Air temperature more than drought duration affects litter decomposition under flow intermittency.

Stream intermittency - periodic sequences of water flow cessation and resumption - occurs throughout the year, across seasons. Even though temperature is a known regulator of litter decomposition in both terrestrial and aquatic environments, comparative experiments on drought durations at distinct temperatures on microbial-mediated decomposition in streams experiencing intermittency are still lacking. Here, three drought temperatures (5, 15 and 25 °C) and two durations (short: 2.5 weeks; long: 5 weeks) were applied in a microcosm study to oak leaf discs colonized in a reference stream; mass loss and associated microbial parameters (fungal biomass, microbial activity, and sporulation rates) were evaluated following re-submersion for 2 weeks. Higher mass loss was found at 15 °C than 25 °C. A prolongation of the drought exposure period had no effect on mass loss, suggesting an early (≤ 2.5 weeks) inhibitor effect of drought on microbial-mediated leaf degradation. Fungal biomass was highest at 25 °C following a short drought, and decreased with a longer drought period at both 15 °C and 25 °C. Microbial activity was not affected by either drought duration or temperature. Sporulation rates and fungal diversity were significantly reduced by the longer drought period; in the short treatment, maximum values were found at 15 °C. In contrast to longer droughts, aquatic fungal communities during short dry periods seem to invest in energetically-expensive physiological responses to desiccation (e.g., ergosterol production) promoting biomass accrual at the expense of mass loss and reproductive output. Under more severe desiccation (higher duration and temperature), the lower diversity of fungal communities seem to result in negative legacy effects for fungal growth and reproductive capacity after flow resumption. These results suggest that native riparian vegetation, through its ability to regulate temperature in streams, may be critical in protecting freshwaters from intensified severity of drought periods in streams experiencing intermittency.

Influence of European Beech (Fagales: Fagaceae) Rot Hole Habitat Characteristics on Invertebrate Community Structure and Diversity.

Hollows of veteran trees (i.e., rot holes) provide habitat for many rare and threatened saproxylic invertebrates. Rot holes are highly heterogeneous, particularly in terms of substrate and microclimate conditions. There is, however, a dearth of information regarding the differences in biological communities inhabiting rot holes with different environmental conditions. Invertebrates were sampled from European beech (Fagus sylvatica) rot holes in Windsor, Savernake, and Epping Forests (United Kingdom). For each rot hole, physical and environmental conditions were measured, including tree diameter, rot hole dimensions, rot hole height, substrate density, water content, and water potential. These parameters were used to assess the influence of environmental conditions and habitat characteristics on invertebrate communities. Rot hole invertebrate communities were extremely diverse, containing both woodland generalist and saproxylic specialist taxa. Large variation in community structure was observed between rot holes and across woodlands; all sites supported threatened and endangered taxa. Environmental conditions in rot holes were highly variable within and between woodland sites, and communities were predominantly structured by these environmental conditions. In particular, turnover between invertebrate communities was linked to high ß-diversity. The linked heterogeneity of environmental conditions and invertebrate communities in rot holes suggests that management of deadwood habitats in woodlands should strive to generate environmental heterogeneity to promote invertebrate diversity. Additional research is required to define how management and conservation activities can further promote enhanced biodiversity across rot holes.

Spatial Ecology of Estuarine Crocodile (Crocodylus porosus) Nesting in a Fragmented Landscape.

The role that oil palm plays in the Lower Kinabatangan region of Eastern Sabah is of considerable scientific and conservation interest, providing a model habitat for many tropical regions as they become increasingly fragmented. Crocodilians, as apex predators, widely distributed throughout the tropics, are ideal indicator species for ecosystem health. Drones (or unmanned aerial vehicles (UAVs)) were used to identify crocodile nests in a fragmented landscape. Flights were targeted through the use of fuzzy overlay models and nests located primarily in areas indicated as suitable habitat. Nests displayed a number of similarities in terms of habitat characteristics allowing for refined modelling of survey locations. As well as being more cost-effective compared to traditional methods of nesting survey, the use of drones also enabled a larger survey area to be completed albeit with a limited number of flights. The study provides a methodology for targeted nest surveying, as well as a low-cost repeatable flight methodology. This approach has potential for widespread applicability across a range of species and for a variety of study designs.

