ALBERT

Description: Bacteria are essential for many ecosystem services but our understanding of factors controlling their functioning is incomplete. While biodiversity has been identified as an important driver of ecosystem processes in macrobiotic communities, we know much less about bacterial communities. Due to the high diversity of bacterial communities, high functional redundancy is commonly proposed as explanation for a lack of clear effects of diversity. The generality of this claim has, however, been questioned. We present the results of an outdoor dilution-to-extinction experiment with four lake bacterial communities. The consequences of changes in bacterial diversity in terms of effective number of species, phylogenetic diversity and functional diversity were studied for (i) bacterial abundance, (ii) temporal stability of abundance, (iii) nitrogen concentration, and (iv) multifunctionality. We observed a richness gradient ranging from 15 to 280 operational taxonomic units. Individual relationships between diversity and functioning ranged from negative to positive depending on lake, diversity dimension and aspect of functioning. Only between phylogenetic diversity and abundance did we find a statistically consistent positive relationship across lakes. A literature review of 24 peer-reviewed studies that used dilution-to-extinction to manipulate bacterial diversity corroborated our findings: about 25% found positive relationships. Combined, these results suggest that bacteria-driven community functioning is relatively resistant to reductions in diversity. This article is protected by copyright. All rights reserved.

Description: The degree of ecological specialization plays a crucial role in shaping the structure and functioning of communities. However, comparing specialization within and among groups of organisms is complicated due to methodological issues but also by conceptual and terminological inconsistencies. Environmental predictability has been considered a key determinant of specialization though empirical evidence is still limited. Fungi and their insect consumers provide a poorly studied but promising system to measure host specialization and test the predictability hypothesis. In this study we systematically sampled mushrooms in North European boreal forest, and reared total samples of fungivores colonizing the fruitbodies. Due to the unpredictable nature of mushrooms as a resource, low levels of host specialization can be predicted for these insects which have indeed widely been considered polyphagous. Contrary to expectations, majority of the studied fungus gnats were found not to exploit their host taxa indiscriminately. Not only were some mushroom taxa never colonised, but also the infestation rate of acceptable hosts differed in most of these fungivores. Gnat species themselves formed continua with respect to the estimates of the degree of specialization, derived from parametric individual-based analyses of presence-absence data. In most cases, host use was best explained by models in which the hosts were classified at genus level, with limited support to specialization to particular host species, families or orders. Indeed, most of the common fungivores appeared to preferentially use various species from one or a few mushroom genera while occasionally feeding on members of other host taxa. This pattern has likely evolved as a compromise between selective forces stemming from host unpredictability, and taxon-specific chemical profiles of the mushrooms. Our study highlights the multidimensional nature of ecological specialization: a high number of acceptable hosts does not preclude considerable discrimination among members of the available resource pool. Such situations can only be revealed by individual-based analyses capable of capturing differences in partner-to-partner interaction intensities. This article is protected by copyright. All rights reserved.

Description: Prey at risk of predation may experience stress and respond physiologically by altering their metabolic rates. Theory predicts that such physiological changes should alter prey nutrient demands from N-rich to C-rich macronutrients and shift the balance between maintenance and growth/reproduction. Theory further suggests that for ectotherms, temperature stands to exacerbate this stress. Yet, the interactive effects of predation stress and temperature stress on diet, metabolism, and survival of ectotherms are not well known. This knowledge gap was addressed with a laboratory study in which wild juvenile grasshoppers were collected, assigned to one of three groups, and raised at three different temperatures. All grasshoppers had access to equal quantities of two diets composed of opposite carbohydrate:protein ratios. Half of the individuals in each temperature group were exposed to predation risk cues from spider predators, while the other half were kept in risk free conditions. Grasshoppers consumed more carbohydrates when exposed to predation risk, but consumption favored greater protein intake as temperature increased. Moreover, the difference in carbohydrate intake between risk cue and risk free treatments diminished as temperature increased. Furthermore, variability between individual consumption patterns both within and between treatments decreased markedly as temperature increased, suggesting that higher temperatures promote more consistent individual consumption behaviors. Grasshoppers grew faster and larger as temperature increased, which translated into higher survival rates at higher temperatures. Warmer grasshoppers also did not alter their metabolic rates in response to predation risk cues, in contrast to colder grasshoppers. Digestive efficiency increased with temperature as well, further indicating that lower temperatures were much more stressful than higher temperatures for grasshoppers. The study shows that physiological responses of ectothermic herbivores to predation stress are highly plastic and temperature dependent, with higher temperatures promoting increased protein intake, growth, development, survival, and digestive efficiency relative to colder temperatures. These findings help to reconcile why dietary responses (proportion of protein vs. carbohydrate intake) to predation stress may vary among different prey taxa studied previously. This article is protected by copyright. All rights reserved.

Description: Contradictory evidence from biogeomorphological studies has increased the debate on the extent of lichen contribution to differential rock surface weathering in both natural and cultural settings. This study, undertaken in Côa Valley Archaeological Park, aimed at evaluating the effect of rock surface orientation on the weathering ability of dominant lichens. Hyphal penetration and oxalate formation at the lichen-rock interface were evaluated as proxies of physical and chemical weathering, respectively. A new protocol of pixel-based supervised image classification for the analysis of periodic acid-Schiff stained cross-sections of colonized schist revealed that hyphal spread of individual species was not influenced by surface orientation. However, hyphal spread was significantly higher in species dominant on north-west facing surfaces. An apparently opposite effect was noticed in terms of calcium oxalate accumulation at the lichen-rock interface, detected by Raman spectroscopy and complementary X-ray microdiffraction on south-east facing surfaces only. These results suggest that lichen-induced physical weathering may be most severe on north-west facing surfaces by means of an indirect effect of surface orientation on species abundance, and thus dependent on the species, whereas lichen-induced chemical weathering is apparently higher on south-east facing surfaces and dependent on micro-environmental conditions, giving only weak support to the hypothesis that lichens are responsible for the currently observed pattern of rock-art distribution in Côa Valley. Assumptions about the drivers of open-air rock-art distribution patterns elsewhere should also consider the micro-environmental controls of lichen-induced weathering, to avoid biased measures of lichen contribution to rock-art deterioration. This article is protected by copyright. All rights reserved.

Description: The majority of humanity now lives in cities or towns, with this proportion expected to continue increasing for the foreseeable future. As novel ecosystems, urban areas offer an ideal opportunity to examine multi-scalar processes involved in community assembly as well as the role of human activities in modulating environmental drivers of biodiversity. Although ecologists have made great strides in recent decades at documenting ecological relationships in urban areas, much remains unknown, and we still need to identify the major ecological factors, aside from habitat loss, behind the persistence or extinction of species and guilds of species in cities. Given this paucity of knowledge, there is an immediate need to facilitate collaborative, interdisciplinary research on the patterns and drivers of biodiversity in cities at multiple spatial scales. In this review, we introduce a new conceptual framework for understanding the filtering processes that mold diversity of urban floras and faunas. We hypothesize that the following hierarchical series of filters influence species distributions in cities: 1) regional climatic and biogeographical factors; 2) human facilitation; 3) urban form and development history; 4) socioeconomic and cultural factors; and 5) species interactions. In addition to these filters, life history and functional traits of species are important in determining community assembly and act at multiple spatial scales. Using these filters as a conceptual framework can help frame future research needed to elucidate processes of community assembly in urban areas. Understanding how humans influence community structure and processes will aid in the management, design, and planning of our cities to best support biodiversity. This article is protected by copyright. All rights reserved.

Description: The main goal of this study is to create a database that ultimately serves further studies on riparian vegetation and flow response guilds in the boreal region and on transferability of results across different regions. For this aim we compiled traits for all woody riparian species in northern Sweden that, directly or indirectly, underlie their responses to hydrological and hydraulic conditions, between October 2012 and April 2015. Consulted sources of information were diverse, ranging from scientific to informative and whose accuracy might or might not be verified. They were focused on particular or several traits and species from concrete areas to a worldwide perspective. Sources were characterized by different degrees of accessibility and showed a wide variety of descriptions, categorical and ordinal classifications, and numerical information for each trait. Our effort was to synthesize information for each trait from all sources into the common frame of our own database, following own defined criteria so that comparisons between species are congruent. Therefore, this data set is unique in that it comprehensively combines and homogenizes morphological, phenological, reproductive and ecological data for 59 woody, riparian, boreal species and from 118 sources of information, that would otherwise be scattered and hardly available. This article is protected by copyright. All rights reserved.

Description: Size variation within a population can influence the structure of ecosystem interactions, because ecological performance differs between individuals of different sizes. Although the impact of size variation in a predator species on the structure of interactions is well understood, our knowledge about how size variation in a prey species might modify the interactions between predators and prey is very limited. Here, by examining the interactions between predatory Hynobius retardatus salamander larvae and their prey, Rana pirica frog tadpoles, we investigated how large prey individuals affect the predation mortality of small prey conspecifics. First, in an experiment conducted in a field pond in which we manipulated the presence of salamanders and large tadpoles (i.e., large enough to protect them against salamander predation) with small tadpoles, we showed that in the presence of large tadpoles the mortality of small tadpoles from salamander predation was increased. On the basis of our observations of the activity of individuals, we hypothesized that active large tadpoles caused physical disturbances, which in turn caused the small tadpoles to move, and thus increased their encounter frequency with the predatory salamanders. To test this hypothesis, we conducted a laboratory experiment in small tanks with three players (i.e., one salamander as predator, one small tadpole as focal prey, and either a small or a large tadpole as the prospective movement inducer). In each tank, we manipulated the presence or absence of a movement inducer, and, when present, its size (large or small) and access (caged or uncaged) to the focal prey. In the presence of a large, uncaged movement inducer, the focal prey was more active and suffered from higher predation mortality compared with the other treatments, because the large movement inducer (unlike a small movement inducer) moved actively and, when uncaged, could stimulate movement of the focal prey through direct contact. The results indicated that high activity of large prey individuals and the resulting behavioral interactions with small conspecifics via direct contact indirectly increased the mortality of the small prey. This article is protected by copyright. All rights reserved.

Description: The phyllosphere (comprising the leaf surface and interior) is one of the world's largest microbial habitats and is host to an abundant and diverse array of bacteria. Nonetheless, the degree to which bacterial communities are benign, harmful, or beneficial to plants in situ is unknown. We tested the hypothesis that the net effect of reducing bacterial abundance and diversity would vary substantially among host species (from harmful to beneficial) and this would be strongly mediated by soil resource availability. To test this, we monitored tree seedling growth responses to commercial antibiotics among replicated resource supply treatments (N, P, K) in a tropical forest in Panama for 29 months. We applied either antibiotics or control water to replicated seedlings of five common tree species ( Alseis blackiana , Desmopsis panamensis , Heisteria concinna , Sorocea affinis , and Tetragastris panamensis ). These antibiotic treatments significantly reduced both the abundance and diversity of bacteria epiphytically as well as endophytically. Overall, the effect of antibiotics on performance was highly host specific. Applying antibiotics increased growth for three species by as much as 49% ( Alseis , Heisteria , and Tetragastris ), decreased growth for a fourth species by nearly 20% ( Sorocea ), and had no impact on a fifth species ( Desmopsis ). Perhaps more importantly, the degree to which foliar bacteria were harmful or not varied with soil resource supply. Specifically, applying antibiotics had no effect when potassium was added but increased growth rate by almost 40% in the absence of potassium. Alternatively, phosphorus enrichment caused the effect of bacteria to switch from being primarily beneficial to harmful or vice versa, but this depended entirely on the presence or absence of nitrogen enrichment ( i.e ., important and significant interactions). Our results are the first to demonstrate that the net effect of reducing the abundance and diversity of bacteria can have very strong positive and negative effects on seedling performance. Moreover, these effects were clearly mediated by soil resource availability. Though speculative, we suggest that foliar bacteria may interact with soil fertility to comprise an important, yet cryptic dimension of niche differentiation, which can have important implications for species coexistence. This article is protected by copyright. All rights reserved.

Description: Emerging aquatic insects from streams are important food sources for riparian predators, yet their availability is seasonally limited. Spatial heterogeneity in stream water temperature was found to spatially desynchronize the emergence timing of aquatic insects, and prolong their flight period, potentially enhancing consumer growth. While a mayfly Ephemerella maculata emergence lasted for 12-22 days in local sites along a river, mayflies emerged 19 days earlier from warmer than cooler sites. Therefore, the overall emergence of E. maculata from the river lasted for 37 days, and adult swarms were observed over that same period in an adjacent reproductive habitat. A feeding experiment with the riparian spider Tetragnatha versicolor showed that a prolonged subsidy, as would occur in a heterogeneous river, led to higher juvenile growth than a synchronous pulsed subsidy of equal total biomass, as would typify a more homogeneous river. Since larger female adult spiders produce more eggs, spiders that received prolonged subsidy as juveniles should achieve higher fecundity. Restoring spatial heterogeneity in streams may benefit not only stream communities but also riparian predators. This article is protected by copyright. All rights reserved.

Description: Mixing models are statistical tools that use biotracers to probabilistically estimate the contribution of multiple sources to a mixture. These biotracers may include contaminants, fatty acids, or stable isotopes, the latter of which are widely used in trophic ecology to estimate the mixed diet of consumers. Bayesian implementations of mixing models using stable isotopes (e.g. MixSIR, SIAR) are regularly used by ecologists for this purpose, but basic questions remain about when each is most appropriate. In this study, we describe the structural differences between common mixing model error formulations in terms of their assumptions about the predation process. We then introduce a new parameterization that unifies these mixing model error structures, as well as implicitly estimates the rate at which consumers sample from source populations (i.e. consumption rate). Using simulations and previously published mixing model datasets, we demonstrate that the new error parameterization outperforms existing models and provides an estimate of consumption. Our results suggest that the error structure introduced here will improve future mixing model estimates of animal diet. This article is protected by copyright. All rights reserved.

