Showing papers in "Oecologia in 2018"

Changes in tree resistance, recovery and resilience across three successive extreme droughts in the northeast Iberian Peninsula

Abstract: Understanding which variables affect forest resilience to extreme drought is key to predict future dynamics under ongoing climate change. In this study, we analyzed how tree resistance, recovery and resilience to drought have changed along three consecutive droughts and how they were affected by species, tree size, plot basal area (as a proxy for competition) and climate. We focused on the three most abundant pine species in the northeast Iberian Peninsula: Pinus halepensis, P. nigra and P. sylvestris during the three most extreme droughts recorded in the period 1951-2010 (occurred in 1986, 1994, and 2005-2006). We cored trees from permanent sample plots and used dendrochronological techniques to estimate resistance (ability to maintain growth level during drought), recovery (growth increase after drought) and resilience (capacity to recover pre-drought growth levels) in terms of tree stem basal area increment. Mixed-effects models were used to determine which tree- and plot-level variables were the main determinants of resistance, recovery and resilience, and to test for differences among the studied droughts. Larger trees were significantly less resistant and resilient. Plot basal area effects were only observed for resilience, with a negative impact only during the last drought. Resistance, recovery and resilience differed across the studied drought events, so that the studied populations became less resistant, less resilient and recovered worse during the last two droughts. This pattern suggests an increased vulnerability to drought after successive drought episodes.

Movement of pulsed resource subsidies from kelp forests to deep fjords

Abstract: Resource subsidies in the form of allochthonous primary production drive secondary production in many ecosystems, often sustaining diversity and overall productivity. Despite their importance in structuring marine communities, there is little understanding of how subsidies move through juxtaposed habitats and into recipient communities. We investigated the transport of detritus from kelp forests to a deep Arctic fjord (northern Norway). We quantified the seasonal abundance and size structure of kelp detritus in shallow subtidal (0‒12 m), deep subtidal (12‒85 m), and deep fjord (400‒450 m) habitats using a combination of camera surveys, dive observations, and detritus collections over 1 year. Detritus formed dense accumulations in habitats adjacent to kelp forests, and the timing of depositions coincided with the discrete loss of whole kelp blades during spring. We tracked these blades through the deep subtidal and into the deep fjord, and showed they act as a short-term resource pulse transported over several weeks. In deep subtidal regions, detritus consisted mostly of fragments and its depth distribution was similar across seasons (50% of total observations). Tagged pieces of detritus moved slowly out of kelp forests (displaced 4‒50 m (mean 11.8 m ± 8.5 SD) in 11‒17 days, based on minimum estimates from recovered pieces), and most (75%) variability in the rate of export was related to wave exposure and substrate. Tight resource coupling between kelp forests and deep fjords indicate that changes in kelp abundance would propagate through to deep fjord ecosystems, with likely consequences for the ecosystem functioning and services they provide.

Inferring the source of evaporated waters using stable H and O isotopes

Abstract: Stable isotope ratios of H and O are widely used to identify the source of water, e.g., in aquifers, river runoff, soils, plant xylem, and plant-based beverages. In situations where the sampled water is partially evaporated, its isotope values will have evolved along an evaporation line (EL) in δ2H/δ18O space, and back-correction along the EL to its intersection with a meteoric water line (MWL) has been used to estimate the source water’s isotope ratios. Here, we review the theory underlying isotopic estimation of source water for evaporated samples (iSWE). We note potential for bias from a commonly used regression-based approach for EL slope estimation and suggest that a model-based approach may be preferable if assumptions of the regression approach are not valid. We then introduce a mathematical framework that eliminates the need to explicitly estimate the EL–MWL intersection, simplifying iSWE analysis and facilitating more rigorous uncertainty estimation. We apply this approach to data from the US EPA's 2007 National Lakes Assessment. We find that data for most lakes are consistent with a water source similar to annual runoff, estimated from monthly precipitation and evaporation within the lake basin. Strong evidence for both summer- and winter-biased sources exists, however, with winter bias pervasive in most snow-prone regions. The new analytical framework should improve the rigor of iSWE in ecohydrology and related sciences, and our initial results from US lakes suggest that previous interpretations of lakes as unbiased isotope integrators may only be valid in certain climate regimes.

Climate variability decreases species richness and community stability in a temperate grassland

Abstract: Climate change involves modifications in both the mean and the variability of temperature and precipitation. According to global warming projections, both the magnitude and the frequency of extreme weather events are increasing, thereby increasing climate variability. The previous studies have reported that climate warming tends to decrease biodiversity and the temporal stability of community primary productivity (i.e., community stability), but the effects of the variability of temperature and precipitation on biodiversity, community stability, and their relationship have not been clearly explored. We used a long-term (from 1982 to 2014) field data set from a temperate grassland in northern China to explore the effects of the variability of mean temperature and total precipitation on species richness, community stability, and their relationship. Results showed that species richness promoted community stability through increases in asynchronous dynamics across species (i.e., species asynchrony). Both species richness and species asynchrony were positively associated with the residuals of community stability after controlling for its dependence on the variability of mean temperature and total precipitation. Furthermore, the variability of mean temperature reduced species richness, while the variability of total precipitation decreased species asynchrony and community stability. Overall, the present study revealed that species richness and species asynchrony promoted community stability, but increased climate variability may erode these positive effects and thereby threaten community stability.

Some like it hot: the physiological ecology of C4 plant evolution.

