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There is strong trait dependence in species‐level responses to environmental change and their cascading effects on ecosystem functioning. However, there is little understanding of whether intraspecific trait variation (ITV) can also be an important mechanism mediating environmental effects on ecosystem functioning. This is surprising, given that global change processes such as habitat fragmentation and the creation of forest edges drive strong trait shifts within species. On 20 islands in the Thousand Island Lake, China, we quantified intraspecific leaf trait shifts of a widely distributed shrub species, Vaccinium carlesii, in response to habitat fragmentation. Using a reciprocal transplant decomposition experiment between forest edge and interior on 11 islands with varying areas, we disentangled the relative effects of intraspecific leaf trait variation versus altered environmental conditions on leaf decomposition rates in forest fragments. We found strong intraspecific variation in leaf traits in response to edge effects, with a shift toward recalcitrant leaves with low specific leaf area and high leaf dry matter content from forest interior to the edge. Using structural equation modeling, we showed that such intraspecific leaf trait response to habitat fragmentation had translated into significant plant afterlife effects on leaf decomposition, leading to decreased leaf decomposition rates from the forest interior to the edge. Importantly, the effects of intraspecific leaf trait variation were additive to and stronger than the effects from local environmental changes due to edge effects and habitat loss. Our experiment provides the first quantitative study showing that intraspecific leaf trait response to edge effects is an important driver of the decrease in leaf decomposition rate in fragmented forests. By extending the trait‐based response–effect framework toward the individual level, intraspecific variation in leaf economics traits can provide the missing functional link between environmental change and ecological processes. These findings suggest an important area for future research on incorporating ITV to understand and predict changes in ecosystem functioning in the context of global change. [ABSTRACT FROM AUTHOR]

The impact of multitrophic interactions on metacommunity structure, despite extensive theory and modeling/manipulative studies, has remained largely unexplored within naturally occurring metacommunities. We investigated the impacts of mutualistic partners and predators on a butterfly metacommunity, as well as the impacts that local and landscape characteristics have across three trophic levels: flowering plants, butterflies, and butterfly predators. Using data for butterfly diversity/richness, flowering plant diversity/richness, and butterfly predation (on clay butterfly models) across 15 grassland sites, we posed 3 questions. (1) How do mutualist metacommunity structure, predation pressure, and local and regional habitat characteristics affect butterfly metacommunity structure? (2) How do local and regional habitat characteristics affect flowering plant metacommunity structure? (3) How do local and regional habitat characteristics affect predation pressure? Floral diversity and richness had a positive effect on butterfly diversity and richness (Question 1). Site size positively affected floral diversity and richness (Question 2), and through this relationship site size had an indirect positive effect on butterfly diversity and richness (Question 1). In contrast to previous work, no other variables impacted butterfly diversity/richness. This result was particularly surprising for predation pressure: Our results suggest that within our study system, butterfly community diversity and richness are not strongly impacted by predation. Predator attacks occurred more in larger and more isolated sites (Question 3), suggesting that predators respond more strongly to landscape characteristics than abundance or diversity of butterfly prey species. This decoupling of predation pressure and butterfly communities suggests that conserving and restoring healthy predator populations may not negatively impact butterfly communities. If diverse plant communities are maintained, even small and isolated habitat patches can be valuable for butterfly conservation, which may influence reserve design and habitat restoration strategies. [ABSTRACT FROM AUTHOR]

Recent discussions of model selection and multimodel inference highlight a general challenge for researchers: how to convey the explanatory content of a hypothesized model or set of competing models clearly. The advice from statisticians for scientists employing multimodel inference is to develop a well‐thought‐out set of candidate models for comparison, though precise instructions for how to do that are typically not given. A coherent body of knowledge, which falls under the general term causal analysis, now exists for examining the explanatory scientific content of candidate models. Much of the literature on causal analysis has been recently developed, and we suspect may not be familiar to many ecologists. This body of knowledge comprises a set of graphical tools and axiomatic principles to support scientists in their endeavors to create "well‐formed hypotheses," as statisticians are asking them to do. Causal analysis is complementary to methods such as structural equation modeling, which provides the means for evaluation of proposed hypotheses against data. In this paper, we summarize and illustrate a set of principles that can guide scientists in their quest to develop explanatory hypotheses for evaluation. The principles presented in this paper have the capacity to close the communication gap between statisticians, who urge scientists to develop well‐thought‐out coherent models, and scientists, who would like some practical advice for exactly how to do that. [ABSTRACT FROM AUTHOR]

