Biotic Forces Research Articles

Drivers and consequences of variation in individual social connectivity

There is a growing interest in identifying specific causes and consequences of variation in individual social behaviour as a means of understanding how different individuals balance the costs and benefits of group living. In this study, we used social networks to examine variation in individual social behaviour in wild Grant's gazelles, Nanger granti, and explored potential drivers and consequences of this variation. First, we quantified two aspects of individual network position (weighted degree and closeness) on a monthly basis for 12 consecutive months and examined life-history (age) and abiotic (rainfall) factors that could explain among-individual variation in network position. Next, we examined the level of within-individual repeatability in network position over time. We then tested for potential consequences of this variation focusing on parasite infection and diet quality. Rainfall and age were strong predictors of variation in closeness but not degree. Interestingly, we found that one aspect of individual network position (closeness) varied over time, while another (degree) was moderately repeatable. The difference in within-individual repeatability of the two measures may be explained by the dependence of closeness on rainfall. In addition, we found that individual network position had consequences for both parasitism and diet, but the magnitude and direction of these effects depended on parasite type, connectivity measure and environmental conditions. Overall, our results suggest that environmental and host factors strongly influence variability in certain aspects of social connectivity in Grant's gazelles, and that abiotic and biotic forces, together, mediate the consequences of social network position.

Sulfolobus islandicus meta-populations in Yellowstone National Park hot springs.

Abiotic and biotic forces shape the structure and evolution of microbial populations. We investigated forces that shape the spatial and temporal population structure of Sulfolobus islandicus by comparing geochemical and molecular analysis from seven hot springs in five regions sampled over 3 years in Yellowstone National Park. Through deep amplicon sequencing, we uncovered 148 unique alleles at two loci whose relative frequency provides clear evidence for independent populations in different hot springs. Although geography controls regional geochemical composition and population differentiation, temporal changes in population were not explained by corresponding variation in geochemistry. The data suggest that the influence of extinction, bottleneck events and/or selective sweeps within a spring and low migration between springs shape these populations. We suggest that hydrologic events such as storm events and surface snowmelt runoff destabilize smaller hot spring environments with smaller populations and result in high variation in the S. islandicus population over time. Therefore, physical abiotic features such as hot spring size and position in the landscape are important factors shaping the stability and diversity of the S. islandicus meta-population within Yellowstone National Park.

Genetic diversity of Atlantic Bluefin tuna in the Mediterranean Sea: insights from genome-wide SNPs and microsatellites

BackgroundElucidating the patterns of the Atlantic Bluefin tuna [ABFT, Thunnus thynnus (Linnaeus, 1758)] population structure constitutes a challenging task of great importance. Most of the unique challenges stem from its biology, as well as the attributes of the marine realm in which it disperses. Accurate information is urgently needed for stock assessment, and the identification of critical features to the persistence and adaptation of populations in order to formulate and adopt effective strategies for ABFT conservation and management. Conclusions of a great number of ABFT genetic studies on the Mediterranean Sea stock structure are rather controversial and not yet conclusive. In this study, ABFT genomic diversity was investigated in the Mediterranean Sea, which is the most important area for the species’ reproduction.ResultsAnalyzing genome-wide SNPs and microsatellites from ABFT samples collected throughout the Mediterranean Sea did not provide strong evidence of genetic structure, pointing towards the existence of a single panmictic unit. An alternative view would recognize a failure to reject the null hypothesis of a panmictic unit as an effect of the study’s sampling design, the type of markers used, and the effectiveness/suitability of analysis methods in respect to the species biological characteristics or any combination of the above.ConclusionsUnravelling the drivers of ABFT population diversity would require the consideration of important aspects of the species spawning behavior for the determination of the appropriate sampling design. Novel approaches and methods of analysis that will bring together experts in genetics/-omics, ecology and oceanography are deemed necessary. Analyzing ABFT genetic data under the discipline of seascape genetics could provide the analysis framework under which major abiotic and biotic forces controlling ABFT recruitment could be identified, elucidating the complicated population dynamics of the species, while multiple and continuous fisheries monitoring should in all cases be considered as a prerequisite in order to achieve efficient and long-term ABFT conservation.

