Contrasting responses of small mammals to fire and topographic refugia

Unburnt patches within burnt landscapes are expected to provide an important resource for fauna, potentially acting as a refuge from direct effects of fire and allowing animals to persist in burnt landscapes. Nevertheless, there is little information about the way refugia are used by fauna and how populations may be affected by them. Planned burns are often patchy, with unburnt areas generally associated with gully systems providing a good opportunity to study faunal use of refugia. We used a before–after control‐impact design associated with a planned burn in south eastern Australia to investigate how two small mammal species, the bush rat Rattus fuscipes and agile antechinus Antechinus agilis, used unburnt gully systems within a larger burnt area. We tested three alternative hypotheses relating to post‐fire abundance: (i) active refugia – abundance would increase in unburnt patches because of a post‐fire shift of individuals from burnt to unburnt areas; (ii) passive refugia – abundance in unburnt patches would remain similar to pre‐fire levels; and (iii) limited or no refugia – abundance would reduce in unburnt patches related to the change induced by fire in the wider landscape. We found the two species responded differently to the presence of unburnt refugia in the landscape. Relative to controls, fire had little effect on bush rat abundance in gullies, supporting hypothesis 2. In contrast, agile antechinus abundance increased in gullies immediately post‐fire consistent with a shift of individuals from burnt parts of the landscape, supporting hypothesis 1. Differences in site fidelity, habitat use and intraspecific competition between these species are suggested as likely factors influencing responses to refugia. The way unburnt patches function as faunal refugia and the subsequent influence they have on post‐fire population dynamics, will to some extent depend on the life history attributes of individual species.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Butterfly edge effects are predicted by a simple model in a complex landscape

Edge responses have been studied for decades and form a critical component of our understanding of how organisms respond to landscape structure and habitat fragmentation. Until recently, however, the lack of a general, conceptual framework has made it difficult to make sense of the patterns and variability reported in the edge literature. We present a test of an edge effects model which predicts that organisms should avoid edges with less-preferred habitat, show increased abundance near edges with preferred habitat or habitat containing complementary resources, and show no response to edges with similar-quality habitat that offers only supplementary resources. We tested the predictions of this model against observations of the edge responses of 15 butterfly species at 12 different edge types within a complex, desert riparian landscape. Observations matched model predictions more than would be expected by chance for the 211 species/edge combinations tested over 3 years of study. In cases where positive or negative edge responses were predicted, observed responses matched those predictions 70% of the time. While the model tends to underpredict neutral results, it was rare that an observed edge response contradicted that predicted by the model. This study also supported the two primary ecological mechanisms underlying the model, although not equally. We detected a positive relationship between habitat preferences and the slope of the observed edge response, suggesting that this basic life history trait underlies edge effects and influences their magnitude. Empirical evidence also suggested the presence of complementary resources underlies positive edge responses, but only when completely confined to the adjacent habitat. This multi-species test of a general edge effects model at multiple edge types shows that resource-based mechanisms can explain many edge responses and that a modest knowledge of life history attributes and resource availability is sufficient for predicting and understanding many edge responses in complex landscapes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Upper thermal limits predict herpetofaunal responses to forest edge and cover

Amphibians and reptiles are sensitive to changes in the thermal environment, which varies considerably in human-modified landscapes. Although it is known that thermal traits of species influence their distribution in modified landscapes, how herpetofauna respond specifically to shifts in ambient temperature along forest edges remains unclear. This may be because most studies focus on local-scale metrics of edge exposure, which only account for a single edge or habitat patch. We predicted that accounting for the combined effect of multiple habitat edges in a landscape would best explain herpetofaunal response to thermally mediated edge effects. We (1) surveyed herpetofauna at two lowland, fragmented forest sites in central Colombia, (2) measured the critical thermal maximum (CT_max) of the species sampled, (3) measured their edge exposure at both local and landscape scales, and (4) created a thermal profile of the landscape itself. We found that species with low CT_max occurred both further from forest edges and in areas of denser vegetation, but were unaffected by the landscape-scale configuration of habitat edges. Variation in the thermal landscape was driven primarily by changes in vegetation density. Our results suggest that amphibians and reptiles with low CT_max are limited by both canopy gaps and proximity to edge, making them especially vulnerable to human modification of tropical forest. Abstract in Spanish is available with online material.     