Description: Contagious seed dispersal refers to the tendency for some sites to receive many dispersed seeds while other sites receive few dispersed seeds. Contagious dispersal can lead to interspecific associations in seed arrival, and this in turn might lead to interspecific associations in seedling recruitment. We evaluate the extent of spatially contagious seed arrival, the frequency of positive interspecific associations in seed arrival, and their consequences for seedling recruitment at the community level in a tropical moist forest. We quantified seed arrival to 200 passive seed traps for 28 years of weekly censuses and seedling recruitment to 600 1-m 2 quadrats for 21 years of annual censuses on Barro Colorado Island, Panama. We assessed whether spatially contagious seed dispersal was more important among zoochorous species than among anemochorous species, increased in importance with similarity in fruiting times, and led to interspecific associations in seed arrival and seedling recruitment. We controlled adult seed source associations statistically to evaluate predicted relationships. We found that spatially contagious seed arrival was widespread among zoochorous species, but also occurred among anemochorous species when the strong, consistent trade winds were present. Significant interspecific associations in seed arrival were more likely for pairs of species with zoochorous seeds and similar fruiting times and persisted through seedling recruitment. Thus, interspecifically contagious seed dispersal affects local species composition and alters the mixture of interspecific interactions through the seed, germination and early seedling stages in this forest. Future investigations should consider the implications of interspecific association at the regeneration stages documented here for later life stages and species coexistence. This article is protected by copyright. All rights reserved.

Description: Understanding processes that may stabilize ecological systems confronted with rapidly changing environmental conditions is a key issue in ecology. We studied a system of highly fluctuating populations, the moth Achyra rantalis feeding on the plant Sesuvium portulacastrum in a group of small subtropical islands of the Bahamas. The plant is a prostrate inhabitant of shorelines, and consequently moths are highly vulnerable to being consumed by the ground-foraging lizard Anolis sagrei . We measured the percent ground cover of Sesuvium and abundance of Achyra on 11 islands with lizards present and 21 islands without lizards annually for 10 consecutive years. Overall abundance of Achyra was 4.6 times higher on no-lizard islands than on lizard islands. The percent cover of Sesuvium exhibited lower temporal variability on lizard islands when the study site was undisturbed by hurricanes, and higher recovery rate on lizard islands following hurricanes. We suggest that both of these stabilizing phenomena are linked to a trophic cascade in which predatory lizards control herbivore populations, thereby suppressing outbreaks and enhancing plant recovery following physical disturbance. This article is protected by copyright. All rights reserved.

Description: Seed dispersal effectiveness, which measures the number of adult plant individuals produced by seed dispersal, is the product of the number of seeds dispersed and the probability a seed produces an adult. Directed dispersal to certain habitat types may enhance some stages of recruitment but disfavor others, generating demographic conflicts in plant ontogeny. We asked whether temporal changes in habitat features may affect the distribution of seedlings recruited from dispersed acorns, and whether this could induce shifts in the life-stage conflicts experienced by successive cohorts of naturally recruited plants. As early successional habitats are characterized by rapid change, we used a burnt pine stand in southern Spain to monitor the recruitment and performance of a major tree species ( Quercus ilex ) across seven years in four types of post-fire habitats. These differed in structure and included patches of unburnt forest and three management alternatives of burnt trees: logging, partial cutting, and non-intervention. Young oaks that resprouted after the fire were mainly located near acorn sources, while new seedlings initially emerged mostly in habitats with standing snags due to habitat selection by European jays, Garrulus glandarius, for dispersal. The dead pines gradually collapsed and attracted less dispersal, so subsequent seedling cohorts mainly recruited within patches of unburnt pines. These live pines enhanced the survival of the oaks located beneath their canopy but greatly reduced their growth as compared to the other post-fire habitats, thus representing a demographic conflict that was absent elsewhere. As a consequence of the directional shift in the habitat where seedlings recruited, successive seedling cohorts experienced a gradual improvement in their likelihood of survival but a reduction in growth. The progressive intensification of this life-stage conflict hinged on the reduction of vertical structures in the habitat with standing burnt pines. Recruitment success thus involved temporal variation in the habitat where recruitment occurred, likely resulting from changes in the direction of seed dispersal, and spatial variation in habitat suitability for seedling establishment and growth. Temporal changes in habitat structure can indirectly change the environment in which recruitment occurs, and consequently seed dispersal effectiveness, by shifting the direction of seed dispersal. This article is protected by copyright. All rights reserved.

Description: Long-term experiments provide a way to test presumed causes of successional or environmentally driven vegetation changes. Early-successional nitrogen (N)-fixing plants are widely thought to facilitate productivity and vegetation development on N-poor sites, thus accounting for observed vegetation patterns later in succession. We tested this facilitative impact on vegetation development in a 23-year field experiment on an Interior Alaska (U.S.A.) floodplain. On three replicate early-successional silt bars, we planted late-successional white spruce ( Picea glauca ) seedlings in the presence and absence of planted seedlings of an early-successional N-fixing shrub—thinleaf alder ( Alnus incana ). Alder initially facilitated survivorship and growth of white spruce. Within six years, however, after canopy closure, alder negatively affected spruce survivorship and growth. Our three replicate sites followed different successional trajectories. One site was eliminated by erosion and supported no vegetation development during our study. The other two sites, which differed in site moisture, diverged in vegetation composition. Structural equation modeling (SEM) suggested that, in the drier of these two sites, alder inhibited spruce growth directly (presumably by competition) and indirectly through effects mediated by competition with other woody species. However at the wetter site alder had both positive and negative effects on spruce growth, with negative effects predominating. Snowshoe hares ( Lepus americanus ) in alder thickets further reduced height growth of spruce in the wetter site. We conclude that net effects of alder on white spruce, the late-successional dominant, were primarily inhibitory and indirect, with the mechanisms depending on initial site moisture. Our results highlight the importance of long-term research showing that small differences among initial replicate sites can cause divergence in successional trajectories–consistent with individualistic distributions of species and communities along environmental gradients. This divergence was detectable only decades later. This article is protected by copyright. All rights reserved.

Description: Decades of ecological study have demonstrated the importance of top-down and bottom-up controls on food webs, yet few studies within this context have quantified the magnitude of energy and material fluxes at the whole-ecosystem scale. We examined top-down and bottom-up effects on food web fluxes using a field experiment that manipulated the presence of a consumer, the Trinidadian guppy Poecilia reticulata , and the production of basal resources by thinning the riparian forest canopy to increase incident light. To gauge the effects of these reach-scale manipulations on food web fluxes, we used a nitrogen ( 15 N) stable isotope tracer to compare basal resource treatments (thinned canopy vs. control) and consumer treatments (guppy introduction vs. control). The thinned canopy stream had higher primary production than the natural canopy control, leading to increased N fluxes to invertebrates that feed on benthic biofilms (grazers), fine benthic organic matter (collector-gatherers), and organic particles suspended in the water column (filter-feeders). Stream reaches with guppies also had higher primary productivity and higher N fluxes to grazers and filter feeders. In contrast, N fluxes to collector-gatherers were reduced in guppy introduction reaches relative to upstream controls. N fluxes to leaf shredding invertebrates, predatory invertebrates and the other fish species present (Hart's killifish, Anablepsoides hartii ) did not differ across light or guppy treatments, suggesting that effects on detritus-based linkages and upper trophic levels were not as strong. Effect sizes of guppy and canopy treatments on N flux rates were similar for most taxa, though guppy effects were the strongest for filter feeding invertebrates while canopy effects were the strongest for collector-gatherer invertebrates. Combined, these results extend previous knowledge about top-down and bottom-up controls on ecosystems by providing experimental, reach-scale evidence that both pathways can act simultaneously and have equally strong influence on nutrient fluxes from inorganic pools through primary consumers. This article is protected by copyright. All rights reserved.

Description: Fifty years ago, Ehrlich and Raven proposed that insect herbivores have driven much of plant speciation, particularly at tropical latitudes. There have been no explicit tests of their hypotheses. Indeed there were no proposed mechanisms either at the time or since by which herbivores might generate new plant species. Here we outline two main classes of mechanisms, pre-zygotic and post-zygotic, with a number of scenarios in each by which herbivore-driven changes in host plant secondary chemistry might lead to new plant lineage production. The former apply mainly to a sympatric model of speciation while the latter apply to a parapatric or allopatric model. Our review suggests that the steps of each mechanism are known to occur individually in many different systems, but no scenario has been thoroughly investigated in any one system. Nevertheless, studies of Dalechampia and its herbivores and pollinators, and patterns of defense tradeoffs in trees on different soil types in the Peruvian Amazon provide evidence consistent with the original hypotheses of Ehrlich and Raven. For herbivores to drive sympatric speciation, our findings suggest that interactions with both their herbivores and their pollinators should be considered. In contrast, herbivores may drive speciation allopatrically without any influence by pollinators. Finally, there is evidence that these mechanisms are more likely to occur at low latitudes, and thus more likely to produce new species in the tropics. The mechanisms we outline provide a predictive framework for further study of the general role that herbivores play in diversification of their host plants. This article is protected by copyright. All rights reserved.

Description: Community ecologists have strived to find mechanisms that mediate the assembly of natural communities. Recent evidence suggests that natural enemies could play an important role in the assembly of hyper-diverse tropical plant systems. Classic ecological theory predicts that in order for coexistence to occur, species differences must be maximized across biologically important niche dimensions. For plant-herbivore interactions, it has been recently suggested that, within a particular community, plant species that maximize the difference in chemical defense profiles compared to neighboring taxa will have a relative competitive advantage. Here we tested the hypothesis that plant chemical diversity can affect local community composition in the hyper-diverse genus Piper at a lowland wet forest location in Costa Rica. We first characterized the chemical composition of 27 of the most locally abundant species of Piper . We then tested whether species with different chemical compositions were more likely to coexist. Finally, we assessed the degree to which Piper phylogenetic relationships are related to differences in secondary chemical composition and community assembly. We found that, on average, co-occurring species were more likely to differ in chemical composition than expected by chance. Contrary to expectations, there was no phylogenetic signal for overall secondary chemical composition. In addition we found that species in local communities were, on average, more phylogenetically closely related than expected by chance, suggesting that functional traits other than those measured here also influence local assembly. We propose that selection by herbivores for divergent chemistries between closely related species facilitates the coexistence of a high diversity of congeneric taxa via apparent competition. This article is protected by copyright. All rights reserved.

Description: Invasive alien predators (IAP) are spreading on a global scale and often with devastating ecological effects. One reason for their success may be that prey species fail to recognize them due to a lack of co-evolutionary history. We performed a comprehensive test of this ‘prey naiveté’ hypothesis using a novel approach: we tested whether predator-naive tadpoles of the agile frog ( Rana dalmatina ) display antipredator behaviour upon encountering chemical cues produced by native, invasive (established or recent) or allopatric fishes (four perciforms, four siluriforms and two cypriniforms). We studied the influence of population origin on predator-detection ability by presenting chemical cues to predator-naive tadpoles that originated from fishless hill-ponds or fish-infested floodplain populations. Before trials, we fed fishes with tadpoles or an alternative food to test whether direct chemical cues from the predator's diet influences the tadpoles’ recognition of potential predators. Tadpoles reduced their activity upon exposure to cues from native and long-established invasive perciforms, but not in response to recent invaders, allopatric predators or to any siluriforms. Also, predators that were previously fed with tadpoles did not universally induce behavioural defences upon first encounter. Finally, tadpoles originating from isolated hill-ponds exhibited higher baseline activity and responded weaker than their conspecifics from floodplain populations, which co-exist with predatory fishes. Our results indicate that tadpoles may be vulnerable to invading predatory fishes due to their inability to recognize them as dangerous, though their ability to recognize invasive IAP may evolve rapidly, in less than 30 generations. This article is protected by copyright. All rights reserved.

Description: Chronic wasting disease (CWD) is a fatal neurodegenerative disease affecting free-ranging and captive cervids that now occurs in 24 US States and two Canadian provinces. Despite the potential threat of CWD to deer populations little is known about the rates of infection and mortality caused by this disease. We used epidemiological models to estimate the force-of-infection and disease-associated mortality for white-tailed deer in the Wisconsin and Illinois CWD outbreaks. Models were based on age-prevalence data corrected for bias in aging deer using the tooth wear and replacement method. Both male and female deer in the Illinois outbreak had higher corrected age-specific prevalence with slightly higher female infection than deer in the Wisconsin outbreak. Corrected ages produced more complex models with different infection and mortality parameters than those based on apparent prevalence. We found that adult male deer have 〉 3 fold higher risk of CWD infection than female deer. Males also had higher disease mortality than female deer. As a result CWD prevalence was 2 fold higher in adult males than females. We also evaluated the potential impacts of alternative contact structures on transmission dynamics in Wisconsin deer. Results suggested that transmission of CWD among male deer during the nonbreeding season may be a potential mechanism for producing higher rates of infection and prevalence characteristically found in males. However, alternatives based on high environmental transmission and transmission from females to males during the breeding season may also play a role. This article is protected by copyright. All rights reserved.

Description: Extinctions have no simple determinism, but rather result from complex interplays between environmental factors and demographic-genetic feedback that occur at small population size. Inbreeding depression has been assumed to be a major trigger of extinction vortices, yet very few models have studied its consequences in dynamic populations with realistic population structure. Here we investigate the impact of Complementary Sex Determination (CSD) on extinction in parasitoid wasps and other insects of the order Hymenoptera. CSD is believed to induce enough inbreeding depression to doom simple small populations to extinction, but we suggest that in parasitoids CSD may have the opposite effect. Using a theoretical model combining the genetics of CSD and the population dynamics of host-parasitoid systems, we show that CSD can reduce the risk of parasitoid extinction by reducing fluctuations in population size. Our result suggests that inbreeding depression is not always a threat to population survival, and that considering trophic interactions may reverse some pervasive hypotheses on its demographic impact. This article is protected by copyright. All rights reserved.