Root Herbivores Drive Changes to Plant Primary Chemistry, but Root Loss Is Mitigated under Elevated Atmospheric CO2.

Above- and belowground herbivory represents a major challenge to crop productivity and sustainable agriculture worldwide. How this threat from multiple herbivore pests will change under anthropogenic climate change, via altered trophic interactions and plant response traits, is key to understanding future crop resistance to herbivory. In this study, we hypothesized that atmospheric carbon enrichment would increase the amount (biomass) and quality (C:N ratio) of crop plant resources for above- and belowground herbivore species. In a controlled environment facility, we conducted a microcosm experiment using the large raspberry aphid (Amphorophora idaei), the root feeding larvae of the vine weevil (Otiorhynchus sulcatus), and the raspberry (Rubus idaeus) host-plant. There were four herbivore treatments (control, aphid only, weevil only and a combination of both herbivores) and an ambient (aCO2) or elevated (eCO2) CO2 treatment (390 versus 650 ± 50 µmol/mol) assigned to two raspberry cultivars (cv Glen Ample or Glen Clova) varying in resistance to aphid herbivory. Contrary to our predictions, eCO2 did not increase crop biomass or the C:N ratio of the plant tissues, nor affect herbivore abundance either directly or via the host-plant. Root herbivory reduced belowground crop biomass under aCO2 but not eCO2, suggesting that crops could tolerate attack in a CO2 enriched environment. Root herbivory also increased the C:N ratio in leaf tissue at eCO2, potentially due to decreased N uptake indicated by lower N concentrations found in the roots. Root herbivory greatly increased root C concentrations under both CO2 treatments. Our findings confirm that responses of crop biomass and biochemistry to climate change need examining within the context of herbivory, as biotic interactions appear as important as direct effects of eCO2 on crop productivity.

The Effects of Supplementary Food on the Breeding Performance of Eurasian Reed Warblers Acrocephalus scirpaceus; Implications for Climate Change Impacts.

Understanding the mechanisms by which climate variation can drive population changes requires information linking climate, local conditions, trophic resources, behaviour and demography. Climate change alters the seasonal pattern of emergence and abundance of invertebrate populations, which may have important consequences for the breeding performance and population change of insectivorous birds. In this study, we examine the role of food availability in driving behavioural changes in an insectivorous migratory songbird; the Eurasian reed warbler Acrocephalus scirpaceus. We use a feeding experiment to examine the effect of increased food supply on different components of breeding behaviour and first-brood productivity, over three breeding seasons (2012-2014). Reed warblers respond to food-supplementation by advancing their laying date by up to 5.6 days. Incubation periods are shorter in supplemented groups during the warmest mean spring temperatures. Nestling growth rates are increased in nests provisioned by supplemented parents. In addition, nest predation is reduced, possibly because supplemented adults spend more time at the nest and faster nestling growth reduces the period of vulnerability of eggs and nestlings to predators (and brood parasites). The net effect of these changes is to advance the fledging completion date and to increase the overall productivity of the first brood for supplemented birds. European populations of reed warblers are currently increasing; our results suggest that advancing spring phenology, leading to increased food availability early in the breeding season, could account for this change by facilitating higher productivity. Furthermore, the earlier brood completion potentially allows multiple breeding attempts. This study identifies the likely trophic and behavioural mechanisms by which climate-driven changes in invertebrate phenology and abundance may lead to changes in breeding phenology, nest survival and net reproductive performance of insectivorous birds.

Antagonistic interactions between an invasive alien and a native coccinellid species may promote coexistence.

