Landscape history and soil properties affect grassland decline and plant species richness in rural landscapes

Past semi-natural grassland extent is thought to have a major influence on contemporary species richness in rural landscapes. The loss of grasslands over the last 300 years was reconstructed for 12 rural landscapes in Sweden, ranging from open modern agricultural landscapes to more forested landscapes. Old maps and aerial photographs from 1950s and today were used to reconstruct landscape patterns in four time-steps to investigate how present plant species richness relates to past grassland extent and decline in old and new grassland habitats. The relative importance of soil properties on the timing of grassland decline was assessed. Plant species occurrence was recorded in managed and abandoned grassland habitats in each landscape.Past and present grassland distribution was a major factor in determining plant species patterns found in grasslands today. All landscapes had an average of 80% grassland 300 years ago. Since then grassland has declined by 90% across all landscapes. Proportion of clay soils influenced the timing of grassland decline, where grasslands in landscapes dominated by clay soils were conversed to crop-fields more than 100 years ago. Grasslands on coarser soils declined later, primarily to forest. Landscapes with more than 10% semi-natural grassland left today had 50% higher species richness in all grasslands, including both abandoned and new grassland. Time since major grassland decline also seems to have an effect on the landscapes’ species richness. The results show that plant species patterns in grasslands at local scales are determined by broader landscape processes which may have occurred many centuries ago.     

Changes in Collembola richness and diversity along a gradient of land-use intensity: A pan European study

Changes in Collembola richness and diversity along a land-use intensity gradient were studied in eight European countries (Portugal, Spain, France, Switzerland, Hungary, UK, Ireland and Finland). In each country a set of six 1 km² land-use units (LUUs) were selected forming a gradient ranging from natural forest to agricultural dominated landscapes, passing through mixed-use ones. In addition to data on Collembola, detailed information regarding landscape diversity and structure was collected for each LUU. A total of 47,774 individuals were identified from 281 species. Collembola reacted not only to changes in the diversity of the landscape, but also to the composition of that diversity and the area occupied by each land-use type at each LUU. Although species richness patterns were not concordant among the different countries, the total number of species per LUU (landscape richness) was generally higher in natural forests and mixed-used landscapes, and lower in agricultural dominated landscapes. Moreover, high richness and diversity of Collembola at each LUU were associated with a diverse landscape structure, both in terms of number of patches and patch richness. Despite this comparable species richness between mixed-use landscapes and those dominated by natural forests, average species richness on forested areas (local richness) decreased along the gradient, showing that forest patches on mixed-use landscapes support a lower richness than in landscapes dominated by forest. This aspect is important when addressing the role of native forests in structuring biodiversity in disturbed and fragmented landscapes. Although a diverse landscape can support a high biodiversity, the results suggest that intensive fragmentation should be avoided with the risk of collapsing local species richness with the consequent result for regional biodiversity.     

The influence of field shape, area and surrounding landscape on plant species richness in grazed ex-fields

Over the past 100 years species-rich semi-natural grasslands have decreased dramatically in Western Europe, where former arable fields (ex-fields) are used instead as pasture. The disappearance of semi-natural grasslands has caused a threat to the biodiversity in agricultural landscapes. Many typical grassland plants are dispersal limited, thus grazed ex-fields can be used to investigate if species spontaneously colonise these new grassland habitats. We examined the relationship between surrounding landscape, field area, shape, distance between edge and centre, and plant species diversity in ex-fields that had been grazed for 15-18 years. The results showed that there were 35% more plant species in fields surrounded by commercial forestry production compared to those surrounded by open agricultural landscape. Area and shape did not influence species richness, although there was increasing number of species in the centre with decreasing distance from the edge. Twenty-five percent of the species where typical grassland species, and ex-fields surrounded by forest had 91% more grassland species compared to those in the open landscape. It is possible to increase grassland plant occurrences by grazing ex-fields surrounded by forest or other grassland remnant habitats, particularly in landscapes where grazed semi-natural grasslands are scarce.     

Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment

Calcareous grasslands, most of which are man made and therefore depend on some kind of human interference for their maintenance, are among the most species-rich communities on Earth at a small scale. For many centuries, most of these grasslands have been used as extensive pasture. However, after 1900, and particularly from 1940 onwards, livestock grazing has declined throughout Europe leading to the abandonment of low intensity grasslands over large areas. To conserve the remaining grasslands or to restore recently abandoned grasslands, better insights about the effects of grassland management on above and belowground species diversity are needed. Here, we describe the results of an 11-year experiment to investigate the role of grassland management (grazing, mowing and abandonment) in determining species composition and diversity both in the aboveground vegetation and the seed bank of a calcareous grassland in Belgium. Species diversity declined by about 60% 11 years after abandonment, from 29 species m⁻² to as few as 12 species m⁻². Plots that were grazed remained constant in species richness, whereas mown sites lost about 20% of their original species. Abandoned plots were largely dominated by a few grass species, in particular Festuca rubra. Concomitant with changes in the aboveground vegetation, both the number of species found in the seed bank and seed density (number of seeds m⁻²) had changed significantly 11 years after abandonment. Species diversity and seed density were significantly lower in abandoned plots than in grazed or mown plots. We conclude that abandonment of calcareous grasslands may lead to rapid decline of plant species diversity both in the aboveground vegetation and in the seed bank. As a result, seed banks probably have a limited role to play in the restoration of recently abandoned grasslands.     

Remnant grassland habitats as source communities for plant diversification in agricultural landscapes

Lately there has been a shift in Sweden from grazing species-rich semi-natural grasslands towards grazing ex-arable fields in the modern agricultural landscape. These fields normally contain a fraction of the plant species richness compared to semi-natural grasslands. However, small remnant habitats have been suggested as important for plant species diversity and conservation as they may function as refugia for grassland specialists in fragmented and highly modified agricultural landscapes. In this study, we examined whether plant communities on small remnant habitats, i.e. midfield islets, can function as sources for grassland species to disperse out into surrounding grazed ex-fields (former arable fields). We examined species richness and grassland specialists (species favoured by grazing) and their ability to colonize fields after 5 and 11 years of grazing. The fields that had been grazed for a shorter time were fairly species-poor with few grassland specialists. A longer period of grazing had a positive effect on total and small-scale species diversity in both islets and fields. Species composition became more similar with time, and the number of grassland specialists in both habitats increased. We found that grassland specialists dispersed step-wise into the fields, and the number of grassland specialists decreased with distance from the source. Our study suggests that remnant habitats, such as midfield islets, do function as a source community for grassland specialists and enhance diversification of grassland species when grazing is introduced. For long-term conservation of plant species, incorporating small refugia into larger grazing complexes may thus enhance species richness.     

Land use and habitat gradients determine bird community diversity and abundance in suburban, rural and reserve landscapes of Minnesota, USA

Bird species’ community responses to land use in the suburbanizing Twin Cities, Minnesota, USA, were contrasted among reserves, rural lands, and suburbs. For each land use type, bird composition, diversity, and abundance were recorded for 2 years in ≈99 plots in three sampling units (each ≈4500 ha). A habitat gradient defined by canopy structure (grasslands to savannas to forests) was influenced by land use, so ≈300 plots were used to characterize simultaneous variation in bird communities along land use and habitat gradients. At broad scales (aggregate of 33 plots covering ≈4500 ha) suburbs supported the lowest bird richness and diversity and rural landscapes the most, with reserves slightly below rural. Although reserves were like rural lands in diversity of bird communities, they supported more species of conservation concern, particularly of grasslands and savannas. Differences among land use types varied with habitat structure. Suburbs, rural lands, and reserves had similar forest bird communities, but differed in grassland and savanna bird communities. The extensive rural forests are important for the region’s forest birds. Suburban grasslands and savannas had low shrub abundance, low native bird richness and high non-native bird richness and abundance. However, total bird richness and diversity were as high in suburban as in rural and reserve plots because high native richness in suburban forests and high non-native species richness in suburban grasslands and savannas compensated for lower native richness in suburban grasslands and savannas. Bird conservation here and in the Midwest USA should protect rural forests, expand grasslands and savannas in reserves, and improve habitat quality overall.     