Climate change and the ash dieback crisis.

Beyond the direct influence of climate change on species distribution and phenology, indirect effects may also arise from perturbations in species interactions. Infectious diseases are strong biotic forces that can precipitate population declines and lead to biodiversity loss. It has been shown in forest ecosystems worldwide that at least 10% of trees are vulnerable to extinction and pathogens are increasingly implicated. In Europe, the emerging ash dieback disease caused by the fungus Hymenoscyphus fraxineus, commonly called Chalara fraxinea, is causing a severe mortality of common ash trees (Fraxinus excelsior); this is raising concerns for the persistence of this widespread tree, which is both a key component of forest ecosystems and economically important for timber production. Here, we show how the pathogen and climate change may interact to affect the future spatial distribution of the common ash. Using two presence-only models, seven General Circulation Models and four emission scenarios, we show that climate change, by affecting the host and the pathogen separately, may uncouple their spatial distribution to create a mismatch in species interaction and so a lowering of disease transmission. Consequently, as climate change expands the ranges of both species polewards it may alleviate the ash dieback crisis in southern and occidental regions at the same time.

Sexual versus Asexual Reproduction: Distinct Outcomes in Relative Abundance of Parthenogenetic Mealybugs following Recent Colonization.

Asexual reproduction, including parthenogenesis in which embryos develop within a female without fertilization, is assumed to confer advantages over sexual reproduction, which includes a “cost of males.” Sexual reproduction largely predominates in animals, however, indicating that this cost is outweighed by the genetic and/or ecological benefits of sexuality, including the acquisition of advantageous mutations occurring in different individuals and the elimination of deleterious mutations. But the evolution of sexual reproduction remains unclear, because we have limited examples that demonstrate the relative success of sexual lineages in the face of competition from asexual lineages in the same environment. Here we investigated a sympatric occurrence of sexual and asexual reproduction in the pineapple mealybug, Dysmicoccus brevipes. This pest invaded southwestern Japan, including Okinawa and Ishigaki Islands, in the 1930s in association with imported pineapple plants. Our recent censuses demonstrated that on Okinawa sexually reproducing individuals can coexist with and even dominate asexual individuals in the presence of habitat and resource competition, which is considered to be severe for this nearly immobile insect. Molecular phylogeny based on partial DNA sequences in the mitochondrial and nuclear genomes, as well as the endosymbiotic bacterial genome, revealed that the asexual lineage diverged from a common sexual ancestor in the relatively recent past. In contrast, only the asexual lineage exhibiting obligate apomictic thelytoky was discovered on Ishigaki. Co-existence of the two lineages cannot be explained by the results of laboratory experiments, which showed that the intrinsic rate of increase in the sexual lineage was not obviously superior to that of the asexual lineage. Differences in biotic and/or abiotic selective forces operating on the two islands might be the cause of this discrepancy. This biological system offers a unique opportunity to assess the relative success of sexual versus asexual lineages with an unusual morphology and life cycle.

A comprehensive analysis of mechanical and morphological traits in temperate and tropical seagrass species

Knowledge of plant mechanical traits is important in understanding how plants resistabiotic and biotic forces and in explaining ecological strategies such as leaf lifespan. To date,these traits have not been systematically evaluated in seagrasses. We analysed mechanical(breaking force and tensile strength) and associated traits (thickness, width, length, fibre content,mass area, and lifespan) of leaves in 22 seagrass species (around one-third of all known seagrassspecies) to examine (1) the inter-specific variation of these traits in relation to growth form andbioregions, (2) the contribution of morphology to leaf breaking force, (3) how breaking force scalesto leaf dimensions, and (4) how mechanical and structural traits correlate to leaf longevity. We alsocompared our seagrass dataset with terrestrial plant databases to examine similarities betweenthem. Large variation in leaf breaking force was found among seagrass species but, on average,temperate species resisted higher forces than tropical species. Variation in leaf breaking force waslargely explained by differences in leaf width rather than thickness, likely due to the benefits inleaf reconfiguration and light interception. Species of large dimensions (long leaves) typically hadhigh leaf breaking force, plausibly to tolerate the drag forces they may experience, which are proportionalto the leaf area. Leaves of long-lived species typically had high mass per leaf area andfibre content and they supported high breaking forces. Compared to terrestrial plants, seagrassesare short-lived species with moderately strong fibre-reinforced leaves, which probably evolved towithstand the hydrodynamic forces occurring in the sea, and in response to other environmentalfactors. Overall, our analysis provides new insights into the physical performance of seagrasses inthe marine environment.