Description: Stability in population dynamics is an emergent property of the interaction between direct and delayed density dependence, the strengths of which vary with environmental covariates. Analysis of variation across populations in the strength of direct and delayed density dependence can reveal variation in stability properties of populations at the species level. We examined the stability properties of 22 elk/red deer populations in a two-stage analysis. First, we estimated direct and delayed density dependence applying an AR(2) model in a Bayesian hierarchical framework. Second, we plotted the coefficients of direct and delayed density dependence in the Royama parameter plane. We then used a hierarchical approach to test the significance of environmental covariates of direct and delayed density dependence. Three populations exhibited highly stable and convergent dynamics with strong direct, and weak delayed, density dependence. The remaining 19 populations exhibited more complex dynamics characterized by multi-annual fluctuations. Most (15 of 19) of these exhibited a combination of weak to moderate direct and delayed density dependence. Best-fit models included environmental covariates in 17 populations (77% of the total). Of these, interannual variation in growing-season primary productivity and interannual variation in winter temperature were the most common, performing as the best-fit covariate in six and five populations, respectively. Interannual variation in growing-season primary productivity was associated with the weakest combination of direct and delayed density dependence, while interannual variation in winter temperature was associated with the strongest combination of direct and delayed density dependence. These results accord with a classic theoretical prediction that environmental variability should weaken population stability. They furthermore suggest that two forms of environmental variability, one related to forage resources and the other related to abiotic conditions, both reduce stability but in opposing fashion: one through weakened direct density dependence and the other through strengthened delayed density dependence. Importantly, however, no single abiotic or biotic environmental factor emerged as generally predictive of the strengths of direct or delayed density dependence, nor of the stability properties emerging from their interaction. Our results emphasize the challenges inherent to ascribing primacy to drivers of such parameters at the species level and distribution scale. This article is protected by copyright. All rights reserved.

Description: Ecosystem multifunctionality – the simultaneous production of multiple ecosystem functions – depends on community diversity, composition, productivity and spatial scale. In metacommunities, each of these community properties is affected by how species disperse between local patches to track environmental change. Here we use a consumer-resource metacommunity model of resource competition to show how dispersal affects the link between diversity, composition, and ecosystem multifunctionality. When species differ in their functional traits, and environmental niche, metacommunity multifunctionality becomes highly dependent upon dispersal, which allows community diversity to be maintained when environmental conditions change. Dispersal promotes multifunctionality in two ways: 1) species sorting – whereby species track local environmental changes by shifting in space, thus preserving diversity and ensuring high biomass productivity, and 2) mass effects – whereby source sink dynamics allow species to persist in sub-optimal environments, thus increasing local diversity. Changing the rate at which species disperse affects the strength of these metacommunity processes, and so metacommunity multifunctionality exhibits a unimodal relationship with dispersal, peaking when dispersal is intermediate. Species sorting dynamics also provide spatial insurance whereby compensatory dynamics stabilize the fluctuations of each function through time at the regional scale. However, this does not extend to the local scale, where species sorting results in high temporal variability for each function, even though the overall rates of multifunctionality are high. Our results suggest that metacommunity processes are important determinants of ecosystem multifunctionality, and thus effective management of multiple ecosystem functions requires consideration of landscape connectivity. This article is protected by copyright. All rights reserved.

Description: The cost and difficulty of manipulative field studies makes low statistical power a pervasive issue throughout most sub-disciplines in ecology. Ecologists are already aware that small sample sizes increase the probability of committing Type II errors. In this article, we address a relatively unknown problem with low power: underpowered studies must overestimate small effect sizes in order to achieve statistical significance. First, we describe how low replication coupled with weak effect sizes leads to Type M errors, or exaggerated effect sizes. We then conduct a meta-analysis to determine the average statistical power and Type M error rate for manipulative field experiments that address important questions related to global change; global warming, biodiversity loss, and drought. Finally, we provide recommendations for avoiding Type M errors and constraining estimates of effect size from underpowered studies. This article is protected by copyright. All rights reserved.

Description: Although the functional basis of variable and synchronous seed production (masting behavior) has been extensively investigated, only recently has attention been focused on the proximate mechanisms driving this phenomenon. We analyzed the relationship between weather and acorn production in 15 species of oaks (genus Quercus ) from three geographic regions on two continents with the goals of determining the extent to which similar sets of weather factors affect masting behavior across species and to explore the ecological basis for the similarities detected. Lag-1 temporal autocorrelations were predominantly negative, supporting the hypothesis that stored resources play a role in masting behavior across this genus, and we were able to determine environmental variables correlating with acorn production in all but one of the species. Standard weather variables outperformed “differential-cue” variables based on the difference between successive years in a majority of species, consistent with the hypothesis that weather is linked directly to the proximate mechanism driving seed production and that masting in these species is likely to be sensitive to climate change. Based on the correlations between weather variables and acorn production, cluster analysis failed to generate any obvious groups of species corresponding to phylogeny or life-history. Discriminant function analyses were, however, able to identify the phylogenetic section to which the species belonged and, controlling for phylogeny, the length of time species required to mature acorns, whether they were evergreen or deciduous, and, to a lesser extent, the geographic region to which they are endemic. These results indicate that similar proximate mechanisms are driving acorn production in these species of oaks, that the environmental factors driving seed production in oaks are to some extent phylogenetically conserved, and that the shared mechanisms driving acorn production result in some degree of synchrony among coexisting species in a way that potentially enhances predator satiation, at least when they have acorns requiring the same length of time to mature. This article is protected by copyright. All rights reserved.

Description: Many studies suggest that biodiversity may be particularly important for ecosystem multi functionality, because different species with different traits can contribute to different functions. Support, however, comes mostly from experimental studies conducted at small spatial scales in low-diversity systems. Here, we test whether different species contribute to different ecosystem functions that are important for carbon cycling in a high-diversity human-modified tropical forest landscape in Southern Mexico. We quantified aboveground standing biomass, primary productivity, litter production, and wood decomposition at the landscape level, and evaluated the extent to which tree species contribute to these ecosystem functions. We used simulations to tease apart the effects of species richness, species dominance and species functional traits on ecosystem functions. We found that dominance was more important than species traits in determining a species’ contribution to ecosystem functions. As a consequence of the high dominance in human-modified landscapes, the same small subset of species mattered across different functions. In human-modified landscapes in the tropics, biodiversity may play a limited role for ecosystem multifunctionality due to the potentially large effect of species dominance on biogeochemical functions. However, given the spatial and temporal turnover in species dominance, biodiversity may be critically important for the maintenance and resilience of ecosystem functions. This article is protected by copyright. All rights reserved.

Description: A number of ecological factors have been shown to influence the importance of positive interactions (i.e. facilitation) in nature, including environmental stress and ontogenetic effects, and many more are likely to emerge as facilitation research expands to new ecosystems and taxa. In this study, I used a combination of field surveys and experiments to explore the roles of stress, ontogeny, and organismal movement in determining the importance of mussel ( Mytilus californianus) recruit facilitation in central California. Results indicate that interactions between mussel recruits (shell length 〈 20 mm) and habitat ameliorating neighbors shift from neutral to positive from the low to high mussel zone. I also observed ontogenetic shifts in recruit survival and growth in the upper mussel zone that suggest mussel recruits migrate from algal substrate to adult mussel beds. This type of habitat shift, where an organism moves sequentially from one facilitator to another, may be common in nature and presents an exciting new area for research. This article is protected by copyright. All rights reserved.

Description: Tropical savannas are hypothesized to be hot spots of nitrogen fixer diversity and activity because of the high disturbance and low nitrogen characteristic of savanna landscapes. Here we compare the abundances of nitrogen-fixing and non-fixing trees in both tropical savannas and tropical forests under climatically equivalent conditions, using plant inventory studies across 566 plots in South America and Africa. A single factor – aridity – explained 19-54% of the variance in fixer abundance, and unexpectedly was more important than fire frequency, biome, and continent. Nitrogen fixers were more abundant in arid environments; as a result, African savannas, which tend to be drier, were richer in nitrogen fixers than South American savannas. Fixer abundance converged on similar levels in forests in both continents. We conclude that climate plays a greater role than fire in determining the distribution of nitrogen fixers across tropical savanna and forest biomes. This article is protected by copyright. All rights reserved.

Description: The timing of the fruit-set stage (i.e. start and end of fruit set) is crucial in a plant's life cycle, but its response to temperature change is still unclear. We investigated the timing of seven phenological events, including fruit-set dates during 3 years for six alpine plants transplanted to warmer (+ ~3.5 °C in soils) and cooler (- ~3.5 °C in soils) locations along an altitudinal gradient in the Tibetan area. We found that fruit-set dates remained relatively stable under both warming and cooling during the 3-years transplant experiment. Three earlier phenological events (emergence of first leaf, first bud set, and first flowering) and two later phenological events (first leaf coloring and complete leaf coloring) were earlier by 4.8-8.2 days °C −1 and later by 3.2-7.1 days °C −1 in response to warming. Conversely, cooling delayed the three earlier events by 3.8-6.9 days °C −1 and advanced the two later events by 3.2-8.1 days °C −1 for all plant species. The timing of the first and/or last fruit-set dates, however, did not change significantly compared to earlier and later phenological events. Statistical analyses also showed that the dates of fruit set were not significantly correlated or had lower correlations with changes of soil temperature relative to the earlier and later phenological events. Alpine plants may thus acclimate to changes in temperature for their fruiting function by maintaining relatively stable timings of fruit set compared with other phenological events to maximize the success of seed maturation and dispersal in response to short-term warming or cooling. This article is protected by copyright. All rights reserved.

Description: Statistical models of habitat preference and species distribution (e.g., Resource Selection Functions and Maximum Entropy approaches) perform a quantitative comparison of the use of space with the availability of all habitats in an animal's environment. However, not all of space is accessible all of the time to all individuals, so availability is in fact determined by limitations in animal perception and mobility. Therefore, measuring habitat availability at biologically relevant scales is essential for understanding preference, but herein lies a trade-off: Models fitted at large spatial scales, will tend to average across the responses of different individuals that happen to be in regions with contrasting habitat compositions. We suggest that such models may fail to capture local extremes (hotspots and coldspots) in animal usage and call this potential problem, homogenization . In contrast, models fitted at smaller scales will vary stochastically depending on the particular habitat composition of their narrow spatial neighborhood, and hence fail to describe responses when predicting for different sampling instances. This is the now well-documented issue of non-transferability of habitat models. We illustrate this tradeoff, using a range of simulated experiments, incorporating variations in environmental gradients, richness and fragmentation. We propose diagnostics for detecting the two issues of homogenization and non-transferability and show that these scale-related symptoms are likely to be more pronounced in highly fragmented or steeply graded landscapes. Further, we address these problems by treating the neighborhood of each cell in the landscape grid as an individual sampling instance (with its own neighborhood), hence allowing coefficients to respond to the local expectations of environmental variables according to a Generalized Functional Response (GFR). Under simulation this approach is consistently better at estimating robust (i.e., transferable) habitat models at smaller scales, and less susceptible to homogenization at larger scales. At the same time, it represents the first application of a GFR to continuous space (rather than multiple, spatially distinct datasets), allowing the predictive advantages of this extension of species distribution models to become available to data from large-scale but single-site field studies.

Description: Conspecific negative density dependence (CNDD) has been recognized as a key mechanism underlying species coexistence, especially in tropical forests. Recently, some studies have reported that seedling survival is also negatively correlated with the phylogenetic relatedness between neighbors and focal individuals, termed phylogenetic negative density dependence (PNDD). In contrast to CNDD or PNDD, shared habitat requirements between closely related individuals are thought to be a cause of observed positive effects of closely related neighbors, which may affect the strength and detectability of CNDD or PNDD. In order to investigate the relative importance of these mechanisms for tropical tree seedling survival, we used generalized linear mixed models to analyze how the survival of more than 10 000 seedlings of woody plant species related to neighborhood and habitat variables in a tropical rainforest in southwest China. By comparing models with and without habitat variables, we tested how habitat filtering affected the detection of CNDD and PNDD. The best-fitting model suggested that CNDD and habitat filtering played key roles in seedling survival, but that, contrary to our expectations, phylogenetic positive density dependence (PPDD) had a distinct and important effect. While habitat filtering affected the detection of CNDD by decreasing its apparent strength, it did not explain the positive effects of closely related neighbors. Our results demonstrate that a failure to control for habitat variables and phylogenetic relationships may obscure the importance of conspecific and heterospecific neighbor densities for seedling survival.

Description: Time can be a limiting constraint for consumers, particularly when resource phenology mediates foraging opportunity. Though a large body of research has explored how resource phenology influences trophic interactions, this work has focused on the topics of trophic mismatch or predator swamping, which typically occur over short periods, at small spatial extents or coarse resolutions. In contrast many consumers integrate across landscape heterogeneity in resource phenology, moving to track ephemeral food sources that propagate across space as resource waves. Here we provide a conceptual framework to advance the study of phenological diversity and resource waves. We define resource waves, review evidence of their importance in recent case studies, and demonstrate their broader ecological significance with a simulation model. We found that consumers ranging from fig wasps ( Chalcidoidea) to grizzly bears ( Ursus arctos ) exploit resource waves, integrating across phenological diversity to make resource aggregates available for much longer than their component parts. In model simulations, phenological diversity was often more important to consumer energy gain than resource abundance per se. Current ecosystem-based management assumes that species abundance mediates the strength of trophic interactions. Our results challenge this assumption and highlight new opportunities for conservation and management. Resource waves are an emergent property of consumer–resource interactions and are broadly significant in ecology and conservation.

Description: Essential fatty acids (EFAs) are primarily generated by phytoplankton in aquatic ecosystems, and can limit the growth, development, and reproduction of higher consumers. Among the most critical of the EFAs are highly unsaturated fatty acids (HUFAs), which are only produced by certain groups of phytoplankton. Changing environmental conditions can alter phytoplankton community and fatty acid composition and affect the HUFA content of higher trophic levels. Almost no research has addressed intraspecific variation in HUFAs in zooplankton, nor intraspecific relationships of HUFAs with body size and fecundity. This is despite that intraspecific variation in HUFAs can exceed interspecific variation and that intraspecific trait variation in body size and fecundity is increasingly recognized to have an important role in food web ecology (effect traits). Our study addressed the relative influences of abiotic selection and food web effects associated with climate change on intraspecific differences and interrelationships between HUFA content, body size, and fecundity of freshwater copepods. We applied structural equation modeling and regression analyses to intraspecific variation in a dominant calanoid copepod, Leptodiatomus minutus , among a series of shallow north-temperate ponds. Climate-driven diurnal temperature fluctuations favored the coexistence of diversity of phytoplankton groups with different temperature optima and nutritive quality. This resulted in unexpected positive relationships between temperature, copepod DHA content and body size. Temperature correlated positively with diatom biovolume, and mediated relationships between copepod HUFA content and body size, and between copepod body size and fecundity. The presence of brook trout further accentuated these positive effects in warm ponds, likely through nutrient cycling and stimulation of phytoplankton resources. Climate change may have previously unrecognized positive effects on freshwater copepod DHA content, body size, and fecundity in the small, shallow bodies of inland waters that are commonly found in north-temperate landscapes.