Abstract: The evolution of C4 photosynthesis requires an intermediate phase where photorespiratory glycine produced in the mesophyll cells must flow to the vascular sheath cells for metabolism by glycine decarboxylase. This glycine flux concentrates photorespired CO2 within the sheath cells, allowing it to be efficiently refixed by sheath Rubisco. A modest C4 biochemical cycle is then upregulated, possibly to support the refixation of photorespired ammonia in sheath cells, with subsequent increases in C4 metabolism providing incremental benefits until an optimized C4 pathway is established. ‘Why’ C4 photosynthesis evolved is largely explained by ancestral C3 species exploiting photorespiratory CO2 to improve carbon gain and thus enhance fitness. While photorespiration depresses C3 performance, it produces a resource (photorespired CO2) that can be exploited to build an evolutionary bridge to C4 photosynthesis. ‘Where’ C4 evolved is indicated by the habitat of species branching near C3-to-C4 transitions on phylogenetic trees. Consistent with the photorespiratory bridge hypothesis, transitional species show that the large majority of > 60 C4 lineages arose in hot, dry, and/or saline regions where photorespiratory potential is high. ‘When’ C4 evolved has been clarified by molecular clock analyses using phylogenetic data, coupled with isotopic signatures from fossils. Nearly all C4 lineages arose after 25 Ma when atmospheric CO2 levels had fallen to near current values. This reduction in CO2, coupled with persistent high temperature at low-to-mid-latitudes, met a precondition where photorespiration was elevated, thus facilitating the evolutionary selection pressure that led to C4 photosynthesis.

The role of ants in north temperate grasslands: a review

Abstract: Historic and current land-use changes have altered the landscape for grassland biota, with over 90% of grasslands and savannas converted to agriculture or some other use in north temperate regions. Reintegrating grasslands into agricultural landscapes can increase biodiversity while also providing valuable ecosystem services. In contrast to their well-known importance in tropical and subtropical ecosystems, the role of ants in temperate grasslands is often underappreciated. As consumers and ecosystem engineers, ants in temperate grasslands influence invertebrate, plant, and soil microbial diversity and potentially alter grassland productivity. As common and numerically dominant invertebrates in grasslands, ants can also serve as important indicator species to monitor conservation and management practices. Drawing on examples largely from mesic, north temperate studies, and from other temperate regions where necessary, we review the roles of ants as consumers and ecosystem engineers in grasslands. We also identify five avenues for future research to improve our understanding of the roles of ants in grasslands. This includes identifying how grassland fragmentation may influence ant community assembly, quantifying how ant communities impact ecosystem functions and soil processes, and understanding how ant communities and their associated interactions are impacted by climate change. In synthesizing the role of ants in temperate grasslands and identifying knowledge gaps, we hope this and future work will help inform how land managers maximize grassland conservation value while increasing multiple ecosystem services and minimizing disservices.

Drought neutralises plant–soil feedback of two mesic grassland forbs

Abstract: Plant–soil feedbacks (PSFs) describe the effect of a plant species on soil properties, which affect the performance of future generations. Here we test the hypothesis that drought alters PSFs by reducing plant–microbe associations and nutrient uptake. We chose two grassland forb species, previously shown to respond differently to soil conditioning and drought, to test our hypothesis. We conditioned unsterilised grassland soil with one generation of each species, and left a third soil unconditioned. We grew a second generation consisting of each combination of plant species, soil, and drought in a full factorial design, and measured soil microbial community and nutrient availability. Scabiosa columbaria displayed negative PSF (smaller plants) under non-droughted conditions, but neutral under drought, suggesting that drought disrupts plant–soil interactions and can advantage the plant. Photosynthetic efficiency of S. columbaria was reduced under drought, but recovered on rewetting regardless of soil conditioning, indicating that PSFs do not impede resilience of this species. Sanguisorba minor showed positive PSFs (larger plants), probably due to an increase in soil N in conspecific soil, but neutral PSF under drought. PSF neutralisation appeared to occur through drought-induced change in the soil microbial community for this species. When S. minor was planted in conspecific soil, photosynthetic efficiency declined to almost zero, with no recovery following rewetting. We attributed this to increased demand for water through higher demand for nutrients with positive PSF. Here we show that drought neutralises PSFs of two grassland forbs, which could have implications for plant communities under climate change.

Belowground competition drives invasive plant impact on native species regardless of nitrogen availability

Abstract: Plant invasions and eutrophication are pervasive drivers of global change that cause biodiversity loss. Yet, how invasive plant impacts on native species, and the mechanisms underpinning these impacts, vary in relation to increasing nitrogen (N) availability remains unclear. Competition is often invoked as a likely mechanism, but the relative importance of the above and belowground components of this is poorly understood, particularly under differing levels of N availability. To help resolve these issues, we quantified the impact of a globally invasive grass species, Agrostis capillaris, on two co-occurring native New Zealand grasses, and vice versa. We explicitly separated above- and belowground interactions amongst these species experimentally and incorporated an N addition treatment. We found that competition with the invader had large negative impacts on native species growth (biomass decreased by half), resource capture (total N content decreased by up to 75%) and even nutrient stoichiometry (native species tissue C:N ratios increased). Surprisingly, these impacts were driven directly and indirectly by belowground competition, regardless of N availability. Higher root biomass likely enhanced the invasive grass’s competitive superiority belowground, indicating that root traits may be useful tools for understanding invasive plant impacts. Our study shows that belowground competition can be more important in driving invasive plant impacts than aboveground competition in both low and high fertility ecosystems, including those experiencing N enrichment due to global change. This can help to improve predictions of how two key drivers of global change, plant species invasions and eutrophication, impact native species diversity.