Indirect evidence for an extinction debt of grassland butterflies half century after habitat loss

Habitat loss often leads to a substantial decline in species richness. However, the extinction of species is typically not instant, but rather involves a time lag. Species richness in recently disturbed habitats is therefore expected to reflect past rather than current habitat availability, with the set of species eventually going extinct representing extinction debt. We explored current species richness of butterflies (Lepidoptera: Rhopalocera) and burnet moths (Lepidoptera: Zygaenidae) with respect to past and current habitat characteristics. The study was conducted in calcareous grasslands, a habitat that has suffered a rapid decline in Europe during the last century. We showed that species richness of habitat specialists correlated positively with both past (75 years ago) and current habitat area, and was best explained by a model incorporating both of these variables. The independent effect of past habitat area presumably indicates the presence of extinction debt in the study region. Species requiring large habitat areas were more likely to show signs of extinction debt. Species richness of other grassland butterflies (those not confined to the focal habitat type) was not associated with past or current habitat area, the pattern indicating their lower sensitivity to changes in focal habitats. The response to habitat loss in specialist butterflies appears faster than in specialist plants, studied in the same landscape earlier. We conclude that linking patterns of species diversity with the temporal dynamics of habitats substantially improves the understanding of the variation in species diversity, and thus helps to identify species of conservation concern.     

Landscape elements as potential barriers and corridors for bees, wasps and parasitoids

Habitat loss and fragmentation in agricultural landscapes lead to severe declines of abundance and richness of many insect species in the remaining isolated semi-natural habitats. We analysed possible barrier effects of large hedges and corridor effects of narrow grass strips that were hypothesized to affect foraging and dispersal of hymenopterans. We selected calcareous grasslands in the vicinity of Göttingen (Germany), which harbour high Hymenoptera diversity and are starting points for foraging and dispersal in the landscape. We installed pan traps to sample bees (i) on the grasslands; (ii) on grassland edges behind adjacent hedges (potential barriers) and without hedges; (iii) on grass strips in 100 m distance to the grassland, which were connected or unconnected to the grassland; and (iv) unconnected (isolated) grass strips in 300 m and 750 m distance to test for corridor and isolation effects on abundance and species richness of foraging wild bees. Additionally we provided trap nests for bees, wasps and their parasitoids on the grasslands and the strips. Species abundance and richness declined with increasing isolation from grasslands for foraging solitary bees, trap-nesting bees, wasps and parasitoids, but not for foraging bumblebees. Hedges did not confine movement of foraging bees. We found no mitigating effects of (100 m) corridor strips on any of the observed groups. We conclude that conservation of semi-natural habitats as sources of bee and wasp diversity is important and that grass strips act as sinks rather than corridors when high quality patches are nearby.     

东北平原典型玉米种植区农业景观植物多样性研究——以昌图县为例

非耕作生境是农业景观中的重要组成部分,为生物提供食物和栖息地,对生物多样性保护具有重要意义。本文以辽宁省昌图县为例,选取了20个非耕作生境比例为0~50%的网格(1 km?1 km)为样区,采用典型样地法对调查区域内农业景观中的林地、田间路、草地、沟渠、果园5类主要非耕作生境中的植物群落进行调查,探讨不同非耕作生境类型和比例中植物多样性及其与景观异质性之间的关系。研究结果表明,随着非耕作生境比例增加,植物物种多样性和均匀度先增加后减小,在20%~30%时达到峰值;非耕作生境比例在0~40%范围内时,植物物种丰富度同样呈现先增加后减小,且在20%~30%时达到峰值;非耕作生境比例高于40%时,植物物种丰富度再次出现峰值且高于20%~30%。随着非耕作生境比例的增加,农业景观香浓多样性、香浓均匀度指数和景观斑块形状复杂程度逐渐增加,蔓延度恰好相反。不同生境中重要值最高的均为禾本科植物,但植物种不同;优势种优势度为果园沟渠草地田间路林地。果园、草地和沟渠的物种多样性和均匀度指数显著高于林地和田间路,丰富度指数为草地林地沟渠果园田间路,且草地、果园和田间路的不同调查区域间植物群落指数存在较大的变异性。草地、果园、沟渠、田间路和林地中景观异质性与物种多样性相关性顺次降低。研究结论表明,东北平原典型玉米种植区农业景观中,最适宜非耕作生境比例为20%~30%;非耕作生境中林地、沟渠对物种多样性维持、湿生植物多样性保护具有积极作用;不同生境干扰类型和方式不同,景观异质性与植物物种多样性相关性程度也不同。在今后农业景观生物多样性保护中应综合考虑非耕作生境的类型和比例,干扰的类型和方式等多种因素对生物多样性、生态服务及农业产量的影响。     