We explain how to obtain a generalized maximum‐likelihood chi‐square statistic,XML2, and a full‐model Akaike Information Criterion (AIC) statistic for piecewise structural equation modeling (SEM); that is, structural equations without latent variables whose causal topology can be represented as a directed acyclic graph (DAG). The full piecewise SEM is decomposed into submodels as a Markov network, each of which can have different distributional assumptions or functional links and that can be modeled by any method that produces maximum‐likelihood parameter estimates. The generalized XML2 is a function of the difference in the maximum likelihoods of the model and its saturated equivalent and the full‐model AIC is calculated by summing the AIC statistics of each of the submodels. [ABSTRACT FROM AUTHOR]

Abstract: Here we assess the impact of geographically dependent (latitude, longitude, and altitude) changes in bioclimatic (temperature, precipitation, and primary productivity) variability on fungal fruiting phenology across Europe. Two main nutritional guilds of fungi, saprotrophic and ectomycorrhizal, were further separated into spring and autumn fruiters. We used a path analysis to investigate how biogeographic patterns in fungal fruiting phenology coincided with seasonal changes in climate and primary production. Across central to northern Europe, mean fruiting varied by approximately 25 d, primarily with latitude. Altitude affected fruiting by up to 30 d, with spring delays and autumnal accelerations. Fruiting was as much explained by the effects of bioclimatic variability as by their large‐scale spatial patterns. Temperature drove fruiting of autumnal ectomycorrhizal and saprotrophic groups as well as spring saprotrophic groups, while primary production and precipitation were major drivers for spring‐fruiting ectomycorrhizal fungi. Species‐specific phenology predictors were not stable, instead deviating from the overall mean. There is significant likelihood that further climatic change, especially in temperature, will impact fungal phenology patterns at large spatial scales. The ecological implications are diverse, potentially affecting food webs (asynchrony), nutrient cycling and the timing of nutrient availability in ecosystems. [ABSTRACT FROM AUTHOR]

Knowledge of the density-dependent processes that regulate animal populations is key to understanding, predicting, and conserving populations. In migratory birds, density-dependence is most often studied during the breeding season, yet we still lack a robust understanding of the reproductive traits through which density influences individual reproductive success. We used 27-yr of detailed, individual-level productivity data from an island-breeding population of Savannah sparrows Passerculus sandwichensis to evaluate effects of local and total annual population density on female reproductive success. Local density (number of neighbors within 50 m of a female's nest) had stronger effects on the number of young fledged than did total annual population density. Females nesting in areas of high local density were more likely to suffer nest predation and less likely to initiate and fledge a second clutch, which led to fewer young fledged in a season. Fledging fewer young subsequently decreased the likelihood of a female recruiting offspring into the breeding population in a subsequent year. Collectively, these results provide insight into the scale and reproductive mechanisms mediating density-dependent reproductive success and fitness in songbirds. [ABSTRACT FROM AUTHOR]

Nitrogen (N) and phosphorus (P) concentrations are elevated in many freshwater systems, stimulating breakdown rates of terrestrially derived plant litter; however, the relative importance of N and P in driving litter breakdown via microbial and detritivore processing are not fully understood. Here, we determined breakdown rates of two litter species, Acer rubrum (maple) and Rhododendron maximum (rhododendron), before (PRE) and during two years (YR1, YR2) of experimental N and P additions to five streams, and quantified the relative importance of hypothesized factors contributing to breakdown. Treatment streams received a gradient of P additions (low to high soluble reactive phosphorus [SRP]; ~10-85 μg/L) crossed with a gradient of N additions (high to low dissolved inorganic nitrogen [DIN]; ~472-96 μg/L) to achieve target molar N:P ratios ranging from 128 to 2. Litter breakdown rates increased above pre-treatment levels by an average of 1.1-2.2× for maple, and 2.7-4.9× for rhododendron in YR1 and YR2. We used path analysis to compare fungal biomass, shredder biomass, litter stoichiometry (nutrient content as C:N or C:P), discharge, and streamwater temperature as predictors of breakdown rates and compared models containing streamwater N, P or N þ P and litter C:N or C:P using model selection criteria. Litter breakdown rates were predicted equally with either streamwater N or P (R² = 0.57). In models with N or P, fungal biomass, litter stoichiometry, discharge, and shredder biomass predicted breakdown rates; litter stoichiometry and fungal biomass were most important for model fit. However, N and P effects may have occurred via subtly different pathways. Litter N content increased with fungal biomass (N-driven effects) and litter P content increased with streamwater P availability (P-driven effects), presumably via P storage in fungal biomass. In either case, the effects of N and P through these pathways were associated with higher shredder biomass and breakdown rates. Our results suggest that N and P stimulate litter breakdown rates via mechanisms in which litter stoichiometry is an important nexus for associated microbial and detritivore effects. [ABSTRACT FROM AUTHOR]