Seagrass collapse due to synergistic stressors is not anticipated by phenological changes

Abstract: Seagrasses are globally declining and often their loss is due to synergies among stressors. We investigated the interactive effects of eutrophication and burial on the Mediterranean seagrass, Posidonia oceanica. A field experiment was conducted to estimate whether shoot survival depends on the interactive effects of three levels of intensity of both stressors and to identify early changes in plants (i.e., morphological, physiological and biochemical, and expression of stress-related genes) that may serve to detect signals of imminent shoot density collapse. Sediment burial and nutrient enrichment produced interactive effects on P. oceanica shoot survival, as high nutrient levels had the potential to accelerate the regression of the seagrass exposed to high burial (HB). After 11 weeks, HB in combination with either high or medium nutrient enrichment caused a shoot loss of about 60%. Changes in morphology were poor predictors of the seagrass decline. Likewise, few biochemical variables were associated with P. oceanica survival (the phenolics, ORAC and leaf δ34S). In contrast, the expression of target genes had the highest correlation with plant survival: photosynthetic genes (ATPa, psbD and psbA) were up-regulated in response to high burial, while carbon metabolism genes (CA-chl, PGK and GADPH) were down-regulated. Therefore, die-offs due to high sedimentation rate in eutrophic areas can only be anticipated by altered expression of stress-related genes that may warn the imminent seagrass collapse. Management of local stressors, such as nutrient pollution, may enhance seagrass resilience in the face of the intensification of extreme climate events, such as floods.

Consequences of interspecific variation in defenses and herbivore host choice for the ecology and evolution of Inga, a speciose rainforest tree

Abstract: We summarize work on a speciose Neotropical tree genus, Inga (Fabaceae), examining how interspecific variation in anti-herbivore defenses may have evolved, how defenses shape host choice by herbivores and how they might regulate community composition and influence species radiations. Defenses of expanding leaves include secondary metabolites, extrafloral nectaries, rapid leaf expansion, trichomes, and synchrony and timing of leaf production. These six classes of defenses are orthogonal, supporting independent evolutionary trajectories. Moreover, only trichomes show a phylogenetic signature, suggesting evolutionary lability in nearly all defenses. The interspecific diversity in secondary metabolite profiles does not arise from the evolution of novel compounds, but from novel combinations of common compounds, presumably due to changes in gene regulation. Herbivore host choice is determined by plant defensive traits, not host phylogeny. Neighboring plants escape each other's pests if their defenses differ enough, thereby enforcing the high local diversity typical of tropical forests. Related herbivores feed on hosts with similar defenses, implying that there are phylogenetic constraints placed on the herbivore traits that are associated with host use. Divergence in defensive traits among Inga appears to be driven by herbivore pressure. However, the lack of congruence between herbivore and host phylogeny suggests that herbivores are tracking defenses, choosing hosts based on traits for which they already have adaptations. There is, therefore, an asymmetry in the host-herbivore evolutionary arms race.

Global patterns in the metacommunity structuring of lake macrophytes: regional variations and driving factors.

Abstract: We studied community-environment relationships of lake macrophytes at two metacommunity scales using data from 16 regions across the world. More specifically, we examined (a) whether the lake macrophyte communities respond similar to key local environmental factors, major climate variables and lake spatial locations in each of the regions (i.e., within-region approach) and (b) how well can explained variability in the community-environment relationships across multiple lake macrophyte metacommunities be accounted for by elevation range, spatial extent, latitude, longitude, and age of the oldest lake within each metacommunity (i.e., across-region approach). In the within-region approach, we employed partial redundancy analyses together with variation partitioning to investigate the relative importance of local variables, climate variables, and spatial location on lake macrophytes among the study regions. In the across-region approach, we used adjusted R2 values of the variation partitioning to model the community-environment relationships across multiple metacommunities using linear regression and commonality analysis. We found that niche filtering related to local lake-level environmental conditions was the dominant force structuring macrophytes within metacommunities. However, our results also revealed that elevation range associated with climate (increasing temperature amplitude affecting macrophytes) and spatial location (likely due to dispersal limitation) was important for macrophytes based on the findings of the across-metacommunities analysis. These findings suggest that different determinants influence macrophyte metacommunities within different regions, thus showing context dependency. Moreover, our study emphasized that the use of a single metacommunity scale gives incomplete information on the environmental features explaining variation in macrophyte communities.

Diversity–biomass relationship across forest layers: implications for niche complementarity and selection effects

Abstract: Forest stratification plays a crucial role in light interception and plant photosynthetic activities. However, despite the increased number of studies on biodiversity–ecosystem function, we still lack information on how stratification in tropical forests modulates biodiversity effects. Moreover, there is less investigation and argument on the role of species and functional traits in forest layers. Here, we analysed from a perspective of forest layer (sub-canopy, canopy and emergent species layers), the relationship between diversity and aboveground biomass (AGB), focusing on functional diversity and dominance, and underlying mechanisms such as niche complementarity and selection. The sub-canopy layer had the highest species richness and diversity, while the emergent layer had the highest AGB. Species richness–AGB relationship was positive for each forest layer, but stronger for sub-canopy layer than for canopy and emergent layers. Total AGB was strongly correlated with functional diversity, leaf and wood traits of species in the sub-canopy and canopy layers. This suggests that sub-canopy and canopy species are major drivers of stand diversity–AGB relationship, and that resource filtering by canopy or emergent trees may not reduce the strength of diversity–AGB relationship in the sub-canopy layer. We argue that complementary resource use by sub-canopy species that supports niche complementarity, is a key mechanism driving AGB in natural forests. Selection effects are most evident in emergent species and niche complementarity effects for sub-canopy and canopy species, supporting arguments that AGB is affected by sub-canopy species’ efficient use of limited resources despite competition from emergent species.