Despite the capacity of invasive alien species to alter ecosystems, the mechanisms underlying their impact remain only partly understood. Invasive alien predators, for example, can significantly disrupt recipient communities by consuming prey species or acting as an intraguild predator (IGP). Behavioural interactions are key components of interspecific competition between predators, yet these are often overlooked invasion processes. Here, we show how behavioural, non-lethal IGP interactions might facilitate the establishment success of an invading alien species. We experimentally assessed changes in feeding behaviour (prey preference and consumption rate) of native UK coccinellid species (Adalia bipunctata and Coccinella septempunctata), whose populations are, respectively, declining and stable, when exposed to the invasive intraguild predator, Harmonia axyridis. Using a population dynamics model parameterized with these experimental data, we predicted how intraguild predation, accommodating interspecific behavioural interactions, might impact the abundance of the native and invasive alien species over time. When competing for the same aphid resource, the feeding rate of A. bipunctata significantly increased compared to the feeding in isolation, while the feeding rate of H. axyridis significantly decreased. This suggests that despite significant declines in the UK, A. bipunctata is a superior competitor to the intraguild predator H. axyridis. In contrast, the behaviour of non-declining C. septempunctata was unaltered by the presence of H. axyridis. Our experimental data show the differential behavioural plasticity of competing native and invasive alien predators, but do not explain A. bipunctata declines observed in the UK. Using behavioural plasticity as a parameter in a population dynamic model for A. bipunctata and H. axyridis, coexistence is predicted between the native and invasive alien following an initial period of decline in the native species. We demonstrate how empirical and theoretical techniques can be combined to understand better the processes and consequences of alien species invasions for native biodiversity.

Habitat Use and Body Mass Regulation among Warblers in the Sahel Region during the Non-Breeding Season.

Migratory birds face significant challenges across their annual cycle, including occupying an appropriate non-breeding home range with sufficient foraging resources. This can affect demographic processes such as over-winter survival, migration mortality and subsequent breeding success. In the Sahel region of Africa, where millions of migratory songbirds attempt to survive the winter, some species of insectivorous warblers occupy both wetland and dry-scrubland habitats, whereas other species are wetland or dry-scrubland specialists. In this study we examine evidence for strategic regulation of body reserves and competition-driven habitat selection, by comparing invertebrate prey activity-density, warbler body size and extent of fat and pectoral muscle deposits, in each habitat type during the non-breeding season. Invertebrate activity-density was substantially higher in wetland habitats than in dry-scrubland. Eurasian reed warblers Acrocephalus scirpaceus occupying wetland habitats maintained lower body reserves than conspecifics occupying dry-scrub habitats, consistent with buffering of reserves against starvation in food-poor habitat. A similar, but smaller, difference in body reserves between wet and dry habitat was found among subalpine warblers Sylvia cantillans but not in chiffchaffs Phylloscopus collybita inhabiting dry-scrub and scrub fringing wetlands. Body reserves were relatively low among habitat specialist species; resident African reed warbler A. baeticatus and migratory sedge warbler A. schoenobaenus exclusively occupying wetland habitats, and Western olivaceous warblers Iduna opaca exclusively occupying dry habitats. These results suggest that specialists in preferred habitats and generalists occupying prey-rich habitats can reduce body reserves, whereas generalists occupying prey-poor habitats carry an increased level of body reserves as a strategic buffer against starvation.

The impact of increased food availability on reproduction in a long-distance migratory songbird: implications for environmental change?

Many populations of migratory songbirds are declining or shifting in distribution. This is likely due to environmental changes that alter factors such as food availability that may have an impact on survival and/or breeding success. We tested the impact of experimentally supplemented food on the breeding success over three years of northern wheatears (Oenanthe oenanthe), a species in decline over much of Europe. The number of offspring fledged over the season was higher for food-supplemented birds than for control birds. The mechanisms for this effect were that food supplementation advanced breeding date, which, together with increased resources, allowed further breeding attempts. While food supplementation did not increase the clutch size, hatching success or number of chicks fledged per breeding attempt, it did increase chick size in one year of the study. The increased breeding success was greater for males than females; males could attempt to rear simultaneous broods with multiple females as well as attempting second broods, whereas females could only increase their breeding effort via second broods. Multiple brooding is rare in the study population, but this study demonstrates the potential for changes in food availability to affect wheatear breeding productivity, primarily via phenotypic flexibility in the number of breeding attempts. Our results have implications for our understanding of how wheatears may respond to natural changes in food availability due to climate changes or changes in habitat management.