Asymmetric ecological conditions favor Red-Queen type of continued evolution over stasis

Four decades ago, Leigh Van Valen presented the Red Queen's hypothesis to account for evolution of species within a multispecies ecological community [Van Valen L (1973) Evol Theory 1(1):1-30]. The overall conclusion of Van Valen's analysis was that evolution would continue even in the absence of abiotic perturbations. Stenseth and Maynard Smith presented in 1984 [Stenseth NC, Maynard Smith J (1984) Evolution 38(4):870-880] a model for the Red Queen's hypothesis showing that both Red-Queen type of continuous evolution and stasis could result from a model with biotically driven evolution. However, although that contribution demonstrated that both evolutionary outcomes were possible, it did not identify which ecological conditions would lead to each of these evolutionary outcomes. Here, we provide, using a simple, yet general population-biologically founded eco-evolutionary model, such analytically derived conditions: Stasis will predominantly emerge whenever the ecological system contains only symmetric ecological interactions, whereas both Red-Queen and stasis type of evolution may result if the ecological interactions are asymmetrical, and more likely so with increasing degree of asymmetry in the ecological system (i.e., the more trophic interactions, host-pathogen interactions, and the like there are [i.e., +/- type of ecological interactions as well as asymmetric competitive (-/-) and mutualistic (+/+) ecological interactions]). In the special case of no between-generational genetic variance, our results also predict dynamics within these types of purely ecological systems.

The role of biotic forces in driving macroevolution: beyond the Red Queen.

A multitude of hypotheses claim that abiotic factors are the main drivers of macroevolutionary change. By contrast, Van Valen's Red Queen hypothesis is often put forward as the sole representative of the view that biotic forcing is the main evolutionary driver. This imbalance of hypotheses does not reflect our current knowledge: theoretical work demonstrates the plausibility of biotically driven long-term evolution, whereas empirical work suggests a central role for biotic forcing in macroevolution. We call for a more pluralistic view of how biotic forces may drive long-term evolution that is compatible with both phenotypic stasis in the fossil record and with non-constant extinction rates. Promising avenues of research include contrasting predictions from relevant theories within ecology and macroevolution, as well as embracing both abiotic and biotic proxies while modelling long-term evolutionary data. By fitting models describing hypotheses of biotically driven macroevolution to data, we could dissect their predictions and transcend beyond pattern description, possibly narrowing the divide between our current understanding of micro- and macroevolution.

Evolving insights to understanding mycorrhizas

Evolving insights to understanding mycorrhizas

Synchronous dynamics of zooplankton competitors prevail in temperate lake ecosystems.

Although competing species are expected to exhibit compensatory dynamics (negative temporal covariation), empirical work has demonstrated that competitive communities often exhibit synchronous dynamics (positive temporal covariation). This has led to the suggestion that environmental forcing dominates species dynamics; however, synchronous and compensatory dynamics may appear at different length scales and/or at different times, making it challenging to identify their relative importance. We compiled 58 long-term datasets of zooplankton abundance in north-temperate and sub-tropical lakes and used wavelet analysis to quantify general patterns in the times and scales at which synchronous/compensatory dynamics dominated zooplankton communities in different regions and across the entire dataset. Synchronous dynamics were far more prevalent at all scales and times and were ubiquitous at the annual scale. Although we found compensatory dynamics in approximately 14% of all combinations of time period/scale/lake, there were no consistent scales or time periods during which compensatory dynamics were apparent across different regions. Our results suggest that the processes driving compensatory dynamics may be local in their extent, while those generating synchronous dynamics operate at much larger scales. This highlights an important gap in our understanding of the interaction between environmental and biotic forces that structure communities.