Description: Understanding the impacts of biodiversity loss on ecosystem functioning and services has been a central issue in ecology. Experiments in synthetic communities suggest that biodiversity loss may erode a set of ecosystem functions, but studies in natural communities indicate that the effects of biodiversity loss are usually weak and that multiple functions can be sustained by relatively few species. Yet, the mechanisms by which natural ecosystems are able to maintain multiple functions in the face of diversity loss remain poorly understood. With a long-term and large-scale removal experiment in the Inner Mongolian grassland, here we showed that losses of plant functional groups (PFGs) can reduce multiple ecosystem functions, including biomass production, soil NO 3 -N use, net ecosystem carbon exchange, gross ecosystem productivity, and ecosystem respiration, but the magnitudes of these effects depended largely on which PFGs were removed. Removing the two dominant PFGs (perennial rhizomatous grasses and perennial bunchgrasses) simultaneously resulted in dramatic declines in all examined functions, but such declines were circumvented when either dominant PFG was present. We identify the major mechanism for this as a compensation effect by which each dominant PFG can mitigate the losses of others. This study provides evidence that compensation ensuing from PFG losses can mitigate their negative consequence, and thus natural communities may be more resilient to biodiversity loss than currently thought if the remaining PFGs have strong compensation capabilities. On the other hand, ecosystems without well-developed compensatory functional diversity may be much more vulnerable to biodiversity loss. This article is protected by copyright. All rights reserved.

Description: Understanding processes that promote species coexistence is integral to diversity maintenance. In hyperdiverse tropical forests, local conspecific density (LCD) and light are influential to woody seedling recruitment and soil nutrients are often limiting, yet the simultaneous effects of these factors on seedling survival across time remain unknown. We fit species- and age-specific models to census and resource data of seedlings of 68 woody species from a Costa Rican wet tropical forest. In decreasing order of prevalence, seedling survivorship was related to LCD, soil base cations, irradiance, nitrogen, and phosphorus. Species–specific responses to factors did not covary, providing evidence that species life history strategies have not converged to one continuum of high-surviving stress tolerant to low-surviving stress intolerant species. Survival responses to all factors varied over the average seedling's lifetime, indicating seedling requirements change with age and conclusions drawn about processes important to species coexistence depend on temporal resolution. This article is protected by copyright. All rights reserved.

Description: Sea-to-land nutrient transfers can connect marine food webs to those on land, creating a dependence on marine webs by opportunistic species. We show how nitrogen, imported by grey seals, Halichoerus grypus , and traced through stable isotope (δ 15 N) measurements in marram grass , Ammophila breviligulata , significantly alters foraging behavior of a free-roaming megaherbivore (feral horses, Equus ferus caballus ) on Sable Island, Canada. Values of δ 15 N correlated with protein content of marram and strongly related to pupping-seal densities, and positively influenced selective foraging by horses. The latter was density-dependent consistent with optimal foraging theory. We present the first demonstration of how sea-to-land nutrient transfers can affect the behavioral process of resource selection (resource use relative to availability) of terrestrial consumers. We hypothesize that persistence of horses on Sable Island is being facilitated by N subsidies. Our results have relevance to advancing theory on trophic dynamics in island biogeography and metaecosystem ecology. This article is protected by copyright. All rights reserved.

Description: The “landscape of fear” model, recently advanced in research on the non-lethal effects of carnivores on ungulates, predicts that prey will exhibit detectable antipredator behavior not only during risky times (i.e., predators in close proximity) but also in risky places (i.e., habitat where predators kill prey or tend to occur). Aggregation is an important antipredator response in numerous ungulate species, making it a useful metric to evaluate the strength and scope of the landscape of fear in a multi-carnivore, multi-ungulate system. We conducted ungulate surveys over a 2-year period in South Africa to test the influence of three broad-scale sources of variation in the landscape on spatial patterns in aggregation: (1) habitat structure, (2) where carnivores tended to occur (i.e., population-level utilization distributions), and (3) where carnivores tended to kill ungulate prey (i.e., probabilistic kill site maps). We analyzed spatial variation in aggregation for six ungulate species exposed to predation from recently reintroduced lion ( Panthera leo ) and spotted hyena ( Crocuta crocuta ). Although we did detect larger aggregations of ungulates in “risky places,” these effects existed primarily for smaller-bodied (〈150 kg) ungulates and were relatively moderate (change of ≤4 individuals across all habitats). In comparison, ungulate aggregations tended to increase at a slightly lower rate in habitat that was more open. The lion, an ambush (stalking) carnivore, had stronger influence on ungulate aggregation than the hyena, an active (coursing) carnivore. In addition, places where lions tended to kill prey had a greater effect on ungulate aggregation than places where lions tended to occur, but an opposing pattern existed for hyena. Our study reveals heterogeneity in the landscape of fear and suggests broad-scale risk effects following carnivore reintroduction only moderately influence ungulate aggregation size and vary considerably by predator hunting mode, type of predation risk, and prey species.

Description: Many insect parasitoids are highly specialized and thus develop on only one or a few related host species, yet some hosts are attacked by many different parasitoid species in nature. For this reason, they have been often used to examine the consequences of competitive interactions. Hosts represent limited resources for larval parasitoid development and thus one competitor usually excludes all others. Although parasitoid competition has been debated and studied over the past several decades, understanding the factors that allow for coexistence among species sharing the same host in the field remains elusive. Parasitoids may be able to coexist on the same host species if they partition host resources according to size, age, or stage, or if their dynamics vary at spatial and temporal scales. One area that has thus far received little experimental attention is if competition can alter host usage strategies in parasitoids that in the absence of competitors attack hosts of the same size in the field. Here, we test this hypothesis with two parasitoid species in the genus Aphytis , both of which are specialized on the citrus pest California red scale Aonidiella aurantii . These parasitoids prefer large scales as hosts and yet coexist in sympatry in eastern parts of Spain. Parasitoids and hosts were sampled in 12 replicated orange groves. When host exploitation by the stronger competitor, A. melinus , was high the poorer competitor, A. chrysomphali , changed its foraging strategy to prefer alternative plant substrates where it parasitized hosts of smaller size. Consequently, the inferior parasitoid species shifted both its habitat and host size as a result of competition. Our results suggest that density-dependent size-mediated asymmetric competition is the likely mechanism allowing for the coexistence of these two species, and that the use of suboptimal (small) hosts can be advantageous under conditions imposed by competition where survival in higher quality larger hosts may be greatly reduced.

Description: Long-distance breeding and natal dispersal play central roles in many ecological and evolutionary processes, including gene flow, population dynamics, range expansion, and individual responses to fluctuating biotic and abiotic conditions. However, the relative contribution of long-distance dispersal to these processes depends on the ability of dispersing individuals to successfully reproduce in their new environment. Unfortunately, due to the difficulties associated with tracking dispersal in the field, relatively little is known about its reproductive consequences. Furthermore, because reproductive success is influenced by a variety of processes, disentangling the influence of each of these processes is critical to understanding the direct consequences of dispersal. In this study, we used stable hydrogen and carbon isotopes to estimate long-distance dispersal and winter territory quality in a migratory bird, the American Redstart ( Setophaga ruticilla ). We then applied Aster life-history models to quantify the strength of influence of these factors on apparent reproductive success. We found no evidence that male or female reproductive success was lower for long-distance dispersers relative to non-dispersing individuals. In contrast, carry-over effects from the winter season did influence male, but not female, reproductive success. Use of Aster models further revealed that for adult males, winter territory quality influenced the number of offspring produced whereas for yearling males, high-quality winter territories were associated with higher mating and nesting success. These results suggest that although long-distance natal and breeding dispersal carry no immediate reproductive cost for American Redstarts, reproductive success in this species may ultimately be limited by the quality of winter habitat.

Description: Ecological patterns arise from the interplay of many different processes, and yet the emergence of consistent phenomena across a diverse range of ecological systems suggests that many patterns may in part be determined by statistical or numerical constraints. Differentiating the extent to which patterns in a given system are determined statistically, and where it requires explicit ecological processes, has been difficult. We tackled this challenge by directly comparing models from a constraint-based theory, the Maximum Entropy Theory of Ecology (METE) and models from a process-based theory, the size-structured neutral theory (SSNT). Models from both theories were capable of characterizing the distribution of individuals among species and the distribution of body size among individuals across 76 forest communities. However, the SSNT models consistently yielded higher overall likelihood, as well as more realistic characterizations of the relationship between species abundance and average body size of conspecific individuals. This suggests that the details of the biological processes contain additional information for understanding community structure that are not fully captured by the METE constraints in these systems. Our approach provides a first step towards differentiating between process- and constraint-based models of ecological systems and a general methodology for comparing ecological models that make predictions for multiple patterns.

Description: In the coming century, climate change is projected to impact precipitation and temperature regimes worldwide, with especially large effects in drylands. We use big sagebrush ecosystems as a model dryland ecosystem to explore the impacts of altered climate on ecohydrology and the implications of those changes for big sagebrush plant communities using output from 10 Global Circulation Models (GCMs) for two representative concentration pathways (RCPs). We ask: 1) What is the magnitude of variability in future temperature and precipitation regimes among GCMs and RCPs for big sagebrush ecosystems and 2) How will altered climate and uncertainty in climate forecasts influence key aspects of big sagebrush water balance? We explored these questions across 1980-2010, 2030-2060, and 2070-2100 to determine how changes in water balance might develop through the 21 st century. We assessed ecohydrological variables at 898 sagebrush sites across the western US using a process-based soil water model, SOILWAT to model all components of daily water balance using site-specific vegetation parameters and site-specific soil properties for multiple soil layers. Our modeling approach allowed for changes in vegetation based on climate. Temperature increased across all GCMs and RCPs, while changes in precipitation were more variable across GCMs. Winter and spring precipitation was predicted to increase in the future (7% by 2030-2060, 12% by 2070-2100), resulting in slight increases in soil water potential (SWP) in winter. Despite wetter winter soil conditions, SWP decreased in late spring and summer due to increased evapotranspiration (6% by 2030-2060, 10% by 2070-2100) and groundwater recharge (26% and 30% increase by 2030-2060 and 2070-2100). Thus, despite increased precipitation in the cold season, soils may dry out earlier in the year, resulting in potentially longer drier summer conditions. If winter precipitation cannot offset drier summer conditions in the future, we expect big sagebrush regeneration and survival will be negatively impacted, potentially resulting in shifts in the relative abundance of big sagebrush plant functional groups. Our results also highlight the importance of assessing multiple GCMs to understand the range of climate change outcomes on ecohydrology, which was contingent on the GCM chosen. This article is protected by copyright. All rights reserved.

Description: Mass mortality of the sea urchin Diadema antillarum due to disease outbreaks in 1983 and 1991 decimated populations in the Florida Keys, and they have yet to recover. Here, we use a coupled advection-diffusion and fertilization-kinetics model to test the hypothesis that these populations are fertilization-limited. We found that fertilization success was ≥ 96% prior to the first disease outbreak, decreased substantially following recurrent disease to 3%, and has since remained low. By investigating the combined effects of physical factors (population spatial extent and current velocity) and sea urchin behavior (aggregation) on density-dependent fertilization success, we show that fertilization success at a given density increases with increasing population spatial extent and decreasing current velocity, and is greater under simulated aggregation behavior of D. antillarum . However, at present population densities, the increase in fertilization success due to aggregation is 〈 1%, even under the most favorable physical conditions. These results indicate that populations are severely fertilization-limited, and that Allee effects at low population density will continue to limit recovery. Our results can serve as a practical guide to managers in the development of coral reef restoration strategies, including the design of a D. antillarum restocking program to obtain reproductively viable populations. This article is protected by copyright. All rights reserved.

Description: Theory suggests that high activity levels in animals increase growth at the cost of increased mortality. This growth-mortality trade-off has recently been incorporated into the wider framework of the pace-of-life syndrome (POLS) hypothesis. However, activity is often quantified only in the laboratory and on a diurnal basis, leaving open the possibility that animals manage predation risk and feeding efficiency in the wild by modulating their circadian activity rhythms. Here we investigate how laboratory activity in wild brown trout parr ( Salmo trutta L.) associates with circadian activity, growth, and mortality in their natal stream. We found that individuals with high activity in the laboratory displayed high dispersal and cathemeral activity in their natal stream. In contrast, trout with low laboratory activity showed variation of activity in the wild, which was negatively related to the light intensity. Our results do not support the growth-mortality trade-off of the POLS hypothesis as highly active, fast-growing individuals showed higher survival than inactive conspecifics. These novel results show for the first time that active and inactive individuals, as scored in the lab, can show different circadian patterns of behaviour in the wild driven by light intensity. This implies that studies conducted under a narrow range of light conditions can bias our understanding of individual behavioural variation and its fitness consequences in the wild. This article is protected by copyright. All rights reserved.

Description: Encounter competition is interference competition in which animals directly contend for resources. Ecological theory predicts the trait that determines the resource holding potential (RHP), and hence the winner of encounter competition, is most often body size or mass. The difficulties of observing encounter competition in complex organisms in natural environments, however, has limited opportunities to test this theory across diverse species. We studied the outcome of encounter competition contests among mesocarnivores at deer carcasses in California to determine the most important variables for winning these contests. We found some support for current theory in that body mass is important in determining the winner of encounter competition, but we found that other factors including hunger and species-specific traits were also important. In particular, our top models were ‘strength and hunger’ and ‘size and hunger,’ with models emphasizing the complexity of variables influencing outcomes of encounter competition. In addition, our wins above predicted (WAP) statistic suggests that an important aspect that determines the winner of encounter competition is species-specific advantages that increase their RHP, as bobcats ( Lynx rufus ) and spotted skunks ( Spilogale gracilis ) won more often than predicted based on mass. In complex organisms, such as mesocarnivores, species-specific adaptations, including strategic behaviors, aggressiveness, and weapons, contribute to competitive advantages and may allow certain species to take control or defend resources better than others. Our results help explain how interspecific competition shapes the occurrence patterns of species in ecological communities. This article is protected by copyright. All rights reserved.