Terrestrial–aquatic trophic linkages support fish production in a tropical oligotrophic river

Abstract: Despite low in situ primary productivity, tropical oligotrophic rivers support highly diverse fish assemblages and productive fisheries. This raises the question, what energy sources support fish production in these ecosystems? We sampled fish and food resources in the floodplain of a nearly pristine, large, oligotrophic river in western Amazonia. We combined data from stomach contents and stable isotopes to test the hypothesis that floodplain forests sustain fisheries in tropical oligotrophic rivers. Analysis of stomach contents from > 800 specimens of 12 omnivorous fish species demonstrated that during the annual flood, forest plant matter dominated diets. Yet, our isotope mixing models estimated that arthropods from the forest canopy made a greater proportional contribution to fish biomass. Most of these arthropods are entirely terrestrial and, therefore, serve as trophic links between forests and fishes. Our results suggest that forest vegetation, particularly fruits, may provide much of the energy supporting metabolism and arthropods contribute significant amounts of protein for somatic growth. Moreover, the importance of terrestrial arthropods in support of fish biomass in oligotrophic rivers depends on interactions between riparian vegetation, terrestrial arthropods and flood pulse dynamics affecting accessibility of arthropods to fishes. The apparent paradox of high fish diversity in an oligotrophic river with low primary productivity may be explained, at least partially, by dynamic terrestrial–aquatic trophic linkages. This study further emphasizes the importance of seasonally flooded forests for sustaining fisheries in the Amazon.

DNA metabarcoding of nestling feces reveals provisioning of aquatic prey and resource partitioning among Neotropical migratory songbirds in a riparian habitat.

Abstract: Riparian habitats are characterized by substantial flows of emergent aquatic insects that cross the stream-forest interface and provide an important source of prey for insectivorous birds. The increased availability of prey arising from aquatic subsidies attracts high densities of Neotropical migratory songbirds that are thought to exploit emergent aquatic insects as a nestling food resource; however, the prey preferences and diets of birds in these communities are only broadly understood. In this study, we utilized DNA metabarcoding to investigate the extent to which three syntopic species of migratory songbirds—Acadian Flycatcher, Louisiana Waterthrush, and Wood Thrush—breeding in Appalachian riparian habitats (Pennsylvania, USA) exploit and partition aquatic prey subsidies as a nestling food resource. Despite substantial differences in adult foraging strategies, nearly every nestling in this study consumed aquatic taxa, suggesting that aquatic subsidies are an important prey resource for Neotropical migrants nesting in riparian habitats. While our results revealed significant interspecific dietary niche divergence, the diets of Acadian Flycatcher and Wood Thrush nestlings were strikingly similar and exhibited significantly more overlap than expected. These results suggest that the dietary niches of Neotropical migrants with divergent foraging strategies may converge due to the opportunistic provisioning of non-limiting prey resources in riparian habitats. In addition to providing the first application of DNA metabarcoding to investigate diet in a community of Neotropical migrants, this study emphasizes the importance of aquatic subsidies in supporting breeding songbirds and improves our understanding of how anthropogenic disturbances to riparian habitats may negatively impact long-term avian conservation.

Phytophagy of omnivorous predator Macrolophus pygmaeus affects performance of herbivores through induced plant defences

Abstract: Plants possess various inducible defences that result in synthesis of specialized metabolites in response to herbivory, which can interfere with the performance of herbivores of the same and other species. Much less is known of the effects of plant feeding by omnivores. We found that previous feeding of the omnivorous predator Macrolophus pygmaeus on sweet pepper plants significantly reduced reproduction of the two-spotted spider mite Tetranychus urticae and western flower thrips Frankliniella occidentalis on the same plants, also on leaves that had not been exposed to the omnivore. In contrast, no effect was found on the reproduction of the green peach aphid Myzus persicae. Juvenile survival and developmental time of T. urticae and M. persicae, and larval survival of F. occidentalis were not affected by plant feeding by M. pygmaeus. Larvae of F. occidentalis feeding on leaves previously exposed to M. pygmaeus required longer to develop into adults. Defence-related plant hormones were produced locally and systemically after exposure to M. pygmaeus. The concentrations of 12-oxo-phytodienoic acid and jasmonic acid–isoleucine in the attacked leaves were significantly higher than in the corresponding leaves on the uninfested plants, and jasmonic acid concentrations showed the same trend, suggesting that jasmonic-acid-related defence pathways were activated. In contrast, similar concentrations of salicylic acid were found in the attacked leaves of M. pygmaeus-infested plants and uninfested plants. Our results show that plant feeding by omnivorous predators decreases the performance of herbivores, suggesting that it induces plant defences.

Phytochemical variation in treetops: causes and consequences for tree-insect herbivore interactions.

