Trophic rewilding establishes a landscape of fear: Tasmanian devil introduction increases risk‐sensitive foraging in a key prey species

Global declines of large carnivores have reduced the ‘landscape of fear’ that constrains the behaviour of other species. In recent years, active and passive trophic rewilding have potentially begun restoring these lost top–down controls. The Tasmanian devil Sarcophilus harrisii has declined severely due to a novel transmissible cancer. In response to extinction fears, devils were introduced to the devil‐free Maria Island, where their abundance rapidly increased. We tested how this introduction influenced risk‐sensitive foraging in the common brushtail possum Trichosurus vulpecula, a major prey species for devils, using giving‐up densities (GUDs). Before the introduction of devils, possum GUDs on Maria Island were indistinguishable from the long‐diseased region of Tasmania, where devils have been rare since ~2000. Three years after devil introduction, GUDs were 64% higher on Maria Island than the control region, demonstrating that after an initial period of high mortality, possums quickly adopted risk‐sensitive foraging behaviours. Devil activity across Maria Island was variable, leading to a heterogeneous landscape of fear and highlighting that top predators must be at functional densities to elicit behavioural responses from prey. Our study provides strong evidence that top predators modify the behaviour of prey by instilling fear, causing rapid ecological change following recoveries.     

A long‐term habitat fragmentation experiment leads to morphological change in a species of carabid beetle

1. Habitat fragmentation and transformation are key drivers of species declines in landscapes. Most of the current understanding of species' responses to environmental change originates from studies of populations and communities. However, phenotypic variation offers another key aspect of species responses and could provide additional insights into the functional drivers of population change. 2. The goal of this study was to address this gap by exploring the morphological changes of a species of carabid beetle (Notonomus resplendens) with a known population response to the Wog Wog Habitat Fragmentation Experiment in Australia. We measured morphological traits associated with body size, head width, and dispersal ability. We quantified patterns of morphological variation over time and between native Eucalyptus forest fragments and the surrounding pine plantation matrix and the continuous intact native Eucalyptus forest controls. 3. We found sexually dimorphic morphological changes in response to the experimental treatments. Males increased in size, had larger legs and had smaller interocular widths in the matrix in both the short and long terms. Conversely, females became comparatively smaller and had increased interocular widths in the same treatments. Effects in the fragments were similar to those in the matrix, but exhibited more uncertainty. 4. Our results demonstrate that species can show morphological change in response to environmental change over very short time periods. We demonstrate that using both population and morphological data allows stronger inferences about the mechanisms behind species responses to environmental change.     

Soil fauna responses to invasive alien plants are determined by trophic groups and habitat structure: a global meta‐analysis

Despite increasing frequency of invasions by alien plant species with widespread ecological and economic consequences, it remains unclear how belowground compartments of ecosystems are impacted. In order to synthetize current knowledge and provide future directions for research we performed a meta‐analysis assessing the impact of invasive alien plant species on soil fauna abundance. Compared to previous synthesis on this topic, we included together in our model the trophic group of each soil faunal taxa (from herbivores to predators) and habitat structure, namely open and closed habitats (i.e. grass and shrub dominated areas versus forested areas). In doing so, we highlighted that both moderators strongly interact to determine the response of soil fauna to the presence of invasive alien plants. Soil fauna abundance increase in the presence of invasive species only in closed habitats (+18.2%). This pattern of habitat‐dependent response (positive effect in closed habitats) was only found for primary consumers (i.e. herbivores +29.1% and detritivores +66.7%) within both detritus‐based and live root‐based trophic pathways. Abundances of predators and microbivores did not respond to the presence of IAS irrespective of habitat structure. For several groups, the habitat structure (open or closed) significantly drove their responses to the presence of invasive alien species. In addition, we carefully considered potential sources of bias (e.g. geographic, taxonomic and functional) within the collected data in an attempt to highlight gaps in available knowledge on the subject. Our findings support the conclusions of previous studies on the subject by demonstrating 1) that soil fauna abundance is impacted by biological invasions, 2) that initial habitat structure has a strong influence on the outcome and 3) that responses within the soil fauna differ between trophic levels with a stronger response of primary consumers.     