Description: Ecological theory suggests that pathogens are capable of regulating or limiting host population dynamics, and this relationship has been empirically established in several settings. However, although studies of childhood diseases were integral to the development of disease ecology, few studies show population limitation by a disease affecting juveniles. Here, we present empirical evidence that disease in lambs constrains population growth in bighorn sheep ( Ovis canadensis ) based on 45 years of population-level and 18 years of individual-level monitoring across 12 populations. While populations generally increased (lambda =1.11) prior to disease introduction, most of these same populations experienced an abrupt change in trajectory at the time of disease invasion, usually followed by stagnant-to-declining growth rates (lambda = 0.98) over the next twenty years. Disease-induced juvenile mortality imposed strong constraints on population growth that were not observed prior to disease introduction, even as adult survival returned to pre-invasion levels. Simulations suggested that models including persistent disease-induced mortality in juveniles qualitatively matched observed population trajectories, whereas models that only incorporated all-age disease events did not. We use these results to argue that pathogen persistence may pose a lasting, but under-recognized, threat to host populations, particularly in cases where clinical disease manifests primarily in juveniles. This article is protected by copyright. All rights reserved.

Description: Global environmental change (GEC) is affecting species interactions and causing a rapid decline in biodiversity. In this study, I present a new Ecosystem Disruption Index (EDI) to quantify the impacts of simulated nitrogen (N) deposition (0, 10, 20 and 50 kg N ha -1 yr -1 + 6-7 kg N ha -1 yr -1 background) on abiotic and biotic ecological interactions. This comparative index is based on pairwise linear and quadratic regression matrices. These matrices, calculated at the N treatment level, were constructed using a range of abiotic and biotic ecosystem constituents: soil pH, shrub cover, and the first component of several separate principal component analyses using soil fertility data (total carbon and N) and community data (annual plants; microorganisms; biocrusts; edaphic fauna) for a total of seven ecosystem constituents. Four years of N fertilization in a semiarid shrubland completely disrupted the network of ecological interactions, with a greater proportional increase in ecosystem disruption at low-N addition levels. Biotic interactions, particularly those involving microbes, shrubs and edaphic fauna, were more prone to be lost in response to N, whereas interactions involving soil properties were more resilient. In contrast, edaphic fauna was the only group directly affected by N addition, with mites and collembolans increasing their abundance with up to 20 kg N ha -1 yr -1 and then decreasing, which supports the idea of higher-trophic level organisms being more sensitive to disturbance due to more complex links with other ecosystem constituents. Future experimental studies evaluating the impacts of N deposition, and possibly other GEC drivers, on biodiversity and biotic and abiotic interactions may be able to explain results more effectively in the context of ecological networks as a key feature of ecosystem sensitivity. This article is protected by copyright. All rights reserved.

Description: Edaphic variation in plant community composition is widespread, yet its underlying mechanisms are rarely understood and often assumed to be physiological. In East African savannas, Acacia tree species segregate sharply across soils of differing parent material: the ant-defended whistling thorn, A. drepanolobium (ACDR), is monodominant on clay vertisols that are nutrient rich but physically stressful, whereas poorly defended species such as A. brevispica (ACBR) dominate on nutrient-poor but otherwise less-stressful sandy loams. Using a series of field experiments, we show that large-mammal herbivory interacts with soil properties to maintain this pattern. In the absence of large herbivores, transplanted saplings of both species established on both soil types. Browsers strongly suppressed survival and growth of ACDR saplings on sandy soil, where resource limitation constrained defensive investment. On clay soil, ACBR saplings established regardless of herbivory regime, but elephants prevented recruitment to maturity, apparently because trees could not tolerate the combination of biotic and abiotic stressors. Hence, each tree species was filtered out of one habitat by browsing in conjunction with different edaphic factors and at different ontogenetic stages. Browser abundance was greater on sandy soil, where trees were less defended, consistent with predicted feedbacks between plant community assembly and herbivore distributions. By exploring two inversely related axes of soil “quality” (abiotic stress and nutrient content), our study extends the range of mechanisms by which herbivores are known to promote edaphic specialization, illustrates how the high cost of a protection mutualism can constrain the realized niche of host trees, and shows that large-scale properties of savanna ecosystems are shaped by species interactions in cryptic ways that mimic simple abiotic determinism. These results suggest that ongoing declines in large-herbivore populations may relax spatial heterogeneity in plant assemblages and reduce the beta diversity of communities. This article is protected by copyright. All rights reserved.

Description: Controversy exists over the cause and timing of the extinction of the Pleistocene megafauna. In the tropical Andes, deglaciation and associated rapid climate change began c. 8000 years before human arrival, providing an opportunity to separate the effects of climate change from human hunting on megafaunal extinction. We present a paleoecological record spanning the last 25,000 years from Lake Pacucha, Peru (3100 m elevation). Fossil pollen, charcoal, diatoms, and the dung fungus Sporormiella, chronicle a two-stage megaherbivore population collapse. Sporormiella abundance, the proxy for megafaunal presence, fell sharply at c. 21,000 years ago, but rebounded prior to a permanent decline between c. 16,800 and 15,800 years ago. This two-stage decline in megaherbivores resulted in a functional extinction by c. 15,800 years ago, 3000 years earlier than human occupation of the high Andes. Declining megaherbivore populations coincided with warm, wet intervals. Climatic instability and megafaunal population collapse triggered an ecological cascade that resulted in novel floral assemblages, and increases in woody species, fire frequency, and plant species that were sensitive to trampling. Our data revealed that Andean megafaunal populations collapsed due to positive feedbacks between habitat quality and climate change rather than human activity. This article is protected by copyright. All rights reserved.

Description: Increasing temperatures and a reduction in the frequency and severity of freezing events have been linked to species distribution shifts. Across the globe, mangrove ranges are expanding towards higher latitudes, likely due to diminishing frequency of freezing events associated with climate change. Continued warming will alter coastal wetland plant dynamics both above- and belowground, potentially altering plant capacity to keep up with sea level rise. We conducted an in-situ warming experiment, in Northeast Florida, to determine how increased temperature (+2°C) influences co-occurring mangrove and salt marsh plants. Warming was achieved using passive warming with three treatment levels (ambient, shade control, warmed). Avicennia germinans , the black mangrove , exhibited no differences in growth or height due to experimental warming but displayed a warming-induced increase in leaf production (48%). Surprisingly, Distichlis spicata, the dominant salt marsh grass , increased in biomass (53% in 2013 and 70% in 2014), density (41%) and height (18%) with warming during summer months. Warming decreased plant root mass at depth and changed abundances of anaerobic bacterial taxa. Even while the poleward shift of mangroves is clearly controlled by the occurrences of severe freezes, chronic warming between these freeze events may slow the progression of mangrove dominance within ecotones. This article is protected by copyright. All rights reserved.

Description: In terrestrial ecosystems, a large portion (20-80%) of the dissolved Si (DSi) in soil solution has passed through vegetation. While the importance of this ‘terrestrial Si filter’ is generally accepted, few data exist on the pools and fluxes of Si in forest vegetation and the rate of release of Si from decomposing plant tissues. We quantified the pools and fluxes of Si through vegetation and coarse woody debris (CWD) in a northern hardwood forest ecosystem (Watershed 6, W6) at the Hubbard Brook Experimental Forest (HBEF) in New Hampshire, USA. Previous work suggested that the decomposition of CWD may have significantly contributed to an excess of DSi reported in stream-waters following experimental deforestation of Watershed 2 (W2) at the HBEF. We found that woody biomass (wood + bark) and foliage account for approximately 65% and 31%, respectively, of the total Si in biomass at the HBEF. During the decay of American beech ( Fagus grandifolia ) boles, Si loss tracked the whole-bole mass loss, while yellow birch ( Betula alleghaniensis ) and sugar maple ( Acer saccharum ) decomposition resulted in a preferential Si retention of up to 30% after 16 years. A power-law model for the changes in wood and bark Si concentrations during decomposition, in combination with an exponential model for whole-bole mass loss, successfully reproduced Si dynamics in decaying boles. Our data suggest that a minimum of 50% of the DSi annually produced in the soil of a biogeochemical reference watershed (W6) derives from biogenic Si (BSi) dissolution. The major source is fresh litter, while only ~2% comes from the decay of CWD. Decay of tree boles could only account for 9% of the excess DSi release observed following the experimental deforestation of W2. Therefore, elevated DSi concentrations after forest disturbance are largely derived from other sources (e.g., dissolution of BSi from forest floor soils and/or mineral weathering). This article is protected by copyright. All rights reserved.

Description: Temperature can play an important role in determining the feeding preferences of ectotherms. In light of the warmer temperatures arising with the current climatic changes omnivorous ectotherms may perform diet shifts towards higher herbivory to optimize energetic intake. Such diet shifts may also occur during heat waves, which are projected to become more frequent, intense and longer lasting in the future. Here, we investigated how heat waves of different duration affect feeding preferences in omnivorous anuran tadpoles and how these choices affect larval life history. In laboratory experiments, we fed tadpoles of three species on animal, plant or mixed diet and exposed them to short heat waves (similar to the heat waves these species experience currently) or long heat waves (predicted to increase under climate change). We estimated the dietary choices of tadpoles fed on the mixed diet using stable isotopes and recorded tadpole survival and growth, larval period and mass at metamorphosis. Tadpole feeding preferences were associated with their thermal background, herbivory increasing with breeding temperature in nature. Patterns in survival, growth and development generally support decreased efficiency of carnivorous diets and increased efficiency or higher relative quality of herbivorous diets at higher temperatures. All three species increased herbivory in at least one of the heat wave treatments, but the responses varied among species. Diet shifts towards higher herbivory were maladaptive in one species, but beneficial in the other two. Higher herbivory in omnivorous ectotherms under warmer temperatures may impact species differently and further contribute to changes in the structure and function of freshwater environments. This article is protected by copyright. All rights reserved.

Description: Environmental filtering, including the influence of environmental constraints and biological interactions on species’ survival, is known to significantly affect patterns of community assembly in terrestrial ecosystems. However, its role in regulating patterns and processes of community assembly in deep-sea environments is poorly studied. Here, we investigated the role of wood characteristics in the assembly of deep-sea wood fall communities. Ten different wood species (substrata) that varied in structural complexity were sunk to a depth of 3,100 m near Monterey Bay, CA. In total, 28 wood parcels were deployed on the deep-sea bed. After 2 years, the wood parcels were recovered with over 7,000 attached or colonizing macroinvertebrates. All macroinvertebrates were identified to the lowest taxonomic level possible, and included several undescribed species. Diversity indices and multivariate analyses of variance detected significant variation in the colonizing community assemblages among different wood substrata. Structural complexity seemed to be the primary factor altering community composition between wood substrata. For example, wood-boring clams were most abundant on solid logs, while small arthropods and limpets were more abundant on bundles of branches that provided more surface area and small, protected spaces to occupy. Other factors such as chemical defenses, the presence of bark, and wood hardness likely also played a role. Our finding that characteristics of woody debris entering the marine realm can have significant effects on community assembly supports the notion of ecological and perhaps evolutionarily significant links between land and sea. This article is protected by copyright. All rights reserved.

Description: Central to understanding global C cycle dynamics is the functional relationship between precipitation and net primary production (NPP). At large spatial (regional) scales, the responsiveness of aboveground NPP (ANPP) to interannual variation in annual precipitation (AP; ANPP sens ) is inversely related to site-level ANPP, coinciding with turnover of plant communities along precipitation gradients. Within ecosystems experiencing chronic alterations in water availability, plant community change will also occur with unknown consequences for ANPP sens . To examine the role plant community shifts may play in determining alterations in site-level ANPP sens , we experimentally increased precipitation by ~35% for two decades in a native Central U.S. grassland. Consistent with regional models, ANPP sens decreased initially as water availability and ANPP increased. However, ANPP sens shifted back to ambient levels when mesic species increased in abundance in the plant community. Similarly, in grassland sites with distinct mesic and xeric plant communities and corresponding 50% differences in ANPP, ANPP sens did not differ over almost three decades. We conclude that responses in ANPP sens to chronic alterations in water availability within an ecosystem may not conform to regional AP–ANPP patterns, despite expected changes in ANPP and plant communities. The result is unanticipated functional resistance to climate change at the site scale.

Description: The challenges posed by observing host–pathogen–environment interactions across large geographic extents and over meaningful time scales limit our ability to understand and manage wildland epidemics. We conducted a landscape-scale, longitudinal study designed to analyze the dynamics of sudden oak death (an emerging forest disease caused by Phytophthora ramorum ) across hierarchical levels of ecological interactions, from individual hosts up to the community and across the broader landscape. From 2004 to 2011, we annually assessed disease status of 732 coast live oak, 271 black oak, and 122 canyon live oak trees in 202 plots across a 275-km 2 landscape in central California. The number of infected oak stems steadily increased during the eight-year study period. A survival analysis modeling framework was used to examine which level of ecological heterogeneity best predicted infection risk of susceptible oak species, considering variability at the level of individuals (species identity, stem size), the community (host density, inoculum load, and species richness), and the landscape (seasonal climate variability, habitat connectivity, and topographic gradients). After accounting for unobserved risk shared among oaks in the same plot, survival models incorporating heterogeneity across all three levels better predicted oak infection than did models focusing on only one level. We show that larger oak trees (especially coast live oak) were more susceptible, and that interannual variability in inoculum production by the highly infectious reservoir host, California bay laurel, more strongly influenced disease risk than simply the density of this important host. Concurrently, warmer and wetter rainy-season conditions in consecutive years intensified infection risk, presumably by creating a longer period of inoculum build-up and increased probability of pathogen spillover from bay laurel to oaks. Despite the presence of many alternate host species, we found evidence of pathogen dilution, where less competent hosts in species-rich communities reduce pathogen transmission and overall risk of oak infection. These results identify key parameters driving the dynamics of emerging infectious disease in California woodlands, while demonstrating how multiple levels of ecological heterogeneity jointly determine epidemic trajectories in wildland settings.