Landscape‐level effects on aboveground biomass of tropical forests: A conceptual framework

Despite the general recognition that fragmentation can reduce forest biomass through edge effects, a systematic review of the literature does not reveal a clear role of edges in modulating biomass loss. Additionally, the edge effects appear to be constrained by matrix type, suggesting that landscape composition has an influence on biomass stocks. The lack of empirical evidence of pervasive edge‐related biomass losses across tropical forests highlights the necessity for a general framework linking landscape structure with aboveground biomass. Here, we propose a conceptual model in which landscape composition and configuration mediate the magnitude of edge effects and seed‐flux among forest patches, which ultimately has an influence on biomass. Our model hypothesizes that a rapid reduction of biomass can occur below a threshold of forest cover loss. Just below this threshold, we predict that changes in landscape configuration can strongly influence the patch's isolation, thus enhancing biomass loss. Moreover, we expect a synergism between landscape composition and patch attributes, where matrix type mediates the effects of edges on species decline, particularly for shade‐tolerant species. To test our conceptual framework, we propose a sampling protocol where the effects of edges, forest amount, forest isolation, fragment size, and matrix type on biomass stocks can be assessed both collectively and individually. The proposed model unifies the combined effects of landscape and patch structure on biomass into a single framework, providing a new set of main drivers of biomass loss in human‐modified landscapes. We argue that carbon trading agendas (e.g., REDD+) and carbon‐conservation initiatives must go beyond the effects of forest loss and edges on biomass, considering the whole set of effects on biomass related to changes in landscape composition and configuration.     

Landscape context modulates alien plant invasion in Mediterranean forest edges

Natural habitats in human-altered landscapes are especially vulnerable to biological invasions, especially in their edges. We aim to understand the influence of landscape and local characteristics on biological invasions by exploring the level of plant invasion and alien species traits in forest edges in highly urbanized landscapes. We identified all plant species in 73 paired plots in the edge and 50 m towards the interior of the forest. We explored the association between alien species richness and similarity in species composition between edge and interior plots with landscape and local variables, using generalized linear models and variance partitioning techniques. Then, we performed Fourth-corner analyses to explore the association between alien plant traits and local and landscape variables. In contrast to native species richness, alien species richness was more affected by the surrounding landscape than by the local characteristics of the edge. Road proximity was positively associated with alien species richness and proportion and was its most important correlate, whereas disturbance was negatively associated with native species richness and was its most influential factor. Alien plant traits were also primarily associated with landscape characteristics. For instance, species of Mediterranean origin and introduced for agriculture were associated with higher agriculture use in the landscape. Our findings suggest that risk analyses of habitat vulnerability to invasion must consider the landscape context in order to successfully predict highly invaded areas and identify potentially successful invaders.     