Water flow gradients have been linked to phenotypic differences and swimming performance across a variety of fish assemblages. However, the extent to which water motion shapes patterns of phenotypic divergence within species remains unknown. We tested the generality of the functional relationship between swimming morphology and water flow by exploring the extent of fin and body shape polymorphism in 12 widespread species from three families (Acanthuridae, Labridae, Pomacentridae) of pectoral-fin swimming (labriform) fishes living across localized wave exposure gradients. The pectoral fin shape of Labridae and Acanthuridae species was strongly related to wave exposure: individuals with more tapered, higher aspect ratio (AR) fins were found on windward reef crests, whereas individuals with rounder, lower AR fins were found on leeward, sheltered reefs. Three of seven Pomacentridae species showed similar trends, and pectoral fin shape was also strongly related to wave exposure in pomacentrids when fin aspect ratios of three species were compared across flow habitats at very small spatial scales (,100 m) along a reef profile (reef slope, crest, and back lagoon). Unlike fin shape, there were no intraspecific differences in fish body fineness ratio across habitats or depths. Contrary to our predictions, there was no pattern relating species' abundances to polymorphism across habitats (i.e., abundance was not higher at sites where morphology is better adapted to the environment). This suggests that there are behavioral and/or physiological mechanisms enabling some species to persist across flow habitats in the absence of morphological differences. We suggest that functional relationships between swimming morphology and water flow not only structure species assemblages, but are yet another important variable contributing to phenotypic differences within species. The close links between fin shape polymorphism and local water flow conditions appear to be important for understanding species' distributions as well as patterns of diversification across environmental gradients. [ABSTRACT FROM AUTHOR]

The factors that promote invasive behavior in introduced plant species occur across many scales of biological and ecological organization. Factors that act at relatively small scales, for example, the evolution of biological traits associated with invasiveness, scale up to shape species distributions among different climates and habitats, as well as other characteristics linked to invasion, such as attractiveness for cultivation (and by extension propagule pressure). To identify drivers of invasion it is therefore necessary to disentangle the contribution of multiple factors that are interdependent. To this end, we formulated a conceptual model describing the process of invasion of central European species into North America based on a sequence of ''drivers.'' We then used confirmatory path analysis to test whether the conceptual model is supported by a statistical model inferred from a comprehensive database containing 466 species. The path analysis revealed that naturalization of central European plants in North America, in terms of the number of North American regions invaded, most strongly depends on residence time in the invaded range and the number of habitats occupied by species in their native range. In addition to the confirmatory path analysis, we identified the effects of various biological traits on several important drivers of the conceptualized invasion process. The data supported a model that included indirect effects of biological traits on invasion via their effect on the number of native range habitats occupied and cultivation in the native range. For example, persistent seed banks and longer flowering periods are positively correlated with number of native habitats, while a stress-tolerant life strategy is negatively correlated with native range cultivation. However, the importance of the biological traits is nearly an order of magnitude less than that of the larger scale drivers and highly dependent on the invasion stage (traits were associated only with native range drivers). This suggests that future research should explicitly link biological traits to the different stages of invasion, and that a failure to consider residence time or characteristics of the native range may seriously overestimate the role of biological traits, which, in turn, may result in spurious predictions of plant invasiveness. [ABSTRACT FROM AUTHOR]

