively short time scales. Composition of microbial community members in this study also deviated from those in previous seagrass phyllosphere studies with a higher representation of candidate bacterial phyla and archaea than previously observed. The capacity for seagrass phyllosphere microbial communities to shift dramatically with environmental conditions, including ecosystem perturbations, could significantly affect seagrass-microbe interactions in ways that may influence the health of the seagrass host.

The evolution of niche overlap and competitive differences.

Competition can result in evolutionary changes to coexistence between competitors but there are no theoretical models that predict how the components of coexistence change during this eco-evolutionary process. Here we study the evolution of the coexistence components, niche overlap and competitive differences, in a two-species eco-evolutionary model based on consumer-resource interactions and quantitative genetic inheritance. Species evolve along a one-dimensional trait axis that allows for changes in both niche position and species intrinsic growth rates. There are three main results. First, the breadth of the environment has a strong effect on the dynamics, with broader environments leading to reduced niche overlap and enhanced coexistence. Second, coexistence often involves a reduction in niche overlap while competitive differences stay relatively constant or vice versa; in general changes in competitive differences maintain coexistence only when niche overlap remains constant. Large simultaneous changes in niche overlap and competitive difference often result in one of the species being excluded. Third, provided that the species evolve to a state where they coexist, the final niche overlap and competitive difference values are independent of the system's initial state, although they do depend on the model's parameters. The model suggests that evolution is often a destructive force for coexistence due to evolutionary changes in competitive differences, a finding that expands the paradox of diversity maintenance.

Host and environmental determinants of microbial community structure in the marine phyllosphere.

Although seagrasses are economically and ecologically critical species, little is known about their blade surface microbial communities and how these communities relate to the plant host. To determine microbial community composition and diversity on seagrass blade surfaces and in the surrounding seawater,16S rRNA gene sequencing (iTag) was used for samples collected at five sites along a gradient of freshwater input in the northern Gulf of Mexico on three separate sampling dates. Additionally, seagrass surveys were performed and environmental parameters were measured to characterize host characteristics and the abiotic conditions at each site. Results showed that Thalassia testudinum (turtle grass) blades hosted unique microbial communities that were distinct in composition and diversity from the water column. Environmental conditions, including water depth, salinity, and temperature, influenced community structure as blade surface microbial communities varied among sites and sampling dates in correlation with changes in environmental parameters. Microbial community composition also correlated with seagrass host characteristics, including growth rates and blade nutrient composition. There is some evidence for a core community for T. testudinum as 21 microorganisms from five phyla (Cyanobacteria, Proteobacteria, Planctomycetes, Chloroflexi, and Bacteroidetes) were present in all blade surface samples. This study provides new insights and understanding of the processes that influence the structure of marine phyllosphere communities, how these microbial communities relate to their host, and their role as a part of the seagrass holobiont, which is an important contribution given the current decline of seagrass coverage worldwide.

Species-specific differences determine responses to a resource pulse and predation.

The effects of resource pulses on natural communities are known to vary with the type of pulse. However, less is known about mechanisms that determine the responses of different species to the same pulse. We hypothesized that these differences are related to the size of the species, as increasing size may be correlated with increasing competitive ability and decreasing tolerance to predation. A factorial experiment quantified the magnitude and timing of species' responses to a resource pulse using the aquatic communities found in the leaves of the carnivorous pitcher plant, Sarracenia purpurea. We added prey to leaves and followed the abundances of bacteria and bacterivores (protozoa and rotifers) in the presence and absence of a top predator, larvae of the mosquito Wyeomyia smithii. Resource pulses had significant positive effects on species abundances and diversity in this community; however, the magnitude and timing of responses varied among the bacterivore species and was not related to body size. Larger bacterivores were significantly suppressed by predators, while smaller bacterivores were not; predation also significantly reduced bacterivore species diversity. There were no interactions between the effects of the resource pulse and predation on protozoa abundances. Over 67 days, some species returned to pre-pulse abundances quickly, others did not or did so very slowly, resulting in new community states for extended periods of time. This study demonstrates that species-specific differences in responses to resource pulses and predation are complex and may not be related to simple life history trade-offs associated with size.