How fire interacts with habitat loss and fragmentation

Biodiversity faces many threats and these can interact to produce outcomes that may not be predicted by considering their effects in isolation. Habitat loss and fragmentation (hereafter ‘fragmentation’) and altered fire regimes are important threats to biodiversity, but their interactions have not been systematically evaluated across the globe. In this comprehensive synthesis, including 162 papers which provided 274 cases, we offer a framework for understanding how fire interacts with fragmentation. Fire and fragmentation interact in three main ways: (i) fire influences fragmentation (59% of 274 cases), where fire either destroys and fragments habitat or creates and connects habitat; (ii) fragmentation influences fire (25% of cases) where, after habitat is reduced in area and fragmented, fire in the landscape is subsequently altered because people suppress or ignite fires, or there is increased edge flammability or increased obstruction to fire spread; and (iii) where the two do not influence each other, but fire interacts with fragmentation to affect responses like species richness, abundance and extinction risk (16% of cases). Where fire and fragmentation do influence each other, feedback loops are possible that can lead to ecosystem conversion (e.g. forest to grassland). This is a well-documented threat in the tropics but with potential also to be important elsewhere. Fire interacts with fragmentation through scale-specific mechanisms: fire creates edges and drives edge effects; fire alters patch quality; and fire alters landscape-scale connectivity. We found only 12 cases in which studies reported the four essential strata for testing a full interaction, which were fragmented and unfragmented landscapes that both span contrasting fire histories, such as recently burnt and long unburnt vegetation. Simulation and empirical studies show that fire and fragmentation can interact synergistically, multiplicatively, antagonistically or additively. These cases highlight a key reason why understanding interactions is so important: when fire and fragmentation act together they can cause local extinctions, even when their separate effects are neutral. Whether fire–fragmentation interactions benefit or disadvantage species is often determined by the species' preferred successional stage. Adding fire to landscapes generally benefits early-successional plant and animal species, whereas it is detrimental to late-successional species. However, when fire interacts with fragmentation, the direction of effect of fire on a species could be reversed from the effect expected by successional preferences. Adding fire to fragmented landscapes can be detrimental for species that would normally co-exist with fire, because species may no longer be able to disperse to their preferred successional stage. Further, animals may be attracted to particular successional stages leading to unexpected responses to fragmentation, such as higher abundance in more isolated unburnt patches. Growing human populations and increasing resource consumption suggest that fragmentation trends will worsen over coming years. Combined with increasing alteration of fire regimes due to climate change and human-caused ignitions, interactions of fire with fragmentation are likely to become more common. Our new framework paves the way for developing a better understanding of how fire interacts with fragmentation, and for conserving biodiversity in the face of these emerging challenges.     

Edge effects across boundaries between natural and restored jarrah (Eucalyptus marginata) forests in south‐western Australia

Edge effects are a widespread and ubiquitous ecological phenomenon, yet they remain poorly studied across edges between restored and natural forests. To address this lack of knowledge, we studied vertebrate communities across edges between 3‐year old restored mine‐pits and adjacent unmined forest in the jarrah (Eucalyptus marginata) forest of south‐western Australia. We found that mammal communities showed no edge response but reptile communities did. Overall reptile abundance and Morethia obscura abundance were higher in unmined forest along edges, Egernia napoleonis abundance was lower in unmined forest along edges, while Pogona minor abundance was lower in restored mine‐pits along edges. Predictive models were unable to predict species edge responses, due to the lack of knowledge of the ecology of jarrah forest reptiles, but proved useful in identifying potential ecological mechanisms behind observed edge responses and suggested that potential mechanisms were likely different for each species. Our study is the first to show edge responses in both habitats forming the edge between restored and natural forests, emphasizing the importance of studying both habitats forming the edge. Our results also suggest that, despite being poorly studied, edge responses are common across edges between restored and natural forest and result from a variety of ecological mechanisms. An increased understanding of the ecological mechanisms driving edge responses across edges between restored and natural forests will improve our ability to integrate restored areas into cross‐landscape management and, ultimately, improve our ability to manage landscapes for biodiversity conservation.     

Landscape-Scale Implications of the Edge Effect on Soil Fauna Activity in a Temperate Forest

Although studies on edge effects on species richness and abundance are numerous, the responses of ecosystem processes to these effects have received considerably less attention. How ecosystem processes respond to edge effects is particularly important in temperate forests, where small fragments and edge habitats form a considerable proportion of the total forest area. Soil fauna are key contributors to decomposition and soil biogeochemical cycling processes. Using the bait lamina technique, we quantified soil fauna feeding activity, and its dependence on soil moisture and distance to the edge in a broad-leaved forest in Southern England. Feeding activity was 40% lower at the forest edge than in the interior, and the depth of edge influence was approximately 75 m. A watering treatment showed that moisture limitation was the main driver of the reduced feeding activity at the edge. In England, only 33% of the forest area is greater than 75 m from the edge. Therefore, assuming that the results from this single-site study are representative for the landscape, it implies that only one- third of the forest area in England supports activity levels typical for the forest core, and that edge effects reduce the mean feeding activity across the landscape by 17% (with lower and upper 90% confidence intervals of 1.3 and 23%, respectively). Changing climatic conditions, such as summer droughts may exacerbate such effects as edges lose water faster than the forest interior. The results highlight the importance of taking edge effects into account in ecological studies and forest management planning in highly fragmented landscapes.     