Long-term performance and herbivory of tree seedlings planted into primary and secondary forests of Central Amazonia

Abstract:Plant survival and growth in tropical rain forest are affected by different biotic and abiotic forces. As time elapses and plants grow the relative importance of such forces as regeneration inhibitors and/or facilitators may change according to habitat and species. To detect within- and among-species divergences in performance over time in different habitats we followed, for nearly a decade, the survival, growth and herbivory of seedlings of the native tree species: Chrysophyllum pomiferum, Micropholis venulosa and Pouteria caimito. In Central Amazonia, young seedlings were planted into old-growth and secondary forests dominated by Vismia spp. One year after planting, C. pomiferum ranked first (i.e. fast growth, fewer dead and less herbivory) for both habitats, followed by M. venulosa and P. caimito. Initial trends changed over time. In the long term, M. venulosa ranked first for both habitats, followed by C. pomiferum and P. caimito ranked consistently lowest. Within-species divergences in growth and herbivory were greater in secondary forest. Initial seedling responses cannot always be used to predict species persistence in the long term. Contrary to previous estimations, old-growth-forest species can persist under Vismia spp. stands, at least when planted.

Forest annual carbon cost: comment

Forest annual carbon cost: comment

Comparing the adaptive landscape across trait types: larger QTL effect size in traits under biotic selection

BackgroundIn a spatially and temporally variable adaptive landscape, mutations operating in opposite directions and mutations of large effect should be commonly fixed due to the shifting locations of phenotypic optima. Similarly, an adaptive landscape with multiple phenotypic optima and deep valleys of low fitness between peaks will favor mutations of large effect. Traits under biotic selection should experience a more spatially and temporally variable adaptive landscape with more phenotypic optima than that experienced by traits under abiotic selection. To test this hypothesis, we assemble information from QTL mapping studies conducted in plants, comparing effect directions and effect sizes of detected QTL controlling traits putatively under abiotic selection to those controlling traits putatively under biotic selection.ResultsWe find no differences in the fraction of antagonistic QTL in traits under abiotic and biotic selection, suggesting similar consistency in selection pressure on these two types of traits. However, we find that QTL controlling traits under biotic selection have a larger effect size than those under abiotic selection, supporting our hypothesis that QTL of large effect are more commonly detected in traits under biotic selection than in traits under abiotic selection. For traits under both abiotic and biotic selection, we find a large number of QTL of large effect, with 10.7% of all QTLs detected controlling more than 20% of the variance in phenotype.ConclusionThese results suggest that mutations of large effect are more common in adaptive landscapes strongly determined by biotic forces, but that these types of adaptive landscapes do not result in a higher fraction of mutations acting in opposite directions. The high number of QTL of large effect detected shows that QTL of large effect are more common than predicted by the infinitesimal model of genetic adaptation.

Processes Influencing the Duration and Decline of Epizootics in Schistocephalus solidus

The interplay of intermediate host fish and plerocercoids of diphyllobothriidean cestodes results in epizootics that are deceptively simple, but conceal complex biotic and abiotic interactions shaping each event independently. Although general descriptions of epizootics and some details of biotic interactions between enemies are known, much remains to be discovered about the abiotic and biotic forces and their interactions driving epizootics. This study shows that the duration of an epizootic of Schistocephalus solidus was sustained by high prevalence, mean intensity, and PI (parasite index-parasite : host biomass ratio) levels among young-of-the-year and 1-yr-old threespine sticklebacks. Many infections and most parasite growth in young-of-the-year fish apparently occurred under the ice during the winter. Few new infections appear to have occurred among 1-yr-old fish, which may live 2 yr and sometimes 3 yr. The decline of the epizootic occurred as the recruitment of 1 to 2-yr-old hosts decreased significantly, followed by reduced infections of young-of-the-year fish. Thus, a major factor influencing parasite population dynamics was reduced transmission (probability of infection) as a result of overwinter host mortality among 1-yr-old fish. Mega-epizootics, named and described herein, appear to represent a "perfect storm" phenomenon dependent on a particular and rare combination of circumstances. Less extreme and more gradual epizootics may be more common and play out in myriad ways, because of complex abiotic and biotic factors influencing both parasite and host populations. The interplay of parasite and host resulting in reciprocal effects upon one another occurs during both the emergence and decline phases of an epizootic.