When effective immune defenses against parasites are costly and resources limited, individuals are expected to alter their investment in immunity in response to the risk of infection. As an ecological factor that can affect both food abundance and parasite exposure, host density can play an important role in host immunity and host-parasite interactions. High levels of intraspecific competition for food and social stress at high host density may diminish immune defenses and increase host susceptibility to parasites. At the same time, for contagious and environmentally transmitted parasites, parasite exposure often increases with host density, whereas in mobile parasites that actively search for hosts, parasite exposure can decrease with host density due to the "encounter-dilution effect." To unravel these multiple and potentially opposing effects of host density on immunity, we manipulated density of the common lizard Zootoca vivipara and measured local inflammation in response to PHA injection and levels of infestation by the tick Ixodes ricinus, a mobile ectoparasite for which we expected an encounter-dilution effect to occur. Local inflammation strongly decreased with lizard density in adults, but not in yearlings. Tick infestation (abundance and prevalence) was negatively correlated with lizard density in both age classes. Using path analyses, we found independent, direct negative density feedbacks on immunity and parasite exposure in adults, supporting the hypothesis of energy constraints and/or physiological stress acting on immunity at high density. In contrast, for yearlings, the best path model showed that density diluted exposure to parasites, which themselves down-regulated immune defenses in lizards. These results highlight the importance of investigating the pathways among host density, host immunity, and parasite infestation, while accounting for relevant individual traits such as age. [ABSTRACT FROM AUTHOR]

Temporal changes in phenological traits arising as a consequence of recent rapid environmental change have been widely demonstrated in animal populations. Increasingly, studies are seeking to understand the impact of changes in such traits on individual fitness and population dynamics, with the ultimate aim of predicting population persistence or extinction under different climate scenarios. Here, we examined the effects of environmental change on maternal reproductive traits in a wild population of red deer (Cervus elaphus) and sought to explain why, despite a rapid advance in offspring birth dates, we observed no apparent consequences for offspring fitness. By using path analysis, we identified both direct and indirect paths along which changes in environmental conditions affected birth date, birth mass, juvenile survival, and female fecundity. In general, warmer temperatures were associated with earlier birth dates and greater birth mass, and higher rainfall was associated with reduced juvenile survival and reduced female fecundity. We also examined concurrent effects of population density, maternal age, and reproductive history, and found that temporal stasis in average trait values, at least in part, could be explained by antagonistic roles of direct and indirect effects of changing climate and increasing population density. Identification of the many mechanisms that contribute to the dynamics of phenotypic traits is challenging; this study demonstrates the need to consider both climatic and demographic variation in order to understand the fitness consequences of changes in phenological traits. [ABSTRACT FROM AUTHOR]

The interplay between density and climate in shaping the dynamics of herbivore populations is widely acknowledged, and current research is fueled by the identification of mechanisms underlying their effects on individuals and populations. We assessed whether forage availability mediated the effects of density and winter climate on body mass of whitetailed deer (Odocoileus virginianus) yearlings by experimentally reducing deer density to 7.5 and 15 deer/km2 during eight growing seasons, and by using causal (graphical) hierarchical models and Bayesian hierarchical modeling to assess relationships. The abundance of preferred forage decreased with deer density and varied quadratically (positive parabola) with winter North Atlantic Oscillation (NAO), whereas the fall mass of yearlings increased with forage abundance and spring mass. Fall mass did not differ between experimentally reduced deer densities, yet experimental yearlings were 30% heavier than yearlings harvested at ambient densities. Hence, forage abundance simultaneously mediated the effects of density and climate on fall body mass, which was also infiuenced by carry-over effects of spring body mass. Our findings increase our ability to anticipate how temperate large herbivores will respond to ongoing changes in intrinsic (e.g., large-herbivore density) and extrinsic (e.g., climate) factors. [ABSTRACT FROM AUTHOR]

There is increasing evidence that environmental conditions experienced early in life can markedly affect an organism's life history, but the pathways by which early environment influences adult phenotype are poorly known. We used long-term data from two roe deer (Capreolus capreolus) populations (Chizé and Trois-Fontaines, France) to investigate the direct and indirect (operating through fawn body mass) effects of environmental conditions during early life on adult body mass. We found that environmental conditions (population size and spring temperatures) around birth influenced body mass of adult females through both direct and indirect effects in both populations. The occurrence of direct effects means that, for a given fawn body mass, adult female mass decreases with adverse conditions in early life. In contrast, we found no evidence for direct effects of early-life conditions on adult body mass of males, suggesting the existence of sex-specific long-term responses of body mass to stressful early conditions. Our results provide evidence that early environmental conditions influence the adult phenotype through persistent effects over the body development in wild mammal populations. [ABSTRACT FROM AUTHOR]