Influence of within‐field position and adjoining habitat on carabid beetle assemblages in winter wheat

• 1 The influence of within‐field position and adjoining habitat on carabid beetles was studied in 20 winter wheat fields in ten different Swiss agricultural landscapes. In each landscape, two winter wheat fields (one with adjoining sown wildflower area and one with adjoining grassy margin) were investigated.
• 2 Carabid beetles were caught in pitfall traps 3 and 30 m from the edge in each of the 20 wheat fields. Significantly more individuals were found in the centres (30‐m position) than at the edges (3‐m position). Conversely, species richness was significantly higher at the field edges than in the centres.
• 3 Of the ten most abundant species, Poecilus cupreus, Agonum muelleri and Pterostichus melanarius were significantly more abundant in the field centres than at the edges. Harpalus rufipes was significantly more abundant in the fields adjoining sown wildflower areas than in the fields adjoining grassy margins.
• 4 In conclusion, the response of carabid beetles to within‐field position and adjoining habitats was species specific. This needs to be taken into account in habitat management for biodiversity conservation and pest control.

     

Evolutionary responses of dispersal distance to landscape structure and habitat loss

It is generally well understood that some ecological factors select for increased and others for decreased dispersal. However, it has remained difficult to assess how the evolutionary dynamics are influenced by the spatio-temporal structure of the environment. We address this question with an individual-based model that enables habitat structure to be controlled through variables such as patch size, patch turnover rate, and patch quality. Increasing patch size at the expense of patch density can select for more or less dispersal, depending on the initial configuration. In landscapes consisting of high-quality and long-lived habitat patches, patch degradation selects for increased dispersal, yet patch loss may select for reduced dispersal. These trends do not depend on the component of life-history that is affected by habitat quality or the component of life-history through which density-dependence operates. Our results are based on a mathematical method that enables derivation of both the evolutionary stable strategy and the stationary genotype distribution that evolves in a polymorphic population. The two approaches generally lead to similar predictions. However, the evolutionary stable strategy assumes that the ecological and evolutionary time scales can be separated, and we find that violation of this assumption can critically alter the evolutionary outcome.     

Dispersal of a carabid beetle in farmland is driven by habitat‐specific motility and preference at habitat interfaces

Carabid beetles are common predators of pest insects and weed seeds in agricultural systems. Understanding their dispersal across farmland is important for designing farms and landscapes that support pest and weed biological control. Little is known, however, about the effect of farmland habitat discontinuities on dispersal behaviour and the resulting redistribution of these beetles. We released 1,985 well‐fed and 1,680 food‐deprived individuals of the predatory carabid beetle Pterostichus melanarius (Illiger) (Coleoptera: Carabidae) on a farm in Wageningen, The Netherlands. We recaptured 23.6% of those beetles over a period of 23 days in 2010. The farmland comprised agricultural fields with various crop species and tillage, separated by strips of perennial vegetation. We developed discrete Fokker‐Planck diffusion models to describe dispersal based on motility (m² day^?1) and preferential behaviour at habitat interfaces. We used model selection and Akaike’s information criterion to determine whether movement patterns were driven by variation in motility between habitats, preferential behaviour at habitat interfaces, or both. Model selection revealed differences in motility among habitats and gave strong support for preferential behaviour at habitat interfaces. Behaviour at interfaces between crop and perennial vegetation was asymmetric, with beetles preferentially moving towards the crop. Furthermore, beetles had lower motility in perennial strips than in arable fields. Also between arable habitats movement was asymmetric, with beetles preferentially moving towards the habitat in which motility was lowest. Neither crop type nor tillage explained differences in motility between crop habitats. Recapture data representing dispersal patterns of beetles were best described by a model that accounted for differences in motility between farmland habitats and preferential behaviour at habitat interfaces. Motility in farmland and behaviour at interfaces can also be estimated for other organisms and farmland habitats to support design of farmland conducive to natural pest suppression. Landscape design for early recruitment of carabids into arable fields should take into account the quantity and quality of resource habitats in the landscape, their proximity to crop fields, movement rates, and the possibility of movement responses at interfaces between landscape elements.     