Elevated atmospheric CO2 impairs aphid escape responses to predators and conspecific alarm signals.

Research into the impact of atmospheric change on predator-prey interactions has mainly focused on density dependent responses and trophic linkages. As yet, the chemical ecology underpinning predator-prey interactions has received little attention in environmental change research. Group living animals have evolved behavioral mechanisms to escape predation, including chemical alarm signalling. Chemical alarm signalling between conspecific prey could be susceptible to environmental change if the physiology and behavior of these organisms are affected by changes in dietary quality resulting from environmental change. Using Rubus idaeus plants, we show that elevated concentrations of atmospheric CO2 (eCO2) severely impaired escape responses of the aphid Amphorophora idaei to predation by ladybird larvae (Harmonia axyridis). Escape responses to ladybirds was reduced by >50% after aphids had been reared on plants grown under eCO2. This behavioral response was rapidly induced, occurring within 24 h of being transferred to plants grown at eCO2 and, once induced, persisted even after aphids were transferred to plants grown at ambient CO2. Escape responses were impaired due to reduced sensitivity to aphid alarm pheromone, (E)-ß-farnesene, via an undefined plant-mediated mechanism. Aphid abundance often increases under eCO2, however, reduced efficacy of conspecific signalling may increase aphid vulnerability to predation, highlighting the need to study the chemical ecology of predator-prey interactions under environmental change.

Discontinuity in the responses of ecosystem processes and multifunctionality to altered soil community composition.

Ecosystem management policies increasingly emphasize provision of multiple, as opposed to single, ecosystem services. Management for such "multifunctionality" has stimulated research into the role that biodiversity plays in providing desired rates of multiple ecosystem processes. Positive effects of biodiversity on indices of multifunctionality are consistently found, primarily because species that are redundant for one ecosystem process under a given set of environmental conditions play a distinct role under different conditions or in the provision of another ecosystem process. Here we show that the positive effects of diversity (specifically community composition) on multifunctionality indices can also arise from a statistical fallacy analogous to Simpson's paradox (where aggregating data obscures causal relationships). We manipulated soil faunal community composition in combination with nitrogen fertilization of model grassland ecosystems and repeatedly measured five ecosystem processes related to plant productivity, carbon storage, and nutrient turnover. We calculated three common multifunctionality indices based on these processes and found that the functional complexity of the soil communities had a consistent positive effect on the indices. However, only two of the five ecosystem processes also responded positively to increasing complexity, whereas the other three responded neutrally or negatively. Furthermore, none of the individual processes responded to both the complexity and the nitrogen manipulations in a manner consistent with the indices. Our data show that multifunctionality indices can obscure relationships that exist between communities and key ecosystem processes, leading us to question their use in advancing theoretical understanding--and in management decisions--about how biodiversity is related to the provision of multiple ecosystem services.

Effects of elevated CO2 on litter chemistry and subsequent invertebrate detritivore feeding responses.

Elevated atmospheric CO2 can change foliar tissue chemistry. This alters leaf litter palatability to macroinvertebrate detritivores with consequences for decomposition, nutrient turnover, and food-web structure. Currently there is no consensus on the link between CO2 enrichment, litter chemistry, and macroinvertebrate-mediated leaf decomposition. To identify any unifying mechanisms, we presented eight invertebrate species from aquatic and terrestrial ecosystems with litter from Alnus glutinosa (common alder) or Betula pendula (silver birch) trees propagated under ambient (380 ppm) or elevated (ambient +200 ppm) CO2 concentrations. Alder litter was largely unaffected by CO2 enrichment, but birch litter from leaves grown under elevated CO2 had reduced nitrogen concentrations and greater C/N ratios. Invertebrates were provided individually with either (i) two litter discs, one of each CO2 treatment ('choice'), or (ii) one litter disc of each CO2 treatment alone ('no-choice'). Consumption was recorded. Only Odontocerum albicorne showed a feeding preference in the choice test, consuming more ambient- than elevated-CO2 birch litter. Species' responses to alder were highly idiosyncratic in the no-choice test: Gammarus pulex and O. albicorne consumed more elevated-CO2 than ambient-CO2 litter, indicating compensatory feeding, while Oniscus asellus consumed more of the ambient-CO2 litter. No species responded to CO2 treatment when fed birch litter. Overall, these results show how elevated atmospheric CO2 can alter litter chemistry, affecting invertebrate feeding behaviour in species-specific ways. The data highlight the need for greater species-level information when predicting changes to detrital processing-a key ecosystem function-under atmospheric change.