Abstract: The interaction of plants and their herbivorous opponents has shaped the evolution of an intricate network of defences and counter-defences for millions of years. The result is an astounding diversity of phytochemicals and plant strategies to fight and survive. Trees are specifically challenged to resist the plethora of abiotic and biotic stresses due to their dimension and longevity. Here, we review the recent literature on the consequences of phytochemical variation in trees on insect-tree-herbivore interactions. We discuss the importance of genotypic and phenotypic variation in tree defence against insects and suggest some molecular mechanisms that might bring about phytochemical diversity in crowns of individual trees.

Mycorrhizal associations and the spatial structure of an old-growth forest community.

Abstract: Plant-soil feedbacks are known to play a central role in species co-existence, but conceptual frameworks for predicting their magnitude and direction are lacking. We ask whether co-occurring trees that associate with different types of mycorrhizal fungi, which are hypothesized to differ in terms of nutrient use and plant-soil feedbacks, differ in sapling establishment densities and probability of co-occurrence. Given that ectomycorrhizal (ECM) trees typically have fungal structures that protect roots from pathogens whereas arbuscular mycorrhizal (AM) trees do not, we hypothesized that ECM saplings would be clustered around ECM trees, while AM saplings would be suppressed near AM trees. Most previous studies have focused on seedlings, but here we examine whether the spatial signal is evident in later life stages. We measured the spatial associations of ~ 28,000 trees using point pattern analysis in a 25-ha old-growth forest where ECM trees comprised 72% of total basal area and 42% of the total stems, while AM trees comprised the remainder. Supporting our hypothesis, AM saplings were more inhibited by AM trees, while ECM saplings were more clustered around ECM trees. The spatial patterns of AM and ECM trees on saplings of the alternate mycorrhizal type were inhibited. To the extent that similar types of feedbacks occur for other AM and ECM trees, our results suggest that fundamental differences in the nature of local-scale biotic interactions between trees and their fungal symbionts may influence forest community assembly and ecosystem dynamics.

Anthropogenic fragmentation of landscapes: mechanisms for eroding the specificity of plant–herbivore interactions

Abstract: Reduced ecological specialization is an emerging, general pattern of ecological networks in fragmented landscapes. In plant–herbivore interactions, reductions in dietary specialization of herbivore communities are consistently associated with fragmented landscapes, but the causes remain poorly understood. We propose several hypothetical bottom–up and top–down mechanisms that may reduce the specificity of plant–herbivore interactions. These include empirically plausible applications and extensions of theory based on reduced habitat patch size and isolation (considered jointly), and habitat edge effects. Bottom–up effects in small, isolated habitat patches may limit availability of suitable hostplants, a constraint that increases with dietary specialization. Poor hostplant quality due to inbreeding in such fragments may especially disadvantage dietary specialist herbivores even when their hostplants are present. Size and isolation of habitat patches may change patterns of predation of herbivores, but whether such putative changes are associated with herbivore dietary specialization should depend on the mobility, size, and diet breadth of predators. Bottom–up edge effects may favor dietary generalist herbivores, yet top–down edge effects may favor dietary specialists owing to reduced predation. An increasingly supported edge effect is trophic ricochets generated by large grazers/browsers, which remove key hostplant species of specialist herbivores. We present empirical evidence that greater deer browsing in small forest fragments disproportionately reduces specialist abundances in lepidopteran assemblages in northeastern USA. Despite indirect evidence for these mechanisms, they have received scant direct testing with experimental approaches at a landscape scale. Identifying their relative contributions to reduced specificity of plant–herbivore interactions in fragmented landscapes is an important research goal.

Heated communities: large inter- and intraspecific variation in heat tolerance across trophic levels of a soil arthropod community.

Abstract: Temperature extremes are predicted to increase in frequency, intensity and duration under global warming and are believed to significantly affect community composition and functioning. However, the effect of extreme climatic events on communities remains difficult to predict, especially because species can show dissimilar responses to abiotic changes, which may affect the outcome of species interactions. To anticipate community responses we need knowledge on within and among species variation in stress tolerance. We exposed a soil arthropod community to experimental heat waves in the field and measured heat tolerance of species of different trophic levels from heated and control plots. We measured the critical thermal maximum (CTmax) of individuals to estimate inter- and intraspecific variation in heat tolerance in this community, and how this was affected by experimental heat waves. We found interspecific variation in heat tolerance, with the most abundant prey species, the springtail Isotoma riparia, being more sensitive to high temperatures than its predators (various spider species). Moreover, intraspecific variation in CTmax was substantial, suggesting that individuals within a single species were unequally affected by heat extremes. However, heat tolerance of species did not increase after being exposed to an experimental heat wave. We conclude that interspecific variation in tolerance traits potentially causes trophic mismatches during extreme events, but that intraspecific variation could lessen these effects by enabling partial survival of populations. Therefore, ecophysiological traits can provide a better understanding of abiotic effects on communities, not only within taxonomic or functional groups, but also when comparing different trophic levels.