Forest–farm edge effects on communities of ground beetles (Coleoptera: Carabidae) under different landscape structures

In fragmented landscapes, ecological processes may be significantly influenced by edge effects, but few data are available for edge effects across forest–farmland edges. We investigated patterns of species richness, abundance, and species composition in ground beetles across forest–farm edges in two different agro-forest landscapes in Korea. Nine and five sites were selected from Hwaseong, a fragmented landscape, in 2011 and 2012, respectively, while eight sites were selected from Hoengseong, a relatively well-protected landscape, in 2012. Ground beetles were collected by pitfall trapping. Species richness was higher in the surrounding habitat than in the forest interior or edge in both Hwaseong and Hoengseong. However, in Hwaseong, species richness of the forest edge was similar to that of the forest interior, while in Hoengseong forest edge species richness was intermediate between that of the forest interior and surrounding areas. In addition, non-metric multidimensional scaling based on the combined data of both locations showed that the species composition of ground beetles in the forest edge was more similar to that of the forest interior than the surrounding areas, although some open-habitat species occurred at the forest edges. Three characteristic groups (forest specialists, edge-associated species, and open-habitat species) of ground beetle species were detected by indicator value analysis. In our study, ground beetle assemblages differed in the forest edges of two agro-forest landscapes, suggesting that the edge effect on biota can be influenced by landscape structure.     

How do slow‐growing,fire‐sensitive conifers survive in flammable eucalypt woodlands?

Question: To what extent do low flammability fuel traits enhance the survival and persistence of fire‐sensitive (slowing‐growing, non‐serotinous, non‐resprouting) dominant trees in highly flammable landscapes, under varying fire‐weather conditions? Location: Mixed forests co‐dominated by flammable Eucalyptus species and fire‐sensitive Callitris glaucophylla in Pilliga State Forest, southeast Australia. Methods: The influence of vegetation composition (relative abundance of Callitris and flammable Eucalyptus) on fire intensity and survival of fire‐sensitive Callitris was assessed across gradients of Callitris abundance in mixed Eucalyptus–Callitris forests that burned under low‐moderate and extreme fire‐weather conditions. Results: In areas that burned under low‐moderate fire‐weather conditions, as Callitris abundance increased, fire intensity declined and Callitris survival increased (46%). By comparison, in extreme fire‐weather conditions, lower fire intensity at higher levels of Callitris abundance, was not sufficient to increase Callitris survival (4%). Callitris survival was also positively related to trunk diameter. Ground fuel type, but not biomass, varied with vegetation composition. Conclusions: These results demonstrate that flammable feedbacks, mediated by low flammability fuel traits of dominant trees, can provide an important mechanism for enhancing the survival and persistence of slow‐growing, non‐serotinous, non‐resprouting, fire‐killed trees in highly flammable landscapes. By modifying vegetation and fuel structure, patches of fire‐sensitive Callitris reduce fire intensity, and thereby reduce Callitris mortality, enhancing population persistence. However, this feedback loop is insufficient to ensure Callitris survival under extreme fire‐weather conditions, when fire intensity is greater. After burning, stands remain vulnerable to future fires, until trees grow large enough to modify fuel levels and reduce stand flammability.     