Predator identity more than predator richness structures aquatic microbial assemblages in Sarracenia purpurea leaves.

The importance of predators in influencing community structure is a well-studied area of ecology. However, few studies test ecological hypotheses of predation in multi-predator microbial communities. The phytotelmic community found within the water-filled leaves of the pitcher plant, Sarracenia purpurea, exhibits a simple trophic structure that includes multiple protozoan predators and microbial prey. Using this system, we sought to determine whether different predators target distinct microorganisms, how interactions among protozoans affect resource (microorganism) use, and how predator diversity affects prey community diversity. In particular, we endeavored to determine if protozoa followed known ecological patterns such as keystone predation or generalist predation. For these experiments, replicate inquiline microbial communities were maintained for seven days with five protozoan species. Microbial community structure was determined by 16S rRNA gene amplicon sequencing (iTag) and analysis. Compared to the control (no protozoa), two ciliates followed patterns of keystone predation by increasing microbial evenness. In pairwise competition treatments with a generalist flagellate, prey communities resembled the microbial communities of the respective keystone predator in monoculture. The relative abundance of the most common bacterial Operational Taxonomic Unit (OTU) in our system decreased compared to the control in the presence of these ciliates. This OTU was 98% similar to a known chitin degrader and nitrate reducer, important functions for the microbial community and the plant host. Collectively, the data demonstrated that predator identity had a greater effect on prey diversity and composition than overall predator diversity.

Plant community responses to simultaneous changes in temperature, nitrogen availability, and invasion.

BACKGROUND: Increasing rates of change in climate have been observed across the planet and have contributed to the ongoing range shifts observed for many species. Although ecologists are now using a variety of approaches to study how much and through what mechanisms increasing temperature and nutrient pollution may influence the invasions inherent in range shifts, accurate predictions are still lacking. METHODS AND RESULTS: In this study, we conducted a factorial experiment, simultaneously manipulating warming, nitrogen addition and introduction of Pityopsis aspera, to determine how range-shifting species affect a plant community. We quantified the resident community using ordination scores, then used structural equation modeling to examine hypotheses related to how plants respond to a network of experimental treatments and environmental variables. Variation in soil pH explained plant community response to nitrogen addition in the absence of invasion. However, in the presence of invasion, the direct effect of nitrogen on the community was negligible and soil moisture was important for explaining nitrogen effects. We did not find effects of warming on the native plant community in the absence of invasion. In the presence of invasion, however, warming had negative effects on functional richness directly and invasion and herbivory explained the overall positive effect of warming on the plant community. CONCLUSIONS AND SIGNIFICANCE: This work highlights the variation in the biotic and abiotic factors responsible for explaining independent and collective climate change effects over a short time scale. Future work should consider the complex and non-additive relationships among factors of climate change and invasion in order to capture more ecologically relevant features of our changing environment.

Effects of disturbance on vegetation by sand accretion and erosion across coastal dune habitats on a barrier island.

Coastal geomorphology and vegetation are expected to be particularly sensitive to climate change, because of disturbances caused by sea-level rise and increased storm frequency. Dunes have critical reciprocal interactions with vegetation; dunes create habitats for plants, while plants help to build dunes and promote geomorphological stability. These interactions are also greatly affected by disturbances associated with sand movement, either in accretion (dune building) or in erosion. The magnitude and intensity of disturbances are expected to vary with habitat, from the more exposed and less stable foredunes, to low-lying and flood-prone interdunes, to the protected and older backdunes. Permanent plots were established at three different spatial scales on St George Island, FL, USA, where the vegetation and dune elevation were quantified annually from 2011 to 2013. Change in elevation, either through accretion or erosion, was used as a measure of year-to-year disturbance over the 2 years of the study. At the scale of different dune habitats, foredunes were found to have the greatest disturbance, while interdunes had the least. Elevation and habitat (i.e. foredune, interdune, backdune) were significantly correlated with plant community composition. Generalized linear models conducted within each habitat show that the change in elevation (disturbance) is also significantly correlated with the plant community, but only within foredunes and interdunes. The importance of disturbance in exposed foredunes was expected and was found to be related to an increasing abundance of a dominant species (Uniola paniculata) in eroding areas. The significant effect of disturbance in the relatively stable interdunes was surprising, and may be due to the importance of flooding associated with small changes in elevation in these low-lying areas. Overall, this study documents changes in the plant community associated with elevation, and demonstrates that the foredune and interdune communities are also associated with the responses of specific species to local changes in elevation due to accretion or erosion.