Effectiveness of predator satiation in masting oaks is negatively affected by conspecific density

Abstract: Variation in seed availability shapes plant communities, and is strongly affected by seed predation. In some plant species, temporal variation in seed production is especially high and synchronized over large areas, which is called ‘mast seeding’. One selective advantage of this phenomenon is predator satiation which posits that masting helps plants escape seed predation through starvation of predators in lean years, and satiation in mast years. However, even though seed predation can be predicted to have a strong spatial component and depend on plant densities, whether the effectiveness of predator satiation in masting plants changes according to the Janzen-Connell effect has been barely investigated. We studied, over an 8-year period, the seed production, the spatiotemporal patters of weevil seed predation, and the abundance of adult weevils in a holm oak (Quercus ilex) population that consists of trees interspersed at patches covering a continuum of conspecific density. Isolated oaks effectively satiate predators, but this is trumped by increasing conspecific plant density. Lack of predator satiation in trees growing in dense patches was caused by re-distribution of insects among plants that likely attenuated them against food shortage in lean years, and changed the type of weevil functional response from type II in isolated trees to type III in trees growing in dense patches. This study provides the first empirical evaluation of the notion that masting and predator satiation should be more important in populations that start to dominate their communities, and is consistent with the observation that masting is less frequent and less intense in diverse forests.

Plants learn and remember: lets get used to it

Abstract: In one of the most beloved comic strip of all times, the creator of Charlie Brown and Snoopy said it all. At the most basic level, this “getting used to it”—a decrease in response to a stimulus after repeated exposure—is what behavioural habituation is about. And, it is an extremely important (adaptive) aspect of life; thanks to it, organisms learn to pay attention to stimuli that are truly meaningful in their environment while ignoring those that have proven irrelevant and innocuous. Despite its simplicity, the concept of habituation seems to have always stirred great debate. For example, the earliest descriptions of what we would now call habituation came from studies conducted by Pfeffer using the sensitive plant Mimosa pudica in 1873 and the Peckhams using spiders in 1887; remarkably, no real agreement on the use of the word ‘habituation’ was to be found until (almost) a whole century later (see review by Christoffersen 1997). As Christoffersen (1997) pointed out, the terminology debate regarding habituation could not be settled until the 1960s, because, only then, the number of studies that investigated the behavioural phenomenon had grown sufficiently large, so that Thompson and Spencer (1966) could establish an operational definition. Thompson and Spencer’s definition of habituation was based on a list of nine behavioural characteristics, which have since been confirmed by most of the cases investigated and still constitutes a useful working framework today. While remaining largely unchallenged since its formulation, the list of characteristics was nonetheless expanded by Groves and Thompson in (1970), revised by Christoffersen in 1997 and updated again by Rankin et al. in 2009, for some examples. Hence, while providing a convenient primer for the study of habituation in a wide range of species and paradigms, this framework may not be taken as a set-in-stones rule manual but rather as an evolving guide. From an ecological perspective, such a framework would be most useful and its application most interesting when informing the study of habituation as the process enabling an organism to better respond and adapt to its constantly changing environment. This is how it was applied in the Gagliano et al. (2014) study, whose aim was to investigate the learning process of the Mimosa plant within the context of a trade-off between predation risk and foraging for light, a context that is ecologically relevant to this plant. In considering these issues, Beigler (2017)’s commentary points out that Gagliano et al. (2014) correctly tested the most relevant behavioural characteristics of habituation. Beigler expresses concerns over the omission of characteristics Gagliano et al. (2014) did not test, and over the fact that other tests not mentioned in the list of criteria (as per Rankin et al. 2009) were included instead. As it is perhaps true of most ecological studies investigating habituation, Gagliano et al.’s paper never aimed nor claimed to be a test of all the criteria characterising the process of habituation as defined by Rankin et al. (2009). Besides, several of the characteristics not (expressly) examined by Gagliano et al. had been investigated in earlier studies of this plant (reviewed by Sanberg 1976 and more recently, Abramson and Chicas-Mosier 2016). What Gagliano et al.’s study did examine were the most relevant behavioural characteristics of habituation in the context of a broader ecological question—whether Mimosa’s ability to learn through the habituation of its defensive leaf-folding reflex was mediated by environmental circumstances such as lowand high-light Communicated by Richard Karban.

Predator-prey interaction reveals local effects of high-altitude insect migration

Abstract: High-altitude nocturnal insect migrations are ubiquitous and represent significant pulses of biomass, which impact large areas and multiple trophic levels, yet are difficult to study and poorly understood. Predation on migratory insects by high-flying bats provides potential for investigating flows of migratory insects across a landscape. Brazilian free-tailed bats, Tadarida brasiliensis, provide valuable ecosystem services by consuming migratory pests, and research suggests migratory insects are an important resource to bats in autumn. We sequenced insect DNA from bat feces collected during the 2010–2012 autumn migrations of insects over southern Texas, and tested the utility of predator–prey interactions for monitoring migratory insect populations by asking: 1) how extensively do bats consume migratory insects during autumn? (2) does the prey community reflect known drivers of insect migrations, e.g. cold fronts? and (3) are migratory insects increasingly important to bats when local food resources decline in autumn? Bats consumed at least 21 species of migratory insects and 44 species of agricultural pests. Prey community richness increased with cold front passage. Bats consumed migratory moths over the entire autumn season, and the proportion of migratory moths in the bat diet increased over the course of the autumn season in all 3 years. This study confirms extensive consumption of migratory insects by bats, links patterns in prey communities to mechanisms driving insect migration, and documents a novel approach to tracking patterns of migratory insect movement. As an important resource for T. brasiliensis in autumn, migratory insects provide stabilizing effects to the local animal community.