Arabidopsis thaliana and Pisum sativum models demonstrate that root colonization is an intrinsic trait of Burkholderia cepacia complex bacteria.

Burkholderia cepacia complex (Bcc) bacteria possess biotechnologically useful properties that contrast with their opportunistic pathogenicity. The rhizosphere fitness of Bcc bacteria is central to their biocontrol and bioremediation activities. However, it is not known whether this differs between species or between environmental and clinical strains. We investigated the ability of 26 Bcc strains representing nine different species to colonize the roots of Arabidopsis thaliana and Pisum sativum (pea). Viable counts, scanning electron microscopy and bioluminescence imaging were used to assess root colonization, with Bcc bacteria achieving mean (±sem) levels of 2.49±0.23×10(6) and 5.16±1.87×10(6) c.f.u. per centimetre of root on the A. thaliana and P. sativum models, respectively. The A. thaliana rhizocompetence model was able to reveal loss of colonization phenotypes in Burkholderia vietnamiensis G4 transposon mutants that had only previously been observed in competition experiments on the P. sativum model. Different Bcc species colonized each plant model at different rates, and no statistical difference in root colonization was observed between isolates of clinical or environmental origin. Loss of the virulence-associated third chromosomal replicon (>1 Mb DNA) did not alter Bcc root colonization on A. thaliana. In summary, Bcc bacteria possess intrinsic root colonization abilities irrespective of their species or source. As Bcc rhizocompetence does not require their third chromosomal replicon, the possibility of using synthetic biology approaches to engineer virulence-attenuated biotechnological strains is tractable.

Contrasting effects of elevated temperature and invertebrate grazing regulate multispecies interactions between decomposer fungi.

Predicting the influence of biotic and abiotic factors on species interactions and ecosystem processes is among the primary aims of community ecologists. The composition of saprotrophic fungal communities is a consequence of competitive mycelial interactions, and a major determinant of woodland decomposition and nutrient cycling rates. Elevation of atmospheric temperature is predicted to drive changes in fungal community development. Top-down regulation of mycelial growth is an important determinant of, and moderator of temperature-driven changes to, two-species interaction outcomes. This study explores the interactive effects of a 4 °C temperature increase and soil invertebrate (collembola or woodlice) grazing on multispecies interactions between cord-forming basidiomycete fungi emerging from colonised beech (Fagus sylvatica) wood blocks. The fungal dominance hierarchy at ambient temperature (16 °C; Phanerochaete velutina > Resinicium bicolor > Hypholoma fasciculare) was altered by elevated temperature (20 °C; R. bicolor > P. velutina > H. fasciculare) in ungrazed systems. Warming promoted the competitive ability of the fungal species (R. bicolor) that was preferentially grazed by all invertebrate species. As a consequence, grazing prevented the effect of temperature on fungal community development and maintained a multispecies assemblage. Decomposition of fungal-colonised wood was stimulated by warming, with implications for increased CO2 efflux from woodland soil. Analogous to aboveground plant communities, increasing complexity of biotic and abiotic interactions appears to be important in buffering climate change effects on soil decomposers.

Global climate change and above- belowground insect herbivore interactions.