Effects of a ciliate protozoa predator on microbial communities in pitcher plant (Sarracenia purpurea) leaves.

The aquatic communities found within the water filled leaves of the pitcher plant, Sarracenia purpurea, have a simple trophic structure providing an ideal system to study microscale interactions between protozoan predators and their bacterial prey. In this study, replicate communities were maintained with and without the presence of the bactivorous protozoan, Colpoda steinii, to determine the effects of grazing on microbial communities. Changes in microbial (Archaea and Bacteria) community structure were assessed using iTag sequencing of 16S rRNA genes. The microbial communities were similar with and without the protozoan predator, with>1000 species. Of these species, Archaea were negligible, with Bacteria comprising 99.99% of the microbial community. The Proteobacteria and Bacteroidetes were the most dominant phyla. The addition of a protozoan predator did not have a significant effect on microbial evenness nor richness. However, the presence of the protozoan did cause a significant shift in the relative abundances of a number of bacterial species. This suggested that bactivorous protozoan may target specific bacterial species and/or that certain bacterial species have innate mechanisms by which they evade predators. These findings help to elucidate the effect that trophic structure perturbations have on predator prey interactions in microbial systems.

Rethinking niche evolution: experiments with natural communities of Protozoa in pitcher plants.

Classic niche theory predicts that competing species will evolve to use different resources and interact less, whereas recent niche-converge ideas predict that species evolve to use similar resources and interact more. Most data supporting niche evolution are based on observations of contemporary niche use, whereas experimental support is quite sparse. We followed the evolution of four species of Protozoa during succession in the water-filled leaves of the pitcher plant, Sarracenia purpurea, and found that evolution in multispecies systems follows a surprising pattern. Over several hundred generations, weak competitors evolved to be stronger, while strong competitors evolved to become weaker, which does not conform to expectations of either niche divergence or convergence. Evolution in this system appears to occur in response to characteristics of a suite of several competitors in the community, rather than pairwise interactions. Ecologists may need to rethink the roles of competition and evolution in structuring communities.

Assembly-history dynamics of a pitcher-plant protozoan community in experimental microcosms.

BACKGROUND: History drives community assembly through differences both in density (density effects) and in the sequence in which species arrive (sequence effects). Density effects arise from predictable population dynamics, which are free of history, but sequence effects are due to a density-free mechanism, arising solely from the order and timing of immigration events. Few studies have determined how components of immigration history (timing, number of individuals, frequency) alter local dynamics to determine community assembly, beyond addressing when immigration history produces historically contingent assembly. METHODS/FINDINGS: We varied density and sequence effects independently in a two-way factorial design to follow community assembly in a three-species aquatic protozoan community. A superior competitor, Colpoda steinii, mediated alternative community states; early arrival or high introduction density allowed this species to outcompete or suppress the other competitors (Poterioochromonas malhamensis and Eimeriidae gen. sp.). Multivariate analysis showed that density effects caused greater variation in community states, whereas sequence effects altered the mean community composition. CONCLUSIONS: A significant interaction between density and sequence effects suggests that we should refine our understanding of priority effects. These results highlight a practical need to understand not only the "ingredients" (species) in ecological communities but their "recipes" as well.

Testing successional hypotheses of stability, heterogeneity, and diversity in pitcher-plant inquiline communities.