Butterfly distribution and abundance is affected by variation in the Swedish forest-farmland landscape

Remaining patches of semi-natural grasslands are hot spots for biodiversity in modern agricultural landscapes. In Sweden semi-natural pastures cover approximately 500,000 ha. However, power-line corridors, road verges and clear-cuts cover larger areas (in total about 2,000,000 ha), and these open, less intensively managed habitats are potentially important for species associated with taller vegetation and flower resources (e.g. pollinating insects). The aim of the present study was to evaluate the relative importance of semi-natural pastures and the other three open habitats for butterflies in 12 forest-farmland mosaic landscapes in south central Sweden. Species composition differed significantly between habitats in multivariate analyses. Power-line corridors and semi-natural pastures harbored several species that were disproportionally abundant in these habitats (13 and 8 species, respectively), and power-line corridors also harbored several species that were classified as typical in indicator species analyses. There were more butterfly species, higher abundances and a tendency for more individuals of red-listed species in power-line corridors than in the other three habitats. Effects of the surrounding landscape composition seemed to be weaker than that of the local habitat. However, species composition was significantly associated with landscape composition and species with intermediate and low mobility were more abundant in forested landscapes than in landscapes dominated by arable fields. Analyses of flying time and host plants for larvae suggest that early flying species and species associated with dwarf shrubs were more common in power-line corridors than in the other habitats. A landscape perspective, which takes several habitats into account, is needed for conservation of butterfly communities in forest-farmland landscapes. Power-line corridors and road verges offer possibilities for creating habitats that are suitable for pollinating insects through conservation-oriented management.     

The importance of the gravel excavation industry for the conservation of grassland butterflies

Conservation biology often relies on the protection of (semi)natural habitat remnants. However, the ever increasing human population is taking over natural resources and habitats. Here, contrary to most other studies, we ask how human-associated severe changes in the environment can be used to enrich local biodiversity. We tested if industrial activity (gravel excavation) leads to the creation of habitats that support grassland butterflies and how these areas add to the richness of local species when compared to typical semi-natural habitats (grasslands). We also identified key factors affecting the richness, abundance, diversity and commonness of butterfly species to provide practical recommendations. Species richness, diversity index and the occurrence of rare species were higher in gravel-pit shores than in grasslands. The richness of butterfly species and their abundance were positively affected by the richness of plant species, shrub density and age of the gravel-pit but negatively by the cover of water reservoirs in the surrounding area and the isolation of gravel-pits from grasslands. Butterfly diversity was positively influenced by the richness of plant species and proximity of human settlement but negatively by area of the shore and isolation. Our study is the first one to show the high value of gravel-pits for the conservation of butterflies. We recommend the inclusion of gravel-pits in a system of ecological networks and management of their surroundings to improve the colonization rate of rare species. We suggests that directing interest to the possible positive effects of industrial development on biodiversity may support conservation efforts.     