Predicted changes to the Earth's climate are likely to affect above-belowground interactions. Our understanding is limited, however, by past focus on two-species aboveground interactions mostly ignoring belowground influences. Despite their importance to ecosystem processes, there remains a dearth of empirical evidence showing how climate change will affect above-belowground interactions. The responses of above- and belowground organisms to climate change are likely to differ given the fundamentally different niches they inhabit. Yet there are few studies that address the biological and ecological reactions of belowground herbivores to environmental conditions in current and future climates. Even fewer studies investigate the consequences of climate change for above-belowground interactions between herbivores and other organisms; those that do provide no evidence of a directed response. This paper highlights the importance of considering the belowground fauna when making predictions on the effects of climate change on plant-mediated interspecific interactions.

Reciprocal feeding facilitation between above- and below-ground herbivores.

Interspecific interactions between insect herbivores predominantly involve asymmetric competition. By contrast, facilitation, whereby herbivory by one insect benefits another via induced plant susceptibility, is uncommon. Positive reciprocal interactions between insect herbivores are even rarer. Here, we reveal a novel case of reciprocal feeding facilitation between above-ground aphids (Amphorophora idaei) and root-feeding vine weevil larvae (Otiorhynchus sulcatus), attacking red raspberry (Rubus idaeus). Using two raspberry cultivars with varying resistance to these herbivores, we further demonstrate that feeding facilitation occurred regardless of host plant resistance. This positive reciprocal interaction operates via an, as yet, unreported mechanism. Specifically, the aphid induces compensatory growth, possibly as a prelude to greater resistance/tolerance, whereas the root herbivore causes the plant to abandon this strategy. Both herbivores may ultimately benefit from this facilitative interaction.

Bottom-up determination of soil collembola diversity and population dynamics in response to interactive climatic factors.

Soil invertebrate contributions to decomposition are climate dependent. Understanding the influence of abiotic factors on soil invertebrate population dynamics will strengthen predictions regarding ecosystem functioning under climate change. As well as being important secondary decomposers, mycophagous collembola exert a strong influence on the growth and activity of primary decomposers, particularly fungi. Species-specific grazing preferences for different fungi enable fungal community composition to influence the direct impacts of climate change on collembola populations. We investigate the interactive roles of altered abiotic conditions (warming, wetting and drying) and the identity of the dominant decomposer fungus in determining collembola community dynamics in woodland soil mesocosms. The bottom-up influence of the dominant component of the fungal resource base was an important mediator of the direct climatic impacts on collembola populations. The positive influences of warming and wetting, and the negative influence of drying, on collembola abundance and diversity were much less pronounced in fungal-inoculation treatments, in which populations were reduced compared with uninoculated mesocosms. We conclude that the thick, sclerotised cords of the competitively dominant decomposer fungi reduced the biomass of smaller, more palatable soil fungi, limiting the size of collembola populations and their ability to respond to altered abiotic conditions.

The impact of increased food availability on survival of a long-distance migratory bird.

Temperature-, rainfall- and habitat-driven change in food availability is one likely mechanism by which anthropogenic factors may affect animal population dynamics and species distributions. Long-distance migratory birds must synchronize their migrations with food availability at locations hundreds or thousands of kilometers apart, so changes in the overall abundance of food or the phenology of peaks in food availability may be critical factors influencing annual survival. In this study we used experimental food supplementation at a breeding site to examine and compare the impact of changing food supply on annual survival of adult migratory birds and their offspring. We provided supplemental food to Northern Wheatears (Oenanthe oenanthe) breeding on Fair Isle, UK, to infer the sensitivity of annual survival to increases in natural food availability. Food-supplemented wheatears exhibited higher rates of annual survival than control wheatears, and the strength of this effect varied with age. Food supplementation led to 1.56 times higher annual survival of juveniles and 1.22 times higher survival of adults. Survival of juveniles was related to their own food availability as fledglings, but not to whether their parents were food-supplemented or unfed control adults. This increased survival, combined with increased breeding productivity associated with food supplementation, implies that an increase in natural food availability, of the magnitude simulated in our experiment, would increase the population growth rate of wheatears on Fair Isle from approximately lambda = 0.93 (a contracting population) to lambda = 1.14 (an expanding population).