Classical path analysis is a statistical technique used to test causal hypotheses é involving multiple variables without latent variables, assuming linearity, multivariate normality, and a sufficient sample size. The d-separation (d-sep) test is a generalization of path analysis that relaxes these assumptions. Although model selection using Akaike's information criterion (AIC) is well established for classical path analysis, this model selection technique has not yet been developed for d-sep tests. In this paper, I explain how to use the AIC statistic for d-sep tests, give a worked example, and include instructions (supplemental material) to implement the analysis in the R computing language. [ABSTRACT FROM AUTHOR]

While the effects of contemporaneous local environment on species richness have been repeatedly documented, much less is known about historical effects, especially over large temporal scales. Using fen sites in the Western Carpathian Mountains with known radiocarbon-dated ages spanning Late Glacial to modern times (16?975-270 cal years before 2008), we have compiled richness data from the same plots for three groups of taxa with contrasting dispersal modes: (1) vascular plants, which have macroscopic propagules possessing variable, but rather low, dispersal abilities; (2) bryophytes, which have microscopic propagules that are readily transported long distances by air; and (3) terrestrial and freshwater mollusks, which have macroscopic individuals with slow active migration rates, but which also often possess high passive dispersal abilities. Using path analysis we tested the relationships between species richness and habitat age, area, isolation, and altitude for these groups. When only matrix-derived taxa were considered, no significant positive relation was noted between species richness and habitat size or age. When only calcareous-fen specialists were considered, however, habitat age was found to significantly affect vascular plant richness and, marginally, also bryophyte richness, whereas mollusk richness was significantly affected by habitat area. These results suggest that in inland insular systems only habitat specialist (i.e., interpatch disperser and/or relict species) richness is influenced by habitat age and/or area, with habitat age becoming more important as species dispersal ability decreases. [ABSTRACT FROM AUTHOR]

This paper describes how to test, and potentially falsify, a multivariate causal hypothesis involving only observed variables (i.e., a path analysis) when the data have a hierarchical or multilevel structure, when different variables are potentially defined at different levels of such a hierarchy, and when different variables have different sampling distributions. The test is a generalization of Shipley's d-sep test and can be conducted using standard statistical programs capable of fitting generalized mixed models. [ABSTRACT FROM AUTHOR]

Biotic interactions, such as competition and herbivory, can limit plant species ranges to a subset of edaphically suitable habitats, termed the realized niche. Here we explored the role that herbivores play in restricting the niche of serpentine ecotypes of the native California annual Collinsia sparsiflora. We planted seeds from four populations into a range of natural field environments that varied in the presence/absence of naturally occurring C. sparsiflora and in predicted suitability for growth and survival of the serpentine ecotype of C. sparsifIora. Path analysis was then used to model the direct and herbivore-mediated indirect effects of environmental variables on the survival of C. sparsiflora serpentine ecotypes. We found that C. sparsiflora received more herbivory when planted into areas where serpentine ecotypes of C. sparsifIora were not predicted to persist, and that increased herbivory was associated with decreased survival, suggesting that herbivores may limit the distribution of C. sparsiflora serpentine ecotypes. Additionally, we demonstrated that edaphic environmental variables impacted the survival of C. sparsiflora serpentine ecotypes both directly and indirectly, by altering interactions with herbivores. These indirect effects were probably trait-mediated and probably occurred because edaphic factors may influence plant traits that, in turn, alter attractiveness to herbivores. Although the magnitude of direct effects exceeded the magnitude of indirect effects, many strong herbivore-mediated indirect effects were detected. Thus, interactions between the abiotic environment and insect herbivory contributed to restricting the niche of C. sparsiflora serpentine ecotypes to a subset of available habitat. [ABSTRACT FROM AUTHOR]