Microbial community structure across the tree of life in the extreme Río Tinto

Understanding biotic versus abiotic forces that shape community structure is a fundamental aim of microbial ecology. The acidic and heavy metal extreme Río Tinto (RT) in southwestern Spain provides a rare opportunity to conduct an ecosystem-wide biodiversity inventory at the level of all three domains of life, because diversity there is low and almost exclusively microbial. Despite improvements in high-throughput DNA sequencing, environmental biodiversity studies that use molecular metrics and consider entire ecosystems are rare. These studies can be prohibitively expensive if domains are considered separately, and differences in copy number of eukaryotic ribosomal RNA genes can bias estimates of relative abundances of phylotypes recovered. In this study we have overcome these barriers (1) by targeting all three domains in a single polymerase chain reaction amplification and (2) by using a replicated sampling design that allows for incidence-based methods to extract measures of richness and carry out downstream analyses that address community structuring effects. Our work showed that combined bacterial and archaeal richness is an order of magnitude higher than eukaryotic richness. We also found that eukaryotic richness was highest at the most extreme sites, whereas combined bacterial and archaeal richness was highest at less extreme sites. Quantitative community phylogenetics showed abiotic forces to be primarily responsible for shaping the RT community structure. Canonical correspondence analysis revealed co-occurrence of obligate symbionts and their putative hosts that may contribute to biotic forces shaping community structure and may further provide a possible mechanism for persistence of certain low-abundance bacteria encountered in the RT.

An invasive plant–fungal mutualism reduces arthropod diversity

Ecological theory holds that competition and predation are the most important biotic forces affecting the composition of communities. Here, we expand this framework by demonstrating that mutualism can fundamentally alter community and food web structure. In large, replicated field plots, we manipulated the mutualism between a dominant plant (Lolium arundinaceum) and symbiotic fungal endophyte (Neotyphodium coenophialum). The presence of the mutualism reduced arthropod abundance up to 70%, reduced arthropod diversity nearly 20%, shifted arthropod species composition relative to endophyte-free plots and suppressed the biomass and richness of other plant species in the community. Herbivorous arthropods were more strongly affected than carnivores, and for both herbivores and carnivores, effects of the mutualism appeared to propagate indirectly via organisms occurring more basally in the food web. The influence of the mutualism was as great or greater than previously documented effects of competition and predation on arthropod communities. Our work demonstrates that a keystone mutualism can significantly reduce arthropod biodiversity at a broad community scale.

Separate and interacting effects of deer florivory and salinity stress on iris herbivores

Most herbivore studies focus on foliage feeders, despite the potentially strong impact of florivores on plant fitness and ecological communities. Vertebrate florivory is particularly noteworthy because it destroys the inflorescence, and may affect the community of floral arthropods. Strong biotic forces such as florivory can be influenced by abiotic forces. For example, field studies indicate that white‐tailed deer Odocoileus virginianus consume up to 90% of all flowers produced in Iris hexagona populations that inhabit high salinity wetlands, whereas freshwater iris populations suffer relatively little damage. We conducted a common garden experiment to investigate the potential effects of vertebrate florivory and salinity on two iris herbivores, the leafminer Cerodontha iridiphora, and the verbena bud moth, Endothenia hebesana. We administered four levels of florivory: zero (control), one, two, and three flowers removed per stalk, and applied three levels of salinity (0, 4 and 8 parts per thousand NaCl) in a full‐factorial design. Florivory and salinity had separate and interacting effects on herbivore performance. Salinity was more detrimental to herbivores than florivory, which was beneficial at low levels. A positive effect of moderate salinity on sexual reproduction was reversed by florivory. Our research reveals complex interactions between environmental stress and florivory, an understudied form of herbivory. These interactions have implications for wetland ecological communities that are increasingly exposed to biotic and abiotic stress.