Nitrogen enrichment in host plants increases the mortality of common Lepidoptera species

Abstract: The recent decline of Lepidoptera species strongly correlates with the increasing intensification of agriculture in Western and Central Europe. However, the effects of changed host-plant quality through agricultural fertilization on this insect group remain largely unexplored. For this reason, we tested the response of six common butterfly and moth species to host-plant fertilization using fertilizer quantities usually applied in agriculture. The larvae of the study species Coenonympha pamphilus, Lycaena phlaeas, Lycaena tityrus, Pararge aegeria, Rivula sericealis and Timandra comae were distributed according to a split-brood design to three host-plant treatments comprising one control treatment without fertilization and two fertilization treatments with an input of 150 and 300 kg N ha−1 year−1, respectively. In L. tityrus, we used two additional fertilization treatments with an input of 30 and 90 kg N ha−1 year−1, respectively. Fertilization increased the nitrogen concentration of both host-plant species, Rumex acetosella and Poa pratensis, and decreased the survival of larvae in all six Lepidoptera species by at least one-third, without clear differences between sorrel- and grass-feeding species. The declining survival rate in all species contradicts the well-accepted nitrogen-limitation hypothesis, which predicts a positive response in species performance to dietary nitrogen content. In contrast, this study presents the first evidence that current fertilization quantities in agriculture exceed the physiological tolerance of common Lepidoptera species. Our results suggest that (1) the negative effect of plant fertilization on Lepidoptera has previously been underestimated and (2) that it contributes to the range-wide decline of Lepidoptera.

Fear, foraging and olfaction: how mesopredators avoid costly interactions with apex predators

Abstract: Where direct killing is rare and niche overlap low, sympatric carnivores may appear to coexist without conflict. Interference interactions, harassment and injury from larger carnivores may still pose a risk to smaller mesopredators. Foraging theory suggests that animals should adjust their behaviour accordingly to optimise foraging efficiency and overall fitness, trading off harvest rate with costs to fitness. The foraging behaviour of red foxes, Vulpes vulpes, was studied with automated cameras and a repeated measures giving-up density (GUD) experiment where olfactory risk cues were manipulated. In Plitvice Lakes National Park, Croatia, red foxes increased GUDs by 34% and quitting harvest rates by 29% in response to wolf urine. In addition to leaving more food behind, foxes also responded to wolf urine by spending less time visiting food patches each day and altering their behaviour in order to compensate for the increased risk when foraging from patches. Thus, red foxes utilised olfaction to assess risk and experienced foraging costs due to the presence of a cue from gray wolves, Canis lupus. This study identifies behavioural mechanisms which may enable competing predators to coexist, and highlights the potential for additional ecosystem service pathways arising from the behaviour of large carnivores. Given the vulnerability of large carnivores to anthropogenic disturbance, a growing human population and intensifying resource consumption, it becomes increasingly important to understand ecological processes so that land can be managed appropriately.

Plant physical and chemical defence variation along elevation gradients: a functional trait-based approach

Abstract: Predicting variation in plant functional traits related to anti-herbivore defences remains a major challenge in ecological research, considering that multiple traits have evolved in response to both abiotic and biotic conditions. Therefore, understanding variation in plant anti-herbivore defence traits requires studying their expression along steep environmental gradients, such as along elevation, where multiple biotic and abiotic factors co-vary. We expand on plant defence theory and propose a novel conceptual framework to address the sources of variations of plant resistance traits at the community level. We analysed elevation patterns of within-community trait dissimilarity using the RaoQ index, and the community-weighted-mean (CWM) index, on several plant functional traits: plant height, specific leaf area (SLA), leaf-dry-matter-content (LDMC), silicium content, presence of trichomes, carbon-to-nitrogen ratio (CN) and total secondary metabolite richness. We found that at high elevation, where harsh environmental conditions persist, community functional convergence is dictated by traits relating to plant growth (plant height and SLA), while divergence arises for traits relating resource-use (LDMC). At low elevation, where greater biotic pressure occurs, we found a combination of random (plant height), convergence (metabolite richness) and divergence patterns (silicium content). This framework thus combines community assembly rules of ecological filtering and niche partition with plant defence hypotheses to unravel the relationship between environmental variations, biotic pressure and the average phenotype of plants within a community.

The impact of sea ice conditions on breeding decisions is modulated by body condition in an arctic partial capital breeder

Abstract: Determining how environmental conditions interact with individual intrinsic properties is important for unravelling the underlying mechanisms that drive variation in reproductive decisions among migratory species. We investigated the influence of sea ice conditions and body condition at arrival on the breeding propensity, i.e. the decision to reproduce or not within a single breeding season, and timing of laying in migrating common eiders (Somateria mollissima) breeding in the Arctic. Using Radarsat satellite images acquired from 2002 to 2013, we estimated the proportion of open water in the intertidal zone in early summer to track the availability of potential foraging areas for pre-breeding females. Timing of ice-breakup varied by up to 20 days across years and showed strong relationship with both breeding propensity and the timing of laying of eiders: fewer pre-breeding individuals were resighted nesting in the colony and laying was also delayed in years with late ice-breakup. Interestingly, the effect of sea ice dynamics on reproduction was modulated by the state of individuals at arrival on the breeding grounds: females arriving in low condition were more affected by a late ice-breakup. Open water accessibility in early summer, a likely proxy of food availability, is thus crucial for reproductive decisions in a (partial) capital breeder. Our predictive capacity in determining how Arctic-breeding seabirds respond to changes in environmental conditions will require incorporating such cross-seasonal cumulative effects.

Assessing seasonal changes in animal diets with stable-isotope analysis of amino acids: a migratory boreal songbird switches diet over its annual cycle.