Interactions among predators that prey on each other and are potential competitors for shared prey (intraguild [IG] predators) are widespread in terrestrial ecosystems and have the potential to strongly influence the dynamics of terrestrial food webs. Ants and spiders are abundant and ubiquitous terrestrial IG predators, yet the strength and consequences of interactions between them are largely unknown. In the leaf-litter food web of a deciduous forest in Kentucky (USA), we tested the direct and indirect effects of ants on spiders and a category of shared prey (Collembola) by experimentally subsidizing ants in open plots in two field experiments. In the first experiment, ant activity was increased, and the density of ants in the litter was doubled, by placing carbohydrate and protein baits in the center of each plot. Gnaphosa spiders were almost twice as abundant and Schizocosa spiders were half as abundant in baited plots relative to controls. There were more tomocerid Collembola in baited plots, suggesting possible indirect effects on Collembola caused by ant-spider interactions. The second experiment, in which screening of two mesh sizes selectively excluded large and small worker ants from a sugar bait, revealed that the large ants, primarily Camponotus, could alone induce similar effects on spiders. Gnaphosa biomass density was almost twice as high in the plots where large ants were more active, whereas Schizocosa biomass density was reduced by half in these plots. Although tomocerid densities did not differ between treatments, tomocerid numbers were negatively correlated with the activity of Formica, another large ant species. Path analysis failed to support the hypothesis that the ant Camponotus indirectly affected tomocerid Collembola through effects on densities of spiders. However, path analysis also revealed other indirect effects of Camponotus affecting tomocerids. These results illustrate the complexity of interactions between and within two major IG predator groups with disparate predatory behaviors, complexities that will have consequences for functioning of the forest-floor food web. [ABSTRACT FROM AUTHOR]

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community. [ABSTRACT FROM AUTHOR]

Recent work has identified a worldwide "economic" spectrum of correlated leaf traits that affects global patterns of nutrient cycling and primary productivity and that is used to calibrate vegetation-climate models. The correlation patterns are displayed by species from the arctic to the tropics and are largely independent of growth form or phylogeny. This generality suggests that unidentified fundamental constraints control the return of photosynthates on investments of nutrients and dry mass in leaves. Using novel graph theoretic methods and structural equation modeling, we show that the relationships among these variables can best be explained by assuming (1) a necessary trade-off between allocation to structural tissues versus liquid phase processes and (2) an evolutionary trade- off between leaf photosynthetic rates, construction costs, and leaf longevity. [ABSTRACT FROM AUTHOR]

An orientation component leads to directionally biased paths, with major consequences in animal population redistribution. Classical orientation analyses, which focus on the overall direction of motion, are useless for detecting such a component when the preferred direction is not common to the whole population, but differs from one path to another. In-depth path analyses are required in this case. They consist of determining whether paths are more suitably represented as biased or unbiased random walks. The answer is not easy because most animals' paths show some forward persistence propensity that acts as a purely local directional bias and, hence, blurs the possible occurrence of an additional, consistent bias in a preferred direction. I highlight the key differences between biased and unbiased random walks and the different ways orientation mechanisms can generate a consistent directional bias. 1 then examine the strength and weakness of the available methods likely to detect it. Finally, I introduce a new procedure based on the backward evolution of the beeline distance, from the end of the path, which might correspond to a goal toward which the animal orients itself, to each of the animal's preceding locations. This new procedure proves to be very efficient, as it requires only a small sample of short paths for detecting a possible orientation component. [ABSTRACT FROM AUTHOR]

From a functional perspective, changes in abundance, and ultimately species replacement, during succession are a consequence of integrated suites of traits conferring different relative ecological advantages as the environment changes over time. Here we use structural equations to model the interspecific relationships between these integrated functional traits using 34 herbaceous species from a Mediterranean old-field succession and thus quantify the notion of a plant strategy. We measured plant traits related to plant vegetative and reproductive size, leaf functioning, reproductive phenology, seed mass, and production on 15 individuals per species monitored during one growing season. The resulting structural equation model successfully accounts for the pattern of trait covariation during the first 45 years post-abandonment using just two forcing variables: time since site abandonment and seed mass; no association between time since field abandonment and seed mass was observed over these herbaceous stages of secondary succession. All other predicted traits values are determined by these two variables and the cause-effect linkage between them. Adding pre-reproductive vegetative mass as a third forcing variable noticeably increased the predictive power of the model. Increasing the time after abandonment favors species with increasing life span and pre-reproductive biomass and decreasing specific leaf area. Allometric coefficients relating vegetative and reproductive components of plant size were in accordance with allometry theory. The model confirmed the trade-off between seed mass and seed number. Maximum plant height and seed mass were major determinants of reproductive phenology. Our results show that beyond verbal conceptualization, plant ecological strategies can be quantified and modeled. [ABSTRACT FROM AUTHOR]