Description: Although genetics in a single species is known to impact whole communities, little is known about how genetic variation influences species interaction networks in complex ecosystems. Here, we examine the interactions in a community of arthropod species on replicated genotypes (clones) of a foundation tree species, Populus angustifolia James (narrowleaf cottonwood), in a long-term, common garden experiment using a bipartite “genotype–species” network perspective. We combine this empirical work with a simulation experiment designed to further investigate how variation among individual tree genotypes can impact network structure. Three findings emerged: (1) the empirical “genotype–species network” exhibited significant network structure with modularity being greater than the highly conservative null model; (2) as would be expected given a modular network structure, the empirical network displayed significant positive arthropod co-occurrence patterns; and (3) furthermore, the simulations of “genotype–species” networks displayed variation in network structure, with modularity in particular clearly increasing, as genotypic variation increased. These results support the conclusion that genetic variation in a single species contributes to the structure of ecological interaction networks, which could influence ecological dynamics (e.g., assembly and stability) and evolution in a community context.

Description: The latitudinal diversity gradient (LDG) has been one of the most documented patterns in ecology, typically showing decreasing species diversity with increasing latitude. Studies of these patterns also used different spatial scales and dispersal traits to better understand the underpinning ecological factors. Seasonal freshwater ecosystems are less studied and may exhibit different patterns because they are more sensitive to climatic variation, which result in an inundation–desiccation cycle. In California, precipitation increases and temperature decreases with increasing latitude and thus the LDG pattern may be associated with this climatic gradient. Using collected data and United States Fish and Wildlife Service reports across seven degrees of latitude, analysis of California vernal pool invertebrate community (total richness and richness of passive and active dispersers) was conducted using correlations (Spearman rank and partial). Alpha diversity (total and passive dispersers) increased and beta diversity (passive dispersers) decreased with increasing latitude. Vernal pool surface area was correlated with active disperser alpha and passive disperser beta diversity. This suggests that climate-driven habitat size influences alpha and beta diversity patterns depending on dispersal ability. Active dispersers and predators exhibited higher beta diversity than passive dispersers and prey, respectively. Species composition differed among counties and some of these differences were correlated with pool depth and temperature. These results suggest that seasonal habitats will have diversity patterns strongly associated with local scale characteristics (habitat size and hydroperiod) determined by climate variation along the latitudinal gradient. Understanding these diversity patterns along the gradient will also contribute to management and restoration of these ecosystems with high endemism and diversity.

Description: Ecological theory predicts that positive interactions among organisms will increase across gradients of increasing abiotic stress or consumer pressure. This theory has been supported by empirical studies examining the magnitude of ecosystem engineering across environmental gradients and between habitat settings at local scale. Predictions that habitat setting, by modifying both biotic and abiotic factors, will determine large-scale gradients in ecosystem engineering have not been tested, however. A combination of manipulative experiments and field surveys assessed whether along the east Australian coastline: (1) facilitation of invertebrates by the oyster Saccostrea glomerata increased across a latitudinal gradient in temperature; and (2) the magnitude of this effect varied between intertidal rocky shores and mangrove forests. It was expected that on rocky shores, where oysters are the primary ecosystem engineer, they would play a greater role in ameliorating latitudinal gradients in temperature than in mangroves, where they are a secondary ecosystem engineer living under the mangrove canopy. On rocky shores, the enhancement of invertebrate abundance in oysters as compared to bare microhabitat decreased with latitude, as the maximum temperatures experienced by intertidal organisms diminished. By contrast, in mangrove forests, where the mangrove canopy resulted in maximum temperatures that were cooler and of greater humidity than on rocky shores, we found no evidence of latitudinal gradients of oyster effects on invertebrate abundance. Contrary to predictions, the magnitude by which oysters enhanced biodiversity was in many instances similar between mangroves and rocky shores. Whether habitat-context modifies patterns of spatial variation in the effects of ecosystem engineers on community structure will depend, in part, on the extent to which the environmental amelioration provided by an ecosystem engineer replicates that of other co-occurring ecosystem engineers.

Description: Many areas of research in ecology and evolutionary biology depend on the quantification of dietary niche width. For herbivorous insects, diet breadth has most often been measured as simply the number and type of host plant taxa attacked. We propose an index of host range (which we refer to as “ordinated diet breadth”) based on observed associations between plants and insects, and the calculation of multivariate distances among plants in ordination space. Similarities and distances are calculated based on host association and, in this context, potentially encompass multiple properties of plants, including phytochemistry, phenology, and other plant traits. This approach can distinguish between herbivores that utilize suites of hosts that are commonly used together and herbivores that attack unusual host combinations, and thus have a relatively broad diet breadth. For illustration, we use a data set of nymphalid butterfly host records, and compare taxonomic and ordinated host range. For a large number of butterfly taxa, we find that host use is clustered in multivariate space with respect to associations observed across all of the butterfly taxa. Applications are discussed, including a hypothesis test of nonrandom host association, and prediction of shifts and expansions of diet breadth.

Description: Environmental change that disrupts communication during mate choice and alters sexual selection could influence population dynamics. Yet little is known about such long-term effects. We investigated experimentally the consequences that disrupted visual communication during mate choice has for the quantity and viability of offspring produced in a threespine stickleback population ( Gasterosteus aculeatus ). We further related the results to long term monitoring of population dynamics in the field to determine if changes are apparent under natural conditions. The results show that impaired visual communication because of algal blooms reduces reliability of male visual signals as indicators of offspring survival during their first weeks of life. This relaxes sexual selection but has no effect on the number of offspring hatching, as most males have a high hatching success in turbid water. Despite eutrophication and high turbidity levels that interfere with communication during mate choice, the population has grown during recent decades. Large numbers of offspring hatching, combined with high variation in juvenile fitness, has probably shifted selection to later life history stages and maintained a viable population. Together with reduced cost of sexual selection and ongoing ecosystem changes caused by human activities, this could have promoted population growth. These results point to the complexity of ecosystems and the necessity to consider all influencing factors when attempting to understand impacts of human activities on populations.

Description: Predators play a key role in shaping natural ecosystems, and understanding the factors that influence a predator's kill rate is central to predicting predator–prey dynamics. While prey density has a well-established effect on predation, it is increasingly apparent that predator density also can critically influence predator kill rates. The effects of both prey and predator density on the functional response will, however, be determined in part by their distribution on the landscape. To examine this complex relationship we experimentally manipulated prey density, predator density, and prey distribution using a tadpole (prey)–dragonfly nymph (predator) system. Predation was strongly ratio-dependent irrespective of prey distribution, but the shape of the functional response changed from hyperbolic to sigmoidal when prey were clumped in space. This sigmoidal functional response reflected a relatively strong negative effect of predator interference on kill rates at low prey: predator ratios when prey were clumped. Prey aggregation also appeared to promote stabilizing density-dependent intraguild predation in our system. We conclude that systems with highly antagonistic predators and patchily distributed prey are more likely to experience stable dynamics, and that our understanding of the functional response will be improved by research that examines directly the mechanisms generating interference.

Description: Because it modulates the fitness returns of possible options of energy expenditure at each ontogenetic stage, environmental stochasticity is usually considered a selective force in driving or constraining possible life histories. Divergent regimes of environmental fluctuation experienced by populations are expected to generate differences in the resource allocation schedule between survival and reproductive effort and outputs. To our knowledge, no study has previously examined how different regimes of stochastic variation in environmental conditions could result in changes in both the temporal variation and mean of demographic parameters, which could then lead to intraspecific variation along the slow–fast continuum of life history tactics. To investigate these issues, we used capture–recapture data collected on five populations of a long-lived amphibian ( Bombina variegata ) experiencing two distinct levels of stochastic environmental variation: (1) constant availability of breeding sites in space and time (predictable environment), and (2) variable spatio-temporal availability of breeding sites (unpredictable environment). We found that female breeding propensity varied more from year to year in unpredictable than in predictable environments. Although females in unpredictable environments produced on average more viable offspring per year, offspring production was more variable between years. Survival at each ontogenetic stage was slightly lower and varied significantly more from year to year in unpredictable environments. Taken together, these results confirm that increased environmental stochasticity can modify the resource allocation schedule between survival and reproductive effort and outputs and may lead to intraspecific variation along the slow–fast continuum of life history tactics.

Description: Plant–plant and plant–soil interactions can help maintain plant diversity and ecosystem functions. Changes in these interactions may underlie experimentally observed increases in biodiversity effects over time via the selection of genotypes adapted to low or high plant diversity. Little is known, however, about such community-history effects and particularly the role of plant–soil interactions in this process. Soil-legacy effects may occur if co-evolved interactions with soil communities either positively or negatively modify plant biodiversity effects. We tested how plant selection and soil legacy influence biodiversity effects on productivity, and whether such effects increase the resistance of the communities to invasion by weeds. We used two plant selection treatments: parental plants growing in monoculture or in mixture over 8 yr in a grassland biodiversity experiment in the field, which we term monoculture types and mixture types. The two soil-legacy treatments used in this study were neutral soil inoculated with live or sterilized soil inocula collected from the same plots in the biodiversity experiment. For each of the four factorial combinations, seedlings of eight species were grown in monocultures or four-species mixtures in pots in an experimental garden over 15 weeks. Soil legacy (live inoculum) strongly increased biodiversity complementarity effects for communities of mixture types, and to a significantly weaker extent for communities of monoculture types. This may be attributed to negative plant–soil feedbacks suffered by mixture types in monocultures, whereas monoculture types had positive plant–soil feedbacks, in both monocultures and mixtures. Monocultures of mixture types were most strongly invaded by weeds, presumably due to increased pathogen susceptibility, reduced biomass, and altered plant–soil interactions of mixture types. These results show that biodiversity effects in experimental grassland communities can be modified by the evolution of positive vs. negative plant–soil feedbacks of plant monoculture vs. mixture types.

Description: The establishment of protected areas or parks has become an important tool for wildlife conservation. However, frequent occurrences of human–wildlife conflict at the edges of these parks can undermine their conservation goals. Many African protected areas have experienced concurrent declines of apex predators alongside increases in both baboon abundance and the density of humans living near the park boundary. Baboons then take excursions outside of the park to raid crops for food, conflicting with the human population. We model the interactions of mesopredators (baboons), apex predators, and shared prey in the park to analyze how four components affect the proportion of time that mesopredators choose to crop-raid: (1) the presence of apex predators; (2) nutritional quality of the crops; (3) mesopredator “shyness” about leaving the park; and (4) human hunting of mesopredators. We predict that the presence of apex predators in the park is the most effective method for controlling mesopredator abundance, and hence significantly reduces their impact on crops. Human hunting of mesopredators is less effective as it only occurs during crop-raiding excursions. Furthermore, making crops less attractive, for instance by planting crops further from the park boundary or farming less nutritional crops, can reduce the amount of time mesopredators crop-raid.

Description: Predictive classifications of life-histories are essential for evolutionary ecology. While attempts to apply a single approach to all organisms may be overambitious, recent advances suggest that more narrow ordination schemes can be useful. However, these schemes mostly lack easily observable proxies of the position of a species on respective axes. It has been proposed that, in insects, the degree of capital (vs income) breeding – reflecting the importance of adult feeding for reproduction – correlates with various ecological traits at the level of among-species comparison. We sought to prove these ideas via rigorous phylogenetic comparative analyses. We used experimentally derived life-history data for 57 species of European Geometridae (Lepidoptera), and an original phylogenetic reconstruction. The degree of capital breeding was estimated based on morphological proxies, including relative abdomen size of females. Applying Brownian motion based comparative analyses (with an original update to include error estimates), we demonstrated the associations between the degree of capital breeding and larval diet breadth, sexual size dimorphism and reproductive season. Ornstein-Uhlenbeck model based phylogenetic analysis suggested a causal relationship between the degree of capital breeding and diet breadth. Our study indicates that the gradation from capital to income breeding is an informative axis to ordinate life-history strategies in flying insects affected by the fecundity vs mobility trade off, with the availability of easy to record proxies contributing to its predictive power in practical contexts. This article is protected by copyright. All rights reserved.

Description: Despite increasing evidence that herbivory on a focal plant may hinge upon the identity of its neighbors, it is not clear whether predictable mechanisms govern the nature and magnitude of such associational effects. Using a factorial field experiment replicated at 14 sites across 80,000 hectares, we evaluated the mechanisms driving associational effects between two plant species mediated by grasshopper herbivores. Our experiment manipulated local neighborhood plant density (two levels) and frequency (three levels), nested within two larger-scale habitat contexts (habitats that did or did not have past agricultural land use). We found that the more-palatable plant species, Solidago nemoralis, experienced reduced herbivory (associational resistance) when rare due mainly to reduced grasshopper foraging activity. Damage to the less-palatable plant species, S. odora , was affected by the interaction between plant frequency and the land-use history of the site: it experienced increased damage (associational susceptibility) in even-frequency neighborhoods, but only in sites with a history of past agricultural use. Behavioral assays generally corroborated the results from the field, further supporting the importance of foraging behavior in generating associational effects. In finding that associational effects are contingent upon relative palatability among plants and events in the distant past that modify contemporary habitat structure (i.e., past agricultural land use), our work suggests that foraging decisions made at the neighborhood level are important for generating associational effects and that in some cases these neighborhood interactions also depend on the larger-scale environmental context resulting from legacies of past land-use events. This article is protected by copyright. All rights reserved.

Description: The role of theory within ecology has changed dramatically in recent decades. Once primarily a source of qualitative conceptual framing, ecological theories and models are now often used to develop quantitative explanations of empirical patterns and to project future dynamics of specific ecological systems. In this essay, I recount my own experience of this transformation, in which accelerating computing power and the widespread incorporation of stochastic processes into ecological theory combined to create some novel integration of mathematical and statistical models. This stronger integration drives theory towards incorporating more biological realism, and I explore ways in which we can grapple with that realism to generate new general theoretical insights. This enhanced realism, in turn, may lead to frameworks for projecting ecological responses to anthropogenic change, which is, arguably, the central challenge for 21st-century ecology. In an era of big data and synthesis, ecologists are increasingly seeking to infer causality from observational data; but conventional biometry provides few tools for this project. This is a realm where theorists can and should play an important role, and I close by pointing towards some analytical and philosophical approaches developed in our sister discipline of economics that address this very problem. While I make no grand prognostications about the likely discoveries of ecological theory over the coming century, you will find in this essay a scattering of more or less far-fetched ideas that I, at least, think are interesting and (possibly) fruitful directions for our field.