地面节肢动物营养类群对土地覆被变化和管理扰动的响应

近几十年来,黑河中游荒漠绿洲过渡带大面积天然沙质草地被转变为农田和防风固沙人工林,然而目前尚缺乏土地利用/覆被变化和管理措施对地面节肢动物多样性形成和维持影响的系统研究。以天然沙质草地转变为的人工梭梭灌木林、人工杨树林、人工樟子松林和农田为研究对象,以天然草地为对照,通过对5种样地地面节肢动物群落的调查及非生物环境因子的测定,采用典范对应分析(CCA)等方法,系统探讨了土地覆被和管理强度(天然草地和人工梭梭林实施零管理、杨树和樟子松人工林实施灌溉管理、农田实施灌溉和施肥管理)变化对地面节肢动物营养类群组成、数量和物种丰富度的影响。主要结果是:(1) 在干旱环境下,单纯改变土地覆被并不会导致土壤环境的显著变化,而高强度的管理扰动加速了土壤环境的演变过程。(2)不同土地覆被和管理措施对营养类群组成及其数量和丰富度的调控作用不同,将天然草地转变为杨树和樟子松人工林和农田后,显著提高了捕食性和植食性类群的比例,降低了'腐食性+杂食性’类群的比例;转变为人工梭梭林后显著降低了3种营养类群的数量;转变为人工杨树林后显著降低了'腐食性+杂食性’类群数量;转变为人工樟子松林后显著增加了植食性类群数量而降低了其它类群的数量;转变为农田后显著增加了植食性类群数量而降低了'腐食性+杂食性’类群数量。(3)影响捕食性动物分布的关键环境因子是地面温度、土壤硝态氮和田间持水量,影响植食性动物分布的关键因子是土壤粘粉粒、土壤硝态氮、土壤容重和pH值,影响'腐食性+杂食性’动物分布的关键因子是地面温度和田间持水量。主要结论是:土地覆被变化与管理措施相互作用对地面节肢动物群落组装过程及其多样性形成具有重要调控作用,尽管其相对重要性不同。     

Forty years of carabid beetle research in Europe - from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation

‘Carabidologists do it all’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family Carabidae is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.     

Seasonal differences in population‐, ensemble‐ and community‐level responses of bats to landscape structure in Amazonia

The amount (composition) and spatial arrangement (configuration) of forest patches in fragmented landscapes influence the accessibility, as well as the abundance and diversity of resources available to bats. Moreover, tropical fruit and insect abundance differ seasonally in response to changes in precipitation, and many bats in the family Phyllostomidae employ seasonal reproductive strategies. Because reproductive activities involve constraints on time and energy as well as increased nutritional demands, foraging behavior and home range size may differ between wet and dry seasons. Nonetheless, seasonal variation in response to landscape structure by bats has not been examined previously. Consequently, population‐, ensemble‐ and assemblage‐level responses of phyllostomids to landscape composition and configuration were quantified separately during the wet and dry season at three circular focal scales (1, 3 and 5 km radii) for 14 sites in fragmented lowland Amazon forest. Responses to landscape characteristics were scale‐dependent, species‐specific, and seasonal. Abundances of frugivores responded to landscape composition in the dry season and to landscape configuration in the wet season. Conversely, abundances of animalivores responded to landscape configuration in the dry season and to landscape composition in the wet season. Divergent responses to landscape structure between seasons suggest that variation in resource abundance and diversity play a significant role in structuring population‐, ensemble‐ and assemblage‐level patterns. As such, considerations of the effects of dietary flexibility and reproductive constraints on foraging strategies and habitat use may be important when designing management plans that successfully promote long‐term persistence of biodiversity in fragmented landscapes.     

Applying stable isotopes to examine food‐web structure: an overview of analytical tools

Stable isotope analysis has emerged as one of the primary means for examining the structure and dynamics of food webs, and numerous analytical approaches are now commonly used in the field. Techniques range from simple, qualitative inferences based on the isotopic niche, to Bayesian mixing models that can be used to characterize food‐web structure at multiple hierarchical levels. We provide a comprehensive review of these techniques, and thus a single reference source to help identify the most useful approaches to apply to a given data set. We structure the review around four general questions: (1) what is the trophic position of an organism in a food web?; (2) which resource pools support consumers?; (3) what additional information does relative position of consumers in isotopic space reveal about food‐web structure?; and (4) what is the degree of trophic variability at the intrapopulation level? For each general question, we detail different approaches that have been applied, discussing the strengths and weaknesses of each. We conclude with a set of suggestions that transcend individual analytical approaches, and provide guidance for future applications in the field.     

Traits reveal how habitat‐quality gradients structure small mammal communities in a fragmented tropical landscape