Incumbency, diversity, and latitudinal gradients

Physical environmental factors have been seen as paramount in determining many large-scale biodistributional patterns in time and space. Although this is probably correct for many situations, this view has become so pervasive that it has led to the neglect of the role of biotic interactions in setting large-scale diversity patterns. (In this paper diversity denotes taxonomic richness.) New approaches to this perennial debate on the roles of physical and biotic forces in paleoecology and macroevolution are needed, and here we explore an argument for the role of incumbency or priority effects in the dynamics behind the most dramatic spatial pattern in biodiversity, the latitudinal diversity gradient.

Test of local adaptation to biotic interactions and soil abiotic conditions in the ant-tended Chamaecrista fasciculata (Fabaceae)

Few previous studies have assessed the role of herbivores and the third trophic level in the evolution of local adaptation in plants. The overall objectives of this study were to determine (1) whether local adaptation is present in the ant-defended plant, Chamaecrista fasciculata, and (2) the contribution of ant-plant-herbivore interactions and soil source to such adaptation. We used three C. fasciculata populations and performed both a field and a greenhouse experiment. The first involved reciprocally transplanting C. fasciculata seedlings from each population-source to each site, and subsequently applying one of three treatments to one-third of the seedlings of each population-source at each site: control, reduced ant density and reduced folivory. The greenhouse experiment involved reciprocal transplants of population-sources with soil sources to test for a soil-source effect on flower production and local adaptation to soil conditions. Field results showed that ant and herbivore treatments reduced ant density (increasing folivory) and herbivore damage relative to controls, respectively; however, these manipulations did not impact C. fasciculata reproduction or the likelihood of survival. In contrast, greenhouse results showed that soil source significantly affected flower production. Overall, plants in both experiments, regardless of population-source, always had higher reproductive output at one specific site. Native populations did not outperform nonnative ones, causing us to reject the hypothesis of local adaptation. The absence of treatment effects on plant reproduction and the likelihood of survival suggest a limited effect of ants and folivores on C. fasciculata fitness and local adaptation during the study year. Temporally inconsistent effects of biotic forces across years, coupled with the young age of populations, relative proximity of populations and possible counter effects of seed predators may reduce the likelihood of local adaptation in the populations studied.

People as Ecological Participants in Ecological Restoration

Abstract In 1987, Bradshaw proposed that ecological restoration is the ultimate “acid test” of our understanding the functioning of ecosystems (Bradshaw 1987). Although this concept is widely supported academically, how it can be applied by restoration practitioners is still unclear. This is an issue not limited to Bradshaw’s acid test, but moreover, reflects a general difficulty associated with the polarization between conceptual restoration (restoration ecology) and practical restoration (ecological restoration), where each has functioned to certain degree in isolation of the other. Outside of the more obvious pragmatic reasons for the relative independence between ecological restoration and restoration ecology, we propose that a more contentious explanation is that the approach taken toward understanding ecosystem development in restoration ecology is tangential to what actually takes place in ecological restoration. Current paradigms assume that the process of ecosystem development in restoration should follow the developmental trajectories suggested by classical ecological succession models. However, unlike these models, ecosystem development in restoration is, at least initially, largely manipulated by people, rather than by abiotic and biotic forces alone. There has been little research undertaken to explore how restoration activities impact upon or add to the extant ecological processes operating within a restoration site. Consequently, ecological restoration may not be so much an acid test of our understanding the functioning of ecosystems, but rather, an acid test of our understanding mutually beneficial interactions between humans and ecosystems.