Pollination by animals is critical to sexual reproduction of most angiosperms. However, little is known about variation in pollination service to single plant species. We report results of a long-term study of Ipomopsis aggregata, a semelparous montane herb whose flowers are visited by hummingbird and insect pollinators as well as "floral larcenists." We censused flower visitors over seven summers at permanent study sites separated by several hundred meters, and counted pollen delivered to flowers on a subset of plants observed for visitation. The species composition of the community of visitors varied significantly across years and within the flowering season; sites varied significantly only in the magnitude of parallel annual changes in the visitor community. Rates of flower visitation fluctuated over an order of magnitude or more. Variation in mean stigma pollen load among plants flowering in the same site and year was explained by a causal path model in which visitation rates by pollinators and larcenists had linear positive and negative effects, respectively. A simplified model including only pollinators explained almost as much variance as did the full model. However, qualitatively different parameter estimates were produced by an analogous causal model based on population means across site-year combinations. Discrepant results from within- and between-population levels of analysis suggest that pollen receipt is influenced by environmental factors that vary among sites and years, as well as by pollinator visit rates. We present a heuristic causal model that includes such factors, and we note its implications for ecological and evolutionary studies of pollination. [ABSTRACT FROM AUTHOR]

Obligate pollination mutualisms have been central to our understanding of the ecology and evolution of mutualisms. Although usually viewed as pairwise interactions, obligate mutualists also interact with other community members that may impact the mutualism. In this study, we examined the community context of the obligate mutualism between the plant Yucca filamentosa and its yucca moth pollinator Tegeticula cassandra to determine the relationships among plant traits, the community assemblage, and fruit set. In addition to attracting its pollinator, V. filanientosa in central Florida also attracts two insect florivores: the hemipteran Leptoglossus phyllopus and the beetle Hymenorus densus. We surveyed flowering V. filamentosa in 2001 and 2002 to determine the abundance of the three insect species throughout the flowering period. We also measured fruit set and four plant traits thought to be important in attracting the moth pollinator. Path analysis revealed that T. cassandra was the most important determinant of relative fruit set for both years. Peak pollinator abundance was significantly earlier than peak flowering. Consequently, we found a negative relationship between plant flowering date and pollinator abundance. Furthermore, taller plants also attracted more pollinators. Leptoglossus phyllopus had a significantly negative impact on relative fruit set in 2001 but not in 2002, whereas H. densus had no effect in either year. Abundance of the two florivores was more closely tied to peak flowering of V. filamentosa than was the peak abundance of the pollinator moths. Although both florivore species reached high densities on V. filamentosa, their effect on relative fruit set through floral abscission may be partially masked by the characteristically high abscission rates found in yuccas. As a result, florivores may be feeding on flowers that are already destined to be abscised. Similarly, pollinator moths may be preadapted to handle the effects of florivores because females lay eggs across multiple plants and flowers. Hence, the obligate mutualism between yuccas and yucca moths may be somewhat buffered from the effects of local community members depending on whether florivores substantially increase floral abscission rates from year to year. [ABSTRACT FROM AUTHOR]

Natural disturbance boundaries have fundamental influences on landscape function; however, quantitative examination of the factors influencing their formation has been limited. Boundary formation is the final component of a disturbance event trio (initiation, propagation, and cessation) and is least well understood. I examined the influences of five stand-level variables (tree age, tree density, basal area of susceptible trees, slope aspect, and slope angle) and space as a surrogate for the unmeasurable factors on standardized tree mortality (basal area dead/total basal area) using multiple causal path hypotheses with data from seven wildfire and 11 mountain pine beetle (MPB) boundaries in British Columbia, Canada. One hundred percent of fire sites and 91% of MPB sites showed strong support for a positive effect of the basal area of susceptible trees (as defined by their propensity to burn or to be attacked by MPB) on tree mortality (r = 0.21-0.81). In addition, seven of 11 MPB boundaries had mortality patterns that were affected only indirectly by the spatial location across the boundary. This indirect effect of space suggests that other unmeasured spatial factors were not responsible for the pattern of mortality. Thus, MPB dispersal limitation did not appear to be an important factor stopping the outbreaks at these seven sites. A population-based approach would be insufficient for understanding the location of the boundaries. On the other hand, mortality in all fire boundaries was directly affected by the spatial location aéross the boundary. In the context of multiple hypothesis testing of path models, this indicates that there was an important influence of spatially varying but unmodeled factors, such as weather and surface vegetation. Furthermore, path models that included other factors explicitly examined in this study (tree density, tree age, aspect, and slope) were rarely supported by the data, suggesting little contribution to boundary formation. These results demonstrate that the combined use of spatial plots, path analysis, and a priori multiple hypotheses can give important insight into the functioning of ecological processes. [ABSTRACT FROM AUTHOR]