Description: Extreme weather events, such as drought, have marked impacts on biotic communities. In many regions, a predicted increase in occurrence of such events will be imposed on landscapes already heavily modified by human land use. There is an urgency, therefore, to understand the way in which the effects of such events may be exacerbated, or moderated, by different patterns of landscape change. We used empirical data on woodland-dependent birds in southeast Australia, collected during and after a severe drought, to document temporal change in the composition of bird assemblages in 24 landscapes (each 100 km 2 ) representing a gradient in the cover of native wooded vegetation (from 60% to 〈2%). We examined (a) whether drought caused region-wide homogenization of the composition of landscape bird assemblages, and (b) whether landscape properties influenced the way assemblages changed in response to drought. To quantify change, we used pairwise indices of assemblage dissimilarity, partitioned into components that represented change in the richness of assemblages and change in the identity of constituent species (turnover). There was widespread loss of woodland birds in response to drought, with only partial recovery following drought-breaking rains. Region-wide, the composition of landscape assemblages became more different over time, primarily caused by turnover-related differentiation. The response of bird assemblages to drought varied between landscapes and was strongly associated with landscape properties. The extent of wooded vegetation had the greatest influence on assemblage change: landscapes with more native vegetation had more stable bird assemblages over time. However, for the component processes of richness- and turnover-related compositional change, measures of landscape productivity had a stronger effect. For example, landscapes with more riparian vegetation maintained more stable assemblages in terms of richness. These results emphasize the importance of the total extent of native vegetation, both overall cover and that occurring in productive parts of the landscape, for maintaining bird communities whose composition is resistant to severe drought. While extreme climatic events cannot be prevented, their effects can be ameliorated by managing the pattern of native vegetation in anthropogenic landscapes, with associated benefits for maintaining ecological processes and human well-being.

Description: Many empirical studies of metacommunities have focused on the classification of observational patterns into four contrasting paradigms characterized by different levels of movement and habitat heterogeneity. However, deeper insight into the underlying local and regional processes may be derived from a combination of long-term observational data and experimental studies. With the aim of exploring forces structuring the insect metacommunity on oak, we fit a hierarchical Bayesian state-space model to data from observations and experiments. The fitted model reveals large variation in species-specific dispersal abilities and basic reproduction numbers, R 0 . The residuals from the model show only weak correlations among species, suggesting a lack of strong interspecific interactions. Simulations with model-derived parameter estimates indicate that habitat configuration and species attributes both contribute substantially to structuring insect communities. Overall, our findings demonstrate that community-level variation in movement and life history are key drivers of metacommunity dynamics.

Description: Mussel beds and rockweed stands (fucoid algae) have been shown to be alternative states on rocky intertidal shores in New England, and here the hypothesis that variation in recruitment provides opportunity for the development of alternative community states was tested. Disturbance by ice scour opens patches for development of alternative states, and in winter 1996–1997, 60 experimental clearings of differing sizes were established on Swan's Island, Maine, USA. Half of the plots were re-cleared during the winter of 2010–2011. Recruitment data for barnacles, mussels, and fucoid algae collected from 1997 to 2012 were used to (1) test for persistence of scale-dependent thresholds, (2) estimate the magnitudes and sources of variation, (3) fit a surface of alternative states as defined by the cusp catastrophe, and (4) test if 1997 recruitment would predict 2010–2011 recruitment in re-scraped plots (i.e., a test of divergence, which is expected in systems with alternative states). For barnacles and mussels, recruitment varied enormously year to year and among sites, but showed consistent patterns over the long-term with respect to clearing size. Average recruitment prior to re-clearing was a good predictor of recruitment afterwards. In contrast, over 50% of the variance in fucoid recruitment was unexplained with weak effects among years and locations. Past fucoid recruitment was a poor predictor of subsequent recruitment. The cusp analysis indicated that fucoid recruitment defines the alternative states. Fucoid recruitment was largely unpredictable and suggests long-term, small-scale priority effects drive the development of alternative states. These observations strongly reinforce the notion that long-term and well-replicated experiments are necessary to develop robust tests of ecological theory.

Description: Although the effects of individual age, resource availability, and reproductive costs have been extensively studied to understand the causes of variation in reproductive output, there are almost no studies showing how these factors interact in explaining this variation. To examine this interaction, we used longitudinal demographic data from an 18-year study of 53 breeding female wolverines ( Gulo gulo ), and corresponding environmental data from their individual home ranges. Females showed a typical age-related pattern in reproductive output, with an initial increase followed by a senescent decline in later years. This pattern was largely driven by four processes: (1) physiological/behavioral maturation between ages two and three; (2) age-related differences in the costs of reproduction resulting in an initial increase, and then a declining probability of breeding two years in a row as individuals aged; (3) resource availability (reindeer [ Rangifer tarandus ] carcass abundance; mostly Eurasian lynx [ Lynx lynx ] kills) in the months preceding parturition, which influenced the probability of having cubs, but only for individuals that had successfully bred in the previous year; and (4) resource availability also influenced the cost of reproduction in an age-dependent manner, as prime age females that had bred in the previous year were more responsive to resource availability than those at other ages. This study demonstrates that by examining how drivers of reproductive variation interact, we can get a much clearer understanding of the mechanisms responsible for age-related patterns of reproduction. This has implications not only for general ecological theory, but will also allow better predictions of population responses to environmental changes or management based on a population's age-structure.

Description: Ecological theory predicts that demographic connectivity structures the dynamics of local populations within metapopulation systems, but empirical support has been constrained by major limitations in data and methodology. We tested this prediction for giant kelp Macrocystis pyrifera , a key habitat-forming species in temperate coastal ecosystems worldwide, in southern California, USA. We combined a long-term (22 years), large-scale (~500 km coastline), high-resolution census of abundance with novel patch delineation methods and an innovative connectivity measure incorporating oceanographic transport and source fecundity. Connectivity strongly predicted local dynamics (well-connected patches had lower probabilities of extinction and higher probabilities of colonization, leading to greater likelihoods of occupancy) but this relationship was mediated by patch size. Moreover, the relationship between connectivity and local population dynamics varied over time, possibly due to temporal variation in oceanographic transport processes. Surprisingly, connectivity had a smaller influence on colonization relative to extinction, possibly because local ecological factors differ greatly between extinct and extant patches. Our results provide the first comprehensive evidence that southern California giant kelp populations function as a metapopulation system, challenging the view that populations of this important foundation species are governed exclusively by self-replenishment.

Description: Disruption of natural flow regimes, nutrient pollution, and other consequences of human population growth and development have impacted most major rivers of the world. Alarming losses of aquatic biodiversity coincide with human-caused river alteration, but effects of biotic homogenization on aquatic ecosystem processes are not as well documented. This is because unaltered systems for comparison are scarce, and some ecosystem-wide effects may take decades to manifest. We evaluated aquatic ecosystem responses to extensive river-floodplain engineering and nutrient addition in the Rio Grande of southwestern North America as revealed by changes in trophic structure of, and resource availability to, the fish community. Stable Isotope Analysis (SIA) was conducted on museum-preserved fishes collected over a 70-year period of intensive river management and exponential human population growth. Trophic complexity and resource heterogeneity for fish consumers (measured as “isotopic niche breadth”) decreased following sediment deprivation and channelization, and these effects persist into the present. Increased nutrient inputs led to δ 15 N enrichment in the entire fish community at all affected sites, and a shift to autochthonous sources of carbon at the most proximal site downstream of wastewater release, probably via bottom-up transfer. Overall, retrospective SIA of apex consumers suggests radical change and functional impairment of a floodplain river ecosystem already marked by significant biodiversity loss.

Description: Colonization–extinction dynamics and species sorting among habitats determine the distribution of species within metacommunities. Theory suggests that disturbances reduce the importance of species sorting and enhance spatial patterning and stochastic effects, however this has not yet been experimentally shown. We examined how extinctions in a heterogeneous landscape of patches affects the influence of environmental, spatial, and stochastic factors on community composition in a simple two-species, two-habitat, protist metacommunity where each species dominates in a different habitat type. We imposed four different levels of random extinctions on local patches and monitored changes in the metacommunity through time. We found that near-steady state patterns of community variability developed relatively rapidly (within nine colonization–extinction cycles) and that increased extinction rate produced altered patterns of community regulation by reducing environmental control and increasing spatial and stochastic effects. Our results indicate a possible explanation for the combination of environmental, spatial and stochastic effects observed in natural metacommunities.

Description: The curvature of generalized Holling type functional response curves is controlled by a shape parameter b yielding hyperbolic type II ( b = 1) to increasingly sigmoid type III ( b 〉 1) responses. Empirical estimates of b vary considerably across taxa. Larger consumer–resource body mass ratios have been suggested to generate more pronounced type III responses and therefore to promote dynamic stability. The dependence of consumer–resource stability on b has, however, not been systematically explored, and the accurate empirical determination of b is challenging. Specifically, the shape of the functional response of the pelagic grazer Daphnia feeding on phytoplankton, and its consequences for stability, remain controversial. We derive a novel analytical condition relating b to local stability of consumer–resource interactions and use it to predict stability of empirically parameterized models of Daphnia and phytoplankton under enrichment. Functional response parameters were experimentally derived for two species of Daphnia feeding separately on single cultures of two different phytoplankton species. All experimentally studied Daphnia –algae systems exhibited type III responses. Parameterized type III responses are predicted to stabilize the modeled Daphnia –phytoplankton dynamics in some species pairs but not in others. Remarkably, stability predictions differ depending on whether functional response parameters are derived from clearance vs. ingestion rates. Accurate parameter estimation may therefore require fitting to both rates. In addition, our estimates of b for filter-feeding Daphnia are much smaller than predicted for actively hunting predators at similar consumer–resource body mass ratios. This suggests that the relationship between functional response shape and body mass ratios may vary with predation mode.

Description: Temperature imposes a constraint on the rates and outcomes of ecological processes that determine community- and ecosystem-level patterns. The application of metabolic scaling theory has advanced our understanding of the influence of temperature on pattern and process in marine communities. Metabolic scaling theory uses the fundamental and ubiquitous patterns of temperature-dependent metabolism to predict how environmental temperature influences patterns and processes at higher levels of biological organization. Here, we outline some of these predictions to review recent advances and illustrate how scaling theory might be applied to new challenges. For example, warming can alter species interactions and food-web structure and can also reduce total animal biomass supportable by a given amount of primary production by increasing animal metabolism and energetic demand. Additionally, within a species, larval development is faster in warmer water, potentially influencing dispersal and other demographic processes like population connectivity and gene flow. These predictions can be extended further to address major questions in marine ecology, and present an opportunity for conceptual unification of marine ecological research across levels of biological organization. Drawing on work by ecologists and oceanographers over the last century, a metabolic scaling approach represents a promising way forward for applying ecological understanding to basic questions as well as conservation challenges.

Description: Ecosystems are subsidized with inputs of mass and energy from their surroundings. These allochthonous inputs regulate many ecosystem characteristics. In inland waters, terrestrial inputs of organic matter regulate food-web structure, ecosystem metabolism, water clarity, and thermal stratification. Future changes in allochthony may be especially pronounced in high-elevation ecosystems due to increases in vegetation and precipitation associated with climate change. Several techniques exist to characterize the degree of allochthony of organic matter in aquatic systems, including metrics such as Δ H , the net isotopic discrimination between water and particulate organic matter (POM) of deuterium stable isotopes, and the fluorescence index (FI), which characterizes the fluorescence of dissolved organic matter (DOM). Despite the importance of allochthonous organic carbon inputs, little is known about either how allochthony varies across elevation gradients or whether different metrics are similarly related to allochthony. We measured Δ H , FI, and a suite of related water-quality characteristics in 30 lakes across a montane to alpine elevation gradient (2340 to 3205 m) in the Beartooth Mountains of Montana and Wyoming, USA, to understand how FI and Δ H varied with elevation, with one another, and with other allochthony-related water-quality characteristics. We hypothesized that allochthony of POM and DOM would decrease at higher elevations, with alpine lakes above treeline being more autochthonous compared with low-elevation lakes below treeline. We observed a significant inverse linear relationship between Δ H and FI, with both metrics indicating a decrease in allochthony at higher elevations. Characteristics including the natural log of the ratio of concentrations of dissolved organic carbon to chlorophyll a (ln(DOC : Chl)), the spectral slope ratio between different spectra of two wavebands ( S R , ratio of spectra at 275–295 to 350–400 nm), and a ratio of diffuse attenuation coefficients at 320 and 380 nm ( K R , K d320   :   K d380 ) varied with both Δ H and FI while pH varied only with Δ H . High-elevation systems were characterized by low ln(DOC : Chl) and K R , and high S R and pH. These results indicate that high-elevation lakes are more autochthonous than low-elevation lakes. The relationships among Δ H , FI, elevation, and other water-quality characteristics provide important insights to understand future changes in carbon cycling in mountain ecosystems.

Description: Explaining variability in offspring vs. adult size among groups is a necessary step to determine the evolutionary and environmental constraints shaping variability in life history strategies. This is of particular interest for life in the ocean where a diversity of offspring development strategies is observed along with variability in physical and biological forcing factors in space and time. We compiled adult and offspring size for 407 pelagic marine species covering more than 17 orders of magnitude in body mass including Cephalopoda, Cnidaria, Crustaceans, Ctenophora, Elasmobranchii, Mammalia, Sagittoidea, and Teleost. We find marine life following one of two distinct strategies, with offspring size being either proportional to adult size (e.g., Crustaceans, Elasmobranchii, and Mammalia) or invariant with adult size (e.g., Cephalopoda, Cnidaria, Sagittoidea, Teleosts, and possibly Ctenophora). We discuss where these two strategies occur and how these patterns (along with the relative size of the offspring) may be shaped by physical and biological constraints in the organism's environment. This adaptive environment along with the evolutionary history of the different groups shape observed life history strategies and possible group-specific responses to changing environmental conditions (e.g., production and distribution).