Abstract: Tools to study seasonal changes in animal diets are needed to address a wide range of ecological questions. This is especially true of migratory animals that experience distinct environments where diets may be substantially different. However, tracking diets of individuals that move vast distances has proven difficult. Compound-specific isotope analysis has emerged as a valuable tool to study diets but has been little used to study dietary changes of migratory animals. Using this technique, we quantify seasonal variation in the annual diet of a migratory songbird (gray-cheeked thrush, Catharus minimus) and test the hypothesis that migrants change their diet in response to the energetic requirements of different periods of the annual cycle. By measuring δ13C and δ15N values of amino acids from feathers grown on the breeding grounds, blood formed during migration and claw grown on the wintering grounds, we found that migration is associated with greater consumption of fruit, compared to the breeding or wintering periods. This was confirmed by the lower trophic position of blood compared to feather and claw, by a decrease in the δ15N value of the source amino acid phenylalanine in blood as a function of days of stopover, and by the positive correlation between δ15N and δ13C values of phenylalanine in blood, and not in feather or claw. This study illustrates how isotopic analysis of amino acids can contribute to understand food webs, seasonal dietary changes and metabolic routing of nutrients in migratory animals.

Safe sites of pollen placement: a conflict of interest between plants and bees?

Abstract: Plant stigmas and bee pollinators are competitors for pollen. Pollen placed on a pollinator’s body can be picked up by conspecific stigmas or it can be collected by the pollinator as food. Hypothetically, one solution is for pollen to be placed on ‘safe sites’ on the pollinator’s body, sites where the pollinator cannot easily remove it, leaving the pollen for stigmas. We compared 14 sites on the bumblebee body in terms of the ability of the bee to groom off fluorescent power, a dust that may be thought of as analogous to pollen. The safest sites were along the midline of the dorsal thorax, the dorsal abdomen, and the ventral abdomen. Next, we counted how much pollen is borne on the 14 sites by bees visiting one nectariferous and three nectarless Pedicularis species. In the four species, only 7, 26, 28, and 30% of pollen found on the bees were on safe sites. Finally, we observed that the 14 sites were contacted by stigmas of the four Pedicularis species; none of the most contacted sites were safe sites. Across all four Pedicularis species, pollen is mainly positioned on sites of the bee body that were beneficial for both the plant and the bee, not on sites detrimental to either of them. Our analysis showed that the conflict of interest between flowers and bees can be solved by cooperation. Pedicularis pollen is placed where it strengthens the mutualism between plants and pollinators.

Small mammal responses to Amazonian forest islands are modulated by their forest dependence.

Abstract: Hydroelectric dams have induced widespread loss, fragmentation and degradation of terrestrial habitats in lowland tropical forests. Yet their ecological impacts have been widely neglected, particularly in developing countries, which are currently earmarked for exponential hydropower development. Here we assess small mammal assemblage responses to Amazonian forest habitat insularization induced by the 28-year-old Balbina Hydroelectric Dam. We sampled small mammals on 25 forest islands (0.83–1466 ha) and four continuous forest sites in the mainland to assess the overall community structure and species-specific responses to forest insularization. We classified all species according to their degree of forest-dependency using a multi-scale approach, considering landscape, patch and local habitat characteristics. Based on 65,520 trap-nights, we recorded 884 individuals of at least 22 small mammal species. Species richness was best predicted by island area and isolation, with small islands ( 200 ha; 10.8 ± 1.3 species) and continuous forest sites (∞ ha; 12.5 ± 2.5 species) exhibited similarly high species richness. Forest-dependent species showed higher local extinction rates and were often either absent or persisted at low abundances on small islands, where non-forest-dependent species became hyper-abundant. Species capacity to use non-forest habitat matrices appears to dictate small mammal success in small isolated islands. We suggest that ecosystem functioning may be highly disrupted on small islands, which account for 62.7% of all 3546 islands in the Balbina Reservoir.

Immune function and blood parasite infections impact stopover ecology in passerine birds.

Abstract: Stopovers play a crucial role for the success of migrating animals and are key to optimal migration theory. Variation in refuelling rates, stopover duration and departure decisions among individuals has been related to several external factors. The physiological mechanisms shaping stopover ecology are, however, less well understood. Here, we explore how immune function and blood parasite infections relate to several aspects of stopover behaviour in autumn migrating short- and long-distance migrating songbirds. We blood sampled individuals of six species and used an automated radio-telemetry system in the stopover area to subsequently quantify stopover duration, ‘bush-level’ activity patterns (~ 0.1–30 m), landscape movements (~ 30–6000 m), departure direction and departure time. We show that complement activity, the acute phase protein haptoglobin and blood parasite infections were related to prolonged stopover duration. Complement activity (i.e., lysis) and total immunoglobulins were negatively correlated with bush-level activity patterns. The differences partly depended on whether birds were long-distance or short-distance migrants. Birds infected with avian malaria-like parasites showed longer landscape movements during the stopover than uninfected individuals, and birds with double blood parasite infections departed more than 2.5 h later after sunset/sunrise suggesting shorter flight bouts. We conclude that variation in baseline immune function and blood parasite infection status affects stopover ecology and helps explain individual variation in stopover behaviour. These differences affect overall migration speed, and thus can have significant impact on migration success and induce carry-over effects on other annual-cycle stages. Immune function and blood parasites should, therefore, be considered as important factors when applying optimal migration theory.