Disease dynamics are governed by variation of individuals, species, and environmental conditions across space and time. In some cases, an alternate reservoir host amplifies pathogen loads and drives disease transmission to less competent hosts in a process called pathogen spillover. Spillover is frequently associated with multi‐host disease systems where a single species is more tolerant of infection and more competent in pathogen transmission compared to other hosts. Pathogen spillover must be driven by biotic factors, including host and community characteristics, yet biotic factors interact with the abiotic environment (e.g., temperature) to create disease. Despite its fundamental role in disease dynamics, the influence of the abiotic environment on pathogen spillover has seldom been examined. Improving our understanding of disease processes such as pathogen spillover hinges on disentangling the effects of interrelated biotic and abiotic factors over space and time. We applied 10 yr of fine‐scale microclimate, disease, and tree community data in a path analysis to investigate the relative influence of biotic and abiotic factors on pathogen spillover for the emerging infectious forest disease sudden oak death (SOD). Disease transmission in SOD is primarily driven by the reservoir host California bay laurel, which supports high foliar pathogen loads that spillover onto neighboring oak trees and create lethal canker infections. The foliar pathogen load and susceptibility of oaks is expected to be sensitive to forest microclimate conditions. We found that biotic factors of pathogen load and tree diversity had relatively stronger effects on pathogen spillover compared to abiotic microclimate factors, with pathogen load increasing oak infection and tree diversity reducing oak infection. Abiotic factors still had significant effects, with greater heat exposure during summer months reducing pathogen loads and optimal pathogen conditions during the wet season increasing oak infection. Our results offer clues to possible disease dynamics under future climate change where hotter and drier or warmer and wetter conditions could have opposing effects on pathogen spillover in the SOD system. Disentangling direct and indirect effects of biotic and abiotic factors affecting disease processes can provide key insights into disease dynamics including potential avenues for reducing disease spread and predicting future epidemics. [ABSTRACT FROM AUTHOR]

Moving beyond species count data is an essential step to better understand the effects of environmental perturbations on biodiversity and ecosystem functions, and to eventually better predict the strength and direction of those effects. Here, coupling an integrative path analysis approach with data from an extensive countrywide monitoring program, we tested the main spatial, environmental and anthropogenic drivers of change in the functional structure of aquatic macroinvertebrate communities along the entire Swiss Rhine river catchment. Functional structure was largely driven by inherent altitudinal variation influencing and cascading to regional scaled factors such as land use change and position in the riverine network, which, in turn, transformed local habitat structure variables. Those cascading effects across scales propagated through the biotic community, first affecting prey and, in turn, predators. Our results illustrate how seemingly less important local factors can act as essential transmission belts, propagating through direct and indirect pathways across scales to generate the specific context in which each functional group will strive or not, leading to characteristic landscape wide variations in functional community structure. [ABSTRACT FROM AUTHOR]

Moving beyond species count data is an essential step to better understand the effects of environmental perturbations on biodiversity and ecosystem functions, and to eventually better predict the strength and direction of those effects. Here, coupling an integrative path analysis approach with data from an extensive countrywide monitoring program, we tested the main spatial, environmental and anthropogenic drivers of change in the functional structure of aquatic macroinvertebrate communities along the entire Swiss Rhine river catchment. Functional structure was largely driven by inherent altitudinal variation influencing and cascading to regional scaled factors such as land use change and position in the riverine network, which, in turn, transformed local habitat structure variables. Those cascading effects across scales propagated through the biotic community, first affecting prey and, in turn, predators. Our results illustrate how seemingly less important local factors can act as essential transmission belts, propagating through direct and indirect pathways across scales to generate the specific context in which each functional group will strive or not, leading to characteristic landscape wide variations in functional community structure., (© 2019 by the Ecological Society of America.)