Understanding the relationships among community structure, vegetation structure and availability of food resources are a key to unravelling the ecological processes that structure biological communities. In this study, we tested (i) whether the composition of small mammal communities changed across gradients in habitat quality in tropical forest fragments, and (ii) whether any observed change could be explained by the functional traits of species. We sampled 24 trapping grids in fragments of semi‐deciduous forest, in each of two 6‐month periods. We considered each trapping grid as a sampling unit, for which we collected three datasets: an environmental matrix (vegetation structure and food resource availability), the abundance of small mammal species (community structure) and a matrix of functional traits (ecological and morphological traits which express tolerance to habitat disturbance and trophic guild). We used an RLQ approach to evaluate the association between traits and environmental gradients. Forest‐specialist and scansorial–arboreal species were associated with more complex habitat that had greater litter and canopy cover and more fallen logs. In relation to trophic guilds, granivore (fruit seeds), insectivorous and omnivorous species were also associated with higher complexity habitat, while frugivores were associated with shrub cover and availability of fruits. We conclude that functional traits (habitat use, use of vertical strata and diet) provide valuable insights into the distribution of small mammals along gradients of habitat quality in tropical forest fragments. We highlight that communities studies in fragmented landscapes should investigate the different components of biodiversity not only in landscape‐scale but also in habitat scale. Abstract in Portuguese is available with online material.     

Interactions among body size,trophic level,and dispersal traits predict beetle detectability and occurrence responses to fire

1. Testing the extent to which traits act alone or in combination with other traits to influence responses to fire informs the trade-off between increased generalisation using single traits and increased predictive power using interactions. This study investigated the following question: do four traits (body size, trophic group, dispersal ability, and stratum of the ecosystem), alone or in combination, best explain changes in beetle occurrence with time since fire? 2. The data from 4 years and 15 independent fires in southern Australia were analysed using generalised linear mixed models. The study also assessed whether detectability depends on time since fire using multi-year detection models, because detectability has the potential to confound occurrence patterns. 3. The best model included the three-way combination of size, flight, and trophic level interacting with time since fire and with year. The relationship between detectability and time since fire was similar to the occurrence relationship in six of the 10 trait–combination groups, with flightless species generally showing reduced detection probability as time since fire increased. Detectability did not confound occurrence responses for four trait groups, with three increasing with time since fire and one decreasing. 4. Generalisation using main effects of traits risks oversimplifying animal responses to fire, because combinations of traits influence the direction and magnitude of the response. Also, taking detectability into account is critical to correctly interpretating occupancy data. Three-way trait combinations that differ by just one trait, particularly dispersal ability, can result in either negligible effects of disturbance on detectability or strong effects that influence observed occurrence.     

The responses of polyphagous predators to prey spatial heterogeneity: aggregation by carabid and staphylinid beetles to their cereal aphid prey

Abstract. 1. Discrete patches of aphids were artificially created in winter wheat fields in 1982 and in 1983 using field cages.
2. Aggregation by polyphagous predators at these patches following cage removal was measured using pitfall traps in patch and control areas.
3. Several species of polyphagous beetle (Carabidae and Staphylinidae) aggregated in these patches while other species did not.
4. The consequences of aggregation by polyphagous predators to a single prey species are discussed.     

Stage‐dependent responses to emergent habitat heterogeneity: consequences for a predatory insect population in a coffee agroecosystem

Interactions among members of biological communities can create spatial patterns that effectively generate habitat heterogeneity for other members in the community, and this heterogeneity might be crucial for their persistence. For example, stage‐dependent vulnerability of a predatory lady beetle to aggression of the ant, Azteca instabilis, creates two habitat types that are utilized differently by the immature and adult life stages of the beetle. Due to a mutualistic association between A. instabilis and the hemipteran Coccus viridis – which is A. orbigera main prey in the area – only plants around ant nests have high C. viridis populations. Here, we report on a series of surveys at three different scales aimed at detecting how the presence and clustered distribution of ant nests affect the distribution of the different life stages of this predatory lady beetle in a coffee farm in Chiapas, Mexico. Both beetle adults and larvae were more abundant in areas with ant nests, but adults were restricted to the peripheries of highest ant activity and outside the reach of coffee bushes containing the highest densities of lady beetle larvae. The abundance of adult beetles located around trees with ants increased with the size of the ant nest clusters but the relationship is not significant for larvae. Thus, we suggest that A. orbigera undergoes an ontogenetic niche shift, not through shifting prey species, but through stage‐specific vulnerability differences against a competitor that renders areas of abundant prey populations inaccessible for adults but not for larvae. Together with evidence presented elsewhere, this study shows how an important predator is not only dependent on the existence of two qualitatively distinct habitat types, but also on the spatial distribution of these habitats. We suggest that this dependency arises due to the different responses that the predator's life stages have to this emergent spatial pattern.     