Forest fuel management as a conservation tool for early successional species under agricultural abandonment: The case of Mediterranean butterflies

In cultural landscapes there are often negative biodiversity consequences of agricultural abandonment and subsequent scrub and forest encroachment, due to homogenization and the loss of early-successional habitats. The common forestry practice of removing understory vegetation to prevent fire hazard (fuel management) probably has the side-effect of ameliorating these consequences, but it is uncertain whether it effectively restores habitats for early-successional species. Here we examine the influence of time since fuel management and management frequency on butterfly assemblages, using a chronosequence of cork oak (Quercus suber) stands spanning about 70 years. Overall species richness increased immediately after management and abundances peaked about 2–3 years later, while both declined thereafter for about 10–20 years to pre-disturbance levels. Richness and abundances were also much higher in recurrently managed stands. Most life history groups showed successional trends similar to the overall species richness and abundances, though consistent positive effects of fuel management were only observed for species with univoltine life cycle, herbaceous layer feeding, larval overwintering, and intermediate body size. Individual species were largely associated with recent and recurrent management, though a few specialists occurred most often in undisturbed stands. These findings suggest that fuel management at <10 years intervals is strongly positive for butterfly assemblages in landscapes under land abandonment. However, to maintain the overall forest biodiversity it is critical that patches of undisturbed habitat are also retained at the landscape scale.     

Matrix effects on bird assemblages in fragmented Afromontane forests in South Africa

We examine the effects of matrix type on forest bird species richness in historically fragmented forests of the KwaZulu-Natal midlands, South Africa. Bird species-area relationships for forest fragments within natural grassland were compared with those surrounded by plantation forestry (Pinus spp.). While fragments in grasslands displayed a species-area effect, no such effect (slope ∼0) was detected for fragments in plantations. The critical fragment size to avoid an island effect on species richness was 302 ha for fragments in grasslands, and contained 51 of the 61 forest bird species encountered. Small forest fragments (<50 ha) within grasslands were less species-rich than those surrounded by plantation, while the reverse was true for forests larger than 50 ha. Bird density was significantly lower in larger forest patches within plantation when compared to those in grassland. However, a slight decline in species density with increasing species richness in a patch (i.e. weak density compensation) characterised only those bird assemblages surrounded by a grassland matrix, suggesting these are approaching species saturation. Commercial plantations may increase the likelihood of colonisation of, and immigration from, small forest fragments by birds, reducing the incidence of area-dependent extinction in small fragments, but may also result in lower species richness in larger fragments. The latter is likely a consequence of the promotion by plantations of a wider distribution of generalist species in, and the loss of some specialised or rare species from, large patches. Management options are to avoid planting plantations near large forest patches and to increase the size of small patches where possible. Where afforestation is unavoidable, placing plantations in the vicinity of small forest patches rather than large forest patches may be preferable.     

The feasibility of using soil seed bank for natural regeneration of degraded sandy grasslands

Desertification in degraded grasslands is manifested through the development of bare sandy patches, which eventually lead to habitat fragmentation. The ability of these bare sandy patches to regenerate naturally through in-situ soil seed banks is not well understood. To fill this knowledge gap, we randomly selected 24 bare sandy patches with areas ranging from 19 to 898 m² in a desertified grassland of the Horqin sandy land, Northern China to determine whether soil seed bank can be used for natural regeneration of bare sandy patches. Species composition and density of soil seed bank as well as aboveground vegetation composition, abundance and coverage were investigated. We then determined their relationships with in-situ habitat characteristics. Our observations showed that the studied area had low soil seed bank density and species richness, as well as depauperate soil seed bank communities. Consequently, local soil seed bank was not able to provide sufficient seed source for natural regeneration. This was indicated by the relationships between aboveground vegetation, soil seed bank and the in-situ habitat characteristics. For bare patches with an area between 300 m² and 900 m², increase the soil seed bank density and species richness should be the main restoration measures. For bare patches with a small area of less than 50 m², restoration of vegetation density should be the main measure. Our data highlighted that different extents of desertification, indicated by different bare patches, are requiring distinct restoration measures.     