Bats in a changing landscape: Linking occupancy and traits of a diverse montane bat community to fire regime

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Edge influence on plant litter biomass in forest and savanna in the Brazilian cerrado

Edge influence, characterized by differences in ecosystem characteristics between the edge and the interior of remnants in fragmented landscapes, affects a variety of organisms and ecosystem processes. An important feature that may be affected by edges is the amount of plant litter, which provides important habitat for a large variety of organisms and influences ecological processes such as fire dynamics. We studied edge influence on plant litter and fine woody debris in the cerrado of São Paulo state, south‐eastern Brazil. We collected, sorted, dried and weighed plant litter along 180 m‐long transects perpendicular to three savanna and eleven forest edges adjacent to different anthropogenic land uses, with four to five transect per edge. There tended to be less biomass of the finer portions of fine woody debris at both savanna and forest edges. Graminoid litter at savanna edges was greater than in the corresponding interior areas, whereas other litter portions were either unaffected by edges or did not show consistent patterns in either savanna or forest. Edge influence was usually restricted to the first 20 m from the edge, was not influenced by edge characteristics and exhibited no clear differences between savanna and forest areas. Several mechanisms may have led to the variable patterns observed including variation in the plant community, plant architecture, and invasive species. The edge‐related variation in plant litter may putatively lead to, for example, increased fire frequency and intensity at the savanna edges and altered trophic dynamics at forest edges; the mechanisms and consequences of this edge influence should be addressed in future studies.     

Conserving butterflies in fragmented plantation forests: are edge and interior habitats equally important?

Edge effects are increasing in forest-dominated landscapes worldwide, due to increased fragmentation by other land uses. Understanding how species respond to edges is therefore critical to define adequate conservation measures. We compared the relative importance of interior and edge habitats for butterflies in a landscape composed of even-aged pine plantations interspersed with semi-natural habitats. Butterfly assemblages were surveyed simultaneously at the edge and the interior of 68 patches belonging to four main habitat types: herbaceous firebreaks, clearcuts and young pine stands, older pine stands, and deciduous woodlands. Butterfly species richness was higher at edges than in interior habitats, especially for pine stands. Assemblage composition differed significantly between edge and interior habitats, except for firebreaks. Of the 23 most abundant butterfly species, seven were significantly more abundant in one or all edge habitat types, five in interior habitats, and 11 species showed no edge-interior preference. Modelling the presence of individual species in edge habitats revealed the importance of habitat variables such as the abundance of nectar and host-plants, but also of the abundance of the same species in the adjacent interior habitat. Moreover, our results suggest that most species use several, different habitat types to find supplementary or complementary resources, including micro-climatic refuges to escape hot temperatures during summer. The use of adjacent edge and interior habitats by butterflies is probably a key process in such mosaic landscapes and underlines the importance of landscape heterogeneity for butterfly conservation.     

Macrofaunal Responses to Edges Are Independent of Habitat-Heterogeneity in Experimental Landscapes

Despite edges being common features of many natural habitats, there is little general understanding of the ways assemblages respond to them. Every edge between two contrasting habitats has characteristics governed by the composition of adjoining habitats and/or by the nature of any transitions between them. To develop better explanatory theory, we examined the extent to which edges act independently of the composition of the surrounding landscape and to which transitions between different types of habitats affect assemblages. Using experimental landscapes, we measured the responses of assemblages of marine molluscs colonising different experimental landscapes constructed with different compositions (i.e. different types of habitats within the landscape) and different types of transitions between habitats (i.e. sharp vs gradual). Edge effects (i.e. proximity to the edge of the landscape) were independent of the internal composition of experimental landscape; fewer species were found near the edges of landscapes. These reductions may be explained by differences in differential larval settlement between edges and interiors of experimental landscapes. We also found that the sharpness of transitions influenced the magnitude of interactions in the different types of habitats in experimental landscapes, most probably due to the increased number of species in areas of transition between two habitats. Our experiments allowed the effects of composition and transitions between habitats to be disentangled from those of proximity to edges of landscapes. Understanding and making predictions about the responses by species to edges depends on understanding not only the nature of transitions across boundaries, but also the landscape in which the edges are embedded.     