Home‐range size and habitat use of European Nightjars Caprimulgus europaeus nesting in a complex plantation‐forest landscape

In Europe, the consequences of commercial plantation management for birds of conservation concern are poorly understood. The European Nightjar Caprimulgus europaeus is a species of conservation concern across Europe due to population depletion through habitat loss. Pine plantation‐forest is now a key Nightjar nesting habitat, particularly in northwestern Europe, and increased understanding of foraging habitat selection is required. We radiotracked 31 Nightjars in an extensive (185‐km²) complex conifer plantation landscape in 2009 and 2010. Home‐range 95% kernels for females, paired males and unpaired males were an order of magnitude larger than song territories of paired males, emphasizing the importance of habitats beyond the song territory. Nightjars travelled a mean maximum distance of 747 m from the territory centre each night. Home‐range placement relative to landscape composition was examined by compositional analysis. Pre‐closure canopy forest (aged 5–10 years) was selected at all scales (MCP, 95% and 50% kernels), with newly planted forest (aged 0–4 years) also selected within 50% kernels. For telemetry fixes relative to habitat composition within 2 km of their territory centre, individuals again selected pre‐closure and newly planted forest, and also grazed grass heath. Open ungrazed habitat was not selected, with implications for open habitat planning for biodiversity conservation within public‐owned forests. Despite the Nightjars’ selection for younger growth, moth biomass was greater in older forest stands, suggesting that foraging site selection reflects ease of prey capture rather than prey abundance. Within large plantation‐forest landscapes, a variety of growth stages is important for this species and our results suggest that grazing of open habitats within and adjacent to forest will additionally benefit the European Nightjar.     

Organic farming affects the potential of a granivorous carabid beetle to control arable weeds at local and landscape scales

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Using structural sustainability for forest health monitoring and triage: Case study of a mountain pine beetle (Dendroctonus ponderosae)-impacted landscape

Heavy disturbance-induced mortality can negatively impact forest biota, functions, and services by drastically altering the forest structures that create stable environmental conditions. Disturbance impacts on forest structure can be assessed using structural sustainability⿿the degree of balance between living and dead portions of a tree population⿿s size-class distribution⿿which is the basis of baseline mortality analysis. This analysis uses a conditionally calibrated reference level (i.e., baseline mortality) against which to detect significant mortality deviations without need for historical references. Recently, a structural sustainability index was developed by parameterizing results of this analysis to allow spatial and temporal comparisons within or among forested landscapes. The utility of this index as a tool for forest health monitoring programs and triage decisions has not been examined. Here, we investigated this utility by retrospectively analyzing the structural sustainability of a mountain pine beetle (Dendroctonus ponderosae)-impacted, lodgepole pine (Pinus contorta)-dominated landscape annually from 2000 to 2006 as well as among watersheds. We show that temporal patterns of structural sustainability at the landscape-level reflect accumulating beetle-induced mortality as well as beetle-lodgepole pine ecology. At the watershed-level, this sustainability spatially varied with 2006 beetle-induced mortality. Further, structural sustainability satisfies key criteria for effective indicators of ecosystem change. We conclude that structural sustainability is a viable tool upon which to base or supplement forest health monitoring and triage programs, and could potentially increase the efficacy of such programs under current and future climate change-associated disturbance patterns.     

Relative influence of habitat structure,species interactions and rainfall on the post‐fire population dynamics of ground‐dwelling vertebrates

The long‐term impacts of wildfires on animal populations are largely unknown. We used time‐series data based on a tracking index, from coastal NSW spanning 28 years after a wildfire, to investigate the relative influence of habitat structure, species interactions and climate on post‐fire animal population dynamics. The fire had an immediate impact on habitat structure, reducing and simplifying vegetation cover, which then underwent post‐fire successional change including an increase and plateau in tree canopy cover; an increase, stabilization and then decline in shrub cover; and an increase in ground litter cover. Population changes of different animal species were influenced by different components of successional change, but there was also evidence that species interactions were important. For example, bandicoots (Isoodon obesulus and Perameles nasuta combined) increased concurrent with an increase in shrub cover then declined at a faster rate than a direct association with senescing shrub cover would suggest, while the feral cat (Felis catus) population changed with the bandicoot population, suggesting a link between these species. Potoroos (Potorous tridactylus) increased 10 years after the fire concurrent with the closing tree canopy, but there was also evidence of a negative association with feral foxes (Vulpes vulpes). Variation in rainfall did not have significant effects on the population dynamics of any species. Our results suggest that changes in habitat structure play a key role in the post‐fire dynamics of many ground‐dwelling animals and hence different fire regimes are likely to influence animal dynamics through their effects on habitat structure. However, the role of predator–prey interactions, particularly with feral predators, is less clear and further study will require manipulative experiments of predators in conjunction with fire treatments to determine whether feral predator control should be integrated with fire management to improve outcomes for some native species.