Succession is a foundation concept in ecology that describes changes in species composition through time, yet many successional patterns have not been thoroughly investigated. We highlight three hypotheses about succession that are often not clearly stated or tested: (1) individual communities become more stable over time, (2) replicate communities become more similar over time, and (3) diversity peaks at mid-succession. Testing general patterns of succession requires estimates of variation in trajectories within and among replicate communities. We followed replicate aquatic communities found within leaves of purple pitcher plants (Sarracenia purpurea) to test these three hypotheses. We found that stability of individual communities initially decreased, but then increased in older communities. Predation was highest in younger leaves but then declined, while competition was likely strongest in older leaves, as resources declined through time. Higher levels of predation and competition corresponded with periods of higher stability. As predicted, heterogeneity among communities decreased with age, suggesting that communities became more similar over time. Changes in diversity depended on trophic level. The diversity of bacteria slightly declined over time, but the diversity of consumers of bacteria increased linearly and strongly throughout succession. We suggest that studies need to focus on the variety of environmental drivers of succession, which are likely to vary through time and across habitats.

Evolution of prey in ecological time reduces the effect size of predators in experimental microcosms.

Ecologists have long studied the effect of predators on prey population abundance while evolutionary biologists have measured prey trait evolution in response to predation. Ecological and evolutionary processes were generally thought to occur on different time scales, but recent evidence suggests that evolution may alter the ecological effects of predation over the course of ecological experiments. We used a protozoan and its mosquito-larvae predator, naturally found in the water-filled leaves of pitcher plants, to examine the effect of prey evolution on predator-prey interactions. In experiments conducted over 12 days (approximately 50 prey generations, but less than one predator generation), we measured a decrease in the effect of mosquito larvae predators on protozoa prey populations. In a separate set of experiments, we found that the presence of predators corresponded with evolution of smaller cell size and increased population growth rate. In ecological experiments, two situations commonly occur: strong selection pressure applied by the treatment itself and discrepancies in generation times of associate species. Our results suggest that in either situation, the resulting evolutionary patterns may lead to dramatic and important changes in ecological effect size.

Habitat destruction and metacommunity size in pen shell communities.

1. In spatially structured communities, habitat destruction can have two effects: first, a main effect that occurs because of the loss of habitat area within a larger region, and a secondary effect due to changes in the spatial arrangement of local communities. Changes to the spatial arrangement can, in turn, affect the migration and extinction rates within local communities. 2. Our study involved the experimental destruction of entire local communities within larger regions in natural marine microcosms. Large and small arrays of dead pen shells were created in a shallow bay in north Florida, and the colonization by both encrusting and motile species on this empty substrate were followed through time. After most species had become established, half of the large arrays were perturbed to create small arrays by removal of half the shells, simulating habitat destruction. 3. After 48 days of further community development, comparisons of the large arrays, reduced arrays and original small arrays suggested that the mechanisms by which habitat destruction affects diversity could depend upon the size of the region affected and the natural history of the species being studied. 4. Habitat destruction reduced the diversity of motile species to a level lower than that found in the undisturbed small arrays, suggesting that the species that assembled in the original large metacommunities negatively influenced the species that occurred ultimately in the converted small arrays. 5. With sessile species, habitat destruction created richness levels that were intermediate to those of small and large arrays. The initial predestruction richness appears to have had a positive effect; because sessile species cannot disperse as adults, they may not respond to significant shifts in metacommunity size later in succession. Initial metacommunity size may be important for allowing individuals to select appropriate habitats before they settle.

A critical review of twenty years' use of the resource-ratio theory.

A model of species interactions based on their use of shared resources was proposed in 1972 by Robert MacArthur and later expanded in an article (1980) and a book (1982) by David Tilman. This "resource-ratio theory" has been used to make a number of testable predictions about competition and community patterns. We reviewed 1,333 papers that cite Tilman's two publications to determine whether predictions of the resource-ratio theory have been adequately tested and to summarize their general conclusions. Most of the citations do not directly test the theory: only 26 studies provide well-designed tests of one or more predictions, resulting in 42 individual tests of predictions. Most of these tests were conducted in the laboratory or experimental microcosms and used primary producers in freshwater systems. Overall, the predictions of the resource-ratio theory were supported 75% of the time. One of the primary predictions of the model, that species dominance varies with the ratio of resource availabilities, was supported by 13 of 16 tests, but most other predictions have been insufficiently tested. We suggest that more experimental work in a variety of natural systems is seriously needed, especially studies designed to test predictions related to resource supply and consumption rates.