Spatio-temporal variation in species assemblages in field edges: seasonally distinct responses of solitary bees to local habitat characteristics and landscape conditions

The bees (Hymenoptera: Apiformes) are important pollinators in many ecosystems, but their diversity has declined in Europe during the past century, mainly due to habitat loss. However, some of the habitat requirements of wild bees are met in anthropogenic landscape elements, such as road sides, power-line strips and field edges. Moreover, as the bee species assemblages change throughout the season the habitat requirements of the bee fauna may change accordingly. Understanding such seasonally distinct responses of solitary bees with different phenologies may be of high value for local conservation planning. The purpose of this study was to examine if the habitat quality of field edges for solitary bees change throughout the season, and how this temporal variation relates to local habitat and landscape conditions. By sampling solitary bees in 18 field edges in southeast Norway throughout the season we found that the species richness and abundance of bees was highest in sun exposed field edges, independently of the season. However, we found phenologically distinct responses to the landscape context. Moreover, field edges situated in landscapes with a high proportion of forests and semi-natural landscape elements hosted the most phenologically diverse bee species assemblages. We conclude that in order to fulfil the habitat requirements of bee species assemblages throughout the season, one should conserve and direct habitat restoration schemes towards increasing sun exposure at field edges with a diverse flora and a high proportion of semi-natural areas in the vicinity.     

Conserving long unburnt vegetation is important for bird species,guilds and diversity

Landscape-level wildfires have a major role in structuring faunal assemblages, particularly in fire-prone landscapes. These effects are mediated by changes to vegetation structure and composition that directly influence the availability of shelter, feeding and breeding resources. We investigated the response of a semi-arid shrubland bird community in Western Australia to the prevailing fire regime by examining the abundance, diversity and guild structure in relation to time since fire. We also examined vegetation structural attributes in relation to time since fire. We surveyed 32 sites ranging in age from 12 to 84 years since last fire. A total of 845 birds from 40 species were recorded. Vegetation structure varied with fire history with old and very old sites characterised by less bare ground, more leaf litter cover and greater canopy cover. Bird community composition varied with time since fire, driven by increased bird species richness and abundance of insectivores, granivores/frugivores, golden whistlers, grey shrike-thrush and red-capped robins with time since fire. Frequent, intense landscape-scale fires transform the landscape into homogeneous young shrublands, which may render vegetation unsuitable for several species and guilds.     

Urbanization Impacts on Mammals across Urban-Forest Edges and a Predictive Model of Edge Effects

With accelerating rates of urbanization worldwide, a better understanding of ecological processes at the wildland-urban interface is critical to conserve biodiversity. We explored the effects of high and low-density housing developments on forest-dwelling mammals. Based on habitat characteristics, we expected a gradual decline in species abundance across forest-urban edges and an increased decline rate in higher contrast edges. We surveyed arboreal mammals in sites of high and low housing density along 600 m transects that spanned urban areas and areas turn on adjacent native forest. We also surveyed forest controls to test whether edge effects extended beyond our edge transects. We fitted models describing richness, total abundance and individual species abundance. Low-density housing developments provided suitable habitat for most arboreal mammals. In contrast, high-density housing developments had lower species richness, total abundance and individual species abundance, but supported the highest abundances of an urban adapter (Trichosurus vulpecula). We did not find the predicted gradual decline in species abundance. Of four species analysed, three exhibited no response to the proximity of urban boundaries, but spilled over into adjacent urban habitat to differing extents. One species (Petaurus australis) had an extended negative response to urban boundaries, suggesting that urban development has impacts beyond 300 m into adjacent forest. Our empirical work demonstrates that high-density housing developments have negative effects on both community and species level responses, except for one urban adapter. We developed a new predictive model of edge effects based on our results and the literature. To predict animal responses across edges, our framework integrates for first time: (1) habitat quality/preference, (2) species response with the proximity to the adjacent habitat, and (3) spillover extent/sensitivity to adjacent habitat boundaries. This framework will allow scientists, managers and planners better understand and predict both species responses across edges and impacts of development in mosaic landscapes.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.