Description: While nonconsumptive effects of predators may strongly affect prey populations, little is known how future warming will modulate these effects. Such information would be especially relevant with regard to prey physiology and resulting changes in prey stoichiometry. We investigated in Enallagma cyathigerum damselfly larvae the effects of a 4°C warming (20°C vs. 24°C) and predation risk on growth rate, physiology and body stoichiometry, for the first time including all key mechanisms suggested by the general stress paradigm (GSP) on how stressors shape changes in body stoichiometry. Growth rate and energy storage were higher at 24°C. Based on thermodynamic principles and the growth rate hypothesis, we could demonstrate predictable reductions in body C:P under warming and link these to the increase in P-rich RNA; the associated warming-induced decrease in C:N may be explained by the increased synthesis of N-rich proteins. Yet, under predation risk, growth rate instead decreased with warming and the warming-induced decreases in C:N and C:P disappeared. As predicted by the GSP, larvae increased body C:N and C:P at 24°C under predation risk. Notably, we did not detect the assumed GSP-mechanisms driving these changes: despite an increased metabolic rate there was neither an increase of C-rich biomolecules (instead fat and sugar contents decreased under predation risk), nor a decrease of N-rich proteins. We hypothesize that the higher C:N and N:P under predation risk are caused by a higher investment in morphological defense. This may also explain the stronger predator-induced increase in C:N under warming. The expected higher C:P under predation risk was only present under warming and matched the observed growth reduction and associated reduction in P-rich RNA. Our integrated mechanistic approach unraveled novel pathways of how warming and predation risk shape body stoichiometry. Key findings that (1) warming effects on elemental stoichiometry were predictable and only present in the absence of predation risk and that (2) warming reinforced the predator-induced effects on C:N:P, are pivotal in understanding how nonconsumptive predator effects under global warming will shape prey populations.

Description: Few studies measure multiple ecological tracers in individual organisms, thus limiting our ability to differentiate among organic matter source pathways and understand consequences of dietary variation and the use of external subsidies in complex food webs. We combined two tracers, stable isotope (SI) ratios and fatty acids (FA), to investigate linkages among ecological compartments (water column, benthos, riparian zone) in food webs in waterholes of a dryland river network, the Border Rivers in southwestern Queensland, Australia. Comprehensive analyses of sources (plankton, periphyton, leaf litter, riparian grasses) and animals (benthic insects, mollusks, large crustaceans, fishes) for SI and FA showed that all three zones contribute to animal biomass, depending on species and life stage. Large fishes derived a subsidy from the riparian/floodplain zone, likely through the consumption of terrestrial and semi-aquatic insects and prawns that fed on detritivorous insects. Importantly, post-larval bony bream ( Nematalosa erebi ) and golden perch ( Macquaria ambigua ) were tightly connected to the water column, as evidenced by 13 C-depleted, 15 N-enriched isotope ratios and a high content of plankton-derived polyunsaturated fatty acids (eicosapentaenoic acid [EPA; 20:5ω3] and docosahexaenoic acid [DHA; 22:6ω3]). These observations were consistent with expectations from nutritional requirements of fish early life stages and habitat changes associated with maturity. These results highlight the importance of high-quality foods during early development of fishes, and show that attempting to attribute food-web production to a single source pathway overlooks important but often subtle subsidies that maintain viable populations. A complete understanding of food-web dynamics must consider both quantity and quality of different available organic matter sources. This understanding can be achieved with a combined SI and FA approach, but more controlled dietary studies are needed to estimate how FA profiles are modified by animals when consuming a diverse range of diets of variable quality.

Description: Despite decades of monitoring global reef decline, we are still largely unable to explain patterns of reef deterioration at local scales, which precludes the development of effective management strategies. Offshore reefs of the Florida Keys, USA, experience milder temperatures and lower nutrient loads in comparison to inshore reefs yet remain considerably more degraded than nearshore patch reefs. A year-long reciprocal transplant experiment of the mustard hill coral ( Porites astreoides ) involving four source and eight transplant locations reveals that corals adapt and/or acclimatize to their local habitat on a 〈10-km scale. Surprisingly, transplantation to putatively similar environmental types (e.g., offshore corals moved to a novel offshore site, or along-shore transplantation) resulted in greater reductions in fitness proxies, such as coral growth, than cross-channel transplantation between inshore and offshore reefs. The only abiotic factor showing significantly greater differences between along-shore sites was daily temperature range extremes (rather than the absolute high or low temperatures reached), providing a possible explanation for this pattern. Offshore-origin corals exhibited significant growth reductions at sites with greater daily temperature ranges, which explained up to 39% of the variation in their mass gain. In contrast, daily temperature range explained at most 9% of growth variation in inshore-origin corals, suggesting that inshore corals are more tolerant of high-frequency temperature fluctuations. Finally, corals incur trade-offs when specializing to their native reef. Across reef locations the coefficient of selection against coral transplants was 0.07 ± 0.02 (mean ± SE). This selection against immigrants could hinder the ability of corals to recolonize devastated reefs, whether through assisted migration efforts or natural recruitment events, providing a unifying explanation for observed patterns of coral decline in this reef system.

Description: Metacommunity theory indicates that variation in local community structure can be partitioned into components including those related to local environmental conditions vs. spatial effects and that these can be quantified using statistical methods based on variation partitioning. It has been hypothesized that joint associations of community composition with environment and space could be due to patch dynamics involving colonization–extinction processes in environmentally heterogeneous landscapes but this has yet to be theoretically shown. We develop a two-patch, type-two, species competition model in such a “harlequin” landscape (where different patches have different environments) to evaluate how composition is related to environmental and spatial effects as a function of background extinction rate. Using spatially implicit analytical models, we find that the environmental association of community composition declines with extinction rate as expected. Using spatially explicit simulation models, we further find that there is an increase in the spatial structure with extinction due to spatial patterning into clusters that are not related to environmental conditions but that this increase is limited. Natural metacommunities often show both environment and spatial determination even under conditions of relatively high isolation and these could be more easily explained by our model than alternative metacommunity models.

Description: When selecting habitats, herbivores must weigh multiple risks, such as predation, starvation, toxicity, and thermal stress, forcing them to make fitness trade-offs. Here, we applied the method of paired comparisons (PC) to investigate how herbivores make trade-offs between habitat features that influence selection of food patches. The method of PC measures utility and the inverse of utility, relative risk, and makes trade-offs and indifferences explicit by forcing animals to make choices between two patches with different types of risks. Using a series of paired-choice experiments to titrate the equivalence curve and find the marginal rate of substitution for one risk over the other, we evaluated how toxin-tolerant (pygmy rabbit Brachylagus idahoensis ) and fiber-tolerant (mountain cottontail rabbit Sylviagus nuttallii ) herbivores differed in their hypothesized perceived risk of fiber and toxins in food. Pygmy rabbits were willing to consume nearly five times more of the toxin 1,8-cineole in their diets to avoid consuming higher levels of fiber than were mountain cottontails. Fiber posed a greater relative risk for pygmy rabbits than cottontails and cineole a greater risk for cottontails than pygmy rabbits. Our flexible modeling approach can be used to (1) quantify how animals evaluate and trade off multiple habitat attributes when the benefits and risks are difficult to quantify, and (2) integrate diverse risks that influence fitness and habitat selection into a single index of habitat value. This index potentially could be applied to landscapes to predict habitat selection across several scales.

Description: Spatial variation in pollinator communities or behaviors can underlie floral diversification. Floral traits in the UV spectrum are common and mediate plant–pollinator interactions, but the role of pollinators in driving or maintaining their geographic variation has not been fully explored. We identify an altitudinal cline of increasing relative size of the UV bullseye pattern in Argentina anserina (Rosaceae) flowers, and assess whether pollination context contributes to clinal variation. At four sites that varied in altitude, we document the pollinator assemblage, and pollinator preference and visitation behavior. We then determine how pollinator visits affected pollen receipt and export. Finally, we describe how the functional relationship between UV floral phenotype and pollen receipt changed with altitude. Floral UV bullseye size increased with altitude, which corresponded with a change from a hymenopteran- to a dipteran-dominated pollinator assemblage. While dipteran and hymenopteran preferences for bullseye size were similar, flowers with large bullseyes received more foraging visits than those with small bullseyes at higher altitude. The reverse was observed at the lower altitudes; pollinators approached large-bullseye flowers often but rarely foraged. These differences are expected to affect fitness because foraging visits increased pollen export and receipt. Indeed, when natural variation in bullseye size was considered, it had a stronger effect on pollen receipt than other traits (flower size, display, or color). Plants with larger bullseyes tended to receive more pollen at the highest-altitude site, while those with smaller ones received more pollen at the lowest-altitude site. Results suggest that altitudinal changes in preference and behavior of the overall pollinator assemblage, but not differential preferences of pollinator taxonomic groups, could contribute to clinal variation in a UV floral trait for a generalist-pollinated plant.

Description: Understanding how biotic communities respond to landscape spatial structure is critically important for conservation management as natural habitats become increasingly fragmented. However, empirical studies of the effects of spatial structure on plant species richness have found inconsistent results, suggesting that more comprehensive approaches are needed. We asked how landscape structure affects total plant species richness and the richness of a guild of specialized plants in a multivariate context. We sampled herbaceous plant communities at 56 dolomite glades (insular, fire-adapted grasslands) across the Missouri Ozarks, USA, and used structural equation modeling (SEM) to analyze the relative importance of landscape structure, soil resource availability, and fire history for plant communities. We found that landscape spatial structure, defined as the area-weighted proximity of glade habitat surrounding study sites (proximity index), had a significant effect on total plant species richness, but only after we controlled for environmental covariates. Richness of specialist species, but not generalists, was positively related to landscape spatial structure. Our results highlight that local environmental filters must be considered to understand the influence of landscape structure on communities and that unique species guilds may respond differently to landscape structure than the community as a whole. These findings suggest that both local environment and landscape context should be considered when developing management strategies for species of conservation concern in fragmented habitats.

Description: Plant diversity enhances many ecosystem processes, including productivity, but these effects have been studied almost exclusively at the taxonomic scale of species. We explore the effect of intraspecific diversity on the productivity of a widespread and dominant grassland species using accessions collected from populations throughout its range. We found that increasing population/ecotype diversity of Pseudoroegneria spicata increased productivity to a similar degree as that reported for species diversity. However, we did not find evidence that overyielding was related to either resource depletion or to pathogenic soil fungi, two causes of overyielding in species-diverse communities. Instead, larger accessions overyielded at low diversity at the expense of smaller accessions, and small accessions overyielded through complementarity at all levels of diversity. Furthermore, overyielding was stronger for accessions from mesic environments, suggesting that local adaptation might predictably influence how plants respond to increases in diversity. This suggests that mass-based competition or other cryptic accession-specific processes had complex but important effects on overyielding. Our results indicate that the effects of diversity within a species can be substantial but that overyielding by intraspecifically diverse populations may not be through the same processes thought to cause overyielding in species diverse communities.

Description: Aboveground herbivores have strong effects on grassland nitrogen (N) cycling. They can accelerate or slow down soil net N mineralization depending on ecosystem productivity and grazing intensity. Yet, most studies only consider either ungulates or invertebrate herbivores, but not the combined effect of several functionally different vertebrate and invertebrate herbivore species or guilds. We assessed how a diverse herbivore community affects net N mineralization in subalpine grasslands. By using size-selective fences, we progressively excluded large, medium, and small mammals, as well as invertebrates from two vegetation types, and assessed how the exclosure types (ET) affected net N mineralization. The two vegetation types differed in long-term management (centuries), forage quality, and grazing history and intensity. To gain a more mechanistic understanding of how herbivores affect net N mineralization, we linked mineralization to soil abiotic (temperature; moisture; NO 3 − , NH 4 + , and total inorganic N concentrations/pools; C, N, P concentrations; pH; bulk density), soil biotic (microbial biomass; abundance of collembolans, mites, and nematodes) and plant (shoot and root biomass; consumption; plant C, N, and fiber content; plant N pool) properties. Net N mineralization differed between ET, but not between vegetation types. Thus, short-term changes in herbivore community composition and, therefore, in grazing intensity had a stronger effect on net N mineralization than long-term management and grazing history. We found highest N mineralization values when only invertebrates were present, suggesting that mammals had a negative effect on net N mineralization. Of the variables included in our analyses, only mite abundance and aboveground plant biomass explained variation in net N mineralization among ET. Abundances of both mites and leaf-sucking invertebrates were positively correlated with aboveground plant biomass, and biomass increased with progressive exclusion. The negative impact of mammals on net N mineralization may be related partially to (1) differences in the amount of plant material (litter) returned to the belowground subsystem, which induced a positive bottom-up effect on mite abundance, and (2) alterations in the amount and/or distribution of dung, urine, and food waste. Thus, our results clearly show that short-term alterations of the aboveground herbivore community can strongly impact nutrient cycling within ecosystems independent of long-term management and grazing history.

Description: Although we understand the ecological processes eliciting changes in plant community composition during secondary succession, we do not understand whether co-occurring changes in plant detritus shape saprotrophic microbial communities in soil. In this study, we investigated soil microbial composition and function across an old-field chronosequence ranging from 16 to 86 years following agricultural abandonment, as well as three forests representing potential late-successional ecosystems. Fungal and bacterial community composition was quantified from ribosomal DNA, and insight into the functional potential of the microbial community to decay plant litter was gained from shotgun metagenomics and extracellular enzyme assays. Accumulation of soil organic matter across the chronosequence exerted a positive and significant effect on fungal phylogenetic β-diversity and the activity of extracellular enzymes with lignocellulolytic activity. In addition, the increasing abundance of lignin-rich C 4 grasses was positively related to the composition of fungal genes with lignocellulolytic function, thereby linking plant community composition, litter biochemistry, and microbial community function. However, edaphic properties were the primary agent shaping bacterial communities, as bacterial β-diversity and variation in functional gene composition displayed a significant and positive relationship to soil pH across the chronosequence. The late-successional forests were compositionally distinct from the oldest old fields, indicating that substantial changes occur in soil microbial communities as old fields give way to forests. Taken together, our observations demonstrate that plants govern the turnover of soil fungal communities and functional characteristics during secondary succession, due to the continual input of detritus and differences in litter biochemistry among plant species.