Small-scale plant species richness and evenness in semi-natural grasslands respond differently to habitat fragmentation

The study explores whether small-scale species diversity, species evenness and species richness in semi-natural grassland communities are similarly associated with present management regime and/or present and historical landscape context (percentage of different land-cover types in the surroundings). Species diversity, evenness and richness were recorded within 441 50 × 50 cm grassland plots in 4.5 × 4.5 km agricultural landscape on Öland, Sweden. Recent and historical land-cover maps (years 2004, 1959, 1938, 1835, and 1800) were used to characterize the present and past landscape context of the sampled vegetation plots. Partial regression and simultaneous autoregressive models were used to explore the relationships between species diversity measures (Shannon diversity, richness and evenness) and different explanatory variables while accounting for spatial autocorrelation in the data. The results indicated that species richness was relatively sensitive to grassland isolation, while the response of species evenness to isolation was characterized by a degree of inertia. Because the richness and evenness components of species diversity may respond differently to habitat fragmentation, we suggest that monitoring projects and empirical studies that focus on changes in biodiversity in semi-natural grasslands should include the assessment of species evenness - as a complement to the assessment of species richness. In addition, our results indicated that the development and persistence of a species-rich and even grassland vegetation was favoured in areas that have historically (in the 19th century) been surrounded by grasslands. Information on landscape history should, whenever possible, be incorporated into the planning of strategies for grassland conservation.     

Old cultural traditions, in addition to land use and topography, are shaping plant diversity of grasslands in the Alps

Socio-economically motivated land use changes are a major threat for species diversity of grasslands throughout the world. Here, we comprehensively explore how plant species diversity of grasslands in the species-rich cultural landscape of the Swiss Alps depends on recent land use changes, and, neglected in previous studies, on old cultural traditions. We studied diversity in 216 grassland parcels at three altitudinal levels in 12 villages of three cultural traditions (Romanic, Germanic, and Walser). In valleys of Romanic villages more different parcel types tended to occur than in those of Germanic and Walser villages, suggesting that socio-economic differences among cultural traditions still play a role in shaping landscape diversity. Moreover, at the village level, higher man-made landscape diversity was associated with higher plant species richness. All observed changes in land use reduced the farmers’ workload. Plant species richness was lower in fertilized than in unfertilized parcels and in abandoned compared with used parcels. Grazing slightly reduced species richness compared with mowing among unfertilized parcels, while in fertilized parcels it had a positive influence. The highest species diversity was found in mown unfertilized subalpine grasslands. Nevertheless, moderate grazing of former meadows can be a valuable alternative to abandonment. We conclude that the ongoing changes in land use reduce plant species richness within parcels and at the landscape level. To preserve plant species diversity at the landscape level a high diversity of land use types has to be maintained.     

Effects of grassland abandonment, restoration and management on butterflies and vascular plants

When semi-natural pastures are abandoned, specialized grassland species are lost as a consequence of succession. As a counter-measure, previously abandoned grasslands may be restored by clearing shrubs and trees and re-introducing grazing livestock. In order to examine the effects of this type of habitat restoration, we compared species richness of plants and of specialized plants thought to be dependent on continuous management and species richness and abundance of butterflies and red-listed butterflies in 12 sets of matched continuously managed, abandoned and restored grassland in southern Sweden. We found no differences in species richness or abundance between the three grassland types. There were, however, some negative effects of abandonment. The number of management-dependent plants decreased with increasing cover of trees and shrubs, and in restored sites species richness of all groups decreased with increasing cover of trees and shrubs before restoration. Also the present management significantly affected both butterflies and plants. Species richness of both groups increased with increasing vegetation height and differed between sites depending on the species of grazers, with negative effects of sheep compared to cattle or horses. Our study indicates that for grassland management to be efficient, the restoration actions should mainly be directed towards sites where the post-abandonment succession has not proceeded too far.     

Plant species richness in midfield islets and road verges - The effect of landscape fragmentation

Small marginal habitats in the rural landscape may play an important role for plant species richness as refugias. Little is known how the surrounding landscape and landscape history influence these patterns. I analysed how plant species richness was affected by isolation, habitat area, past and present land use, and if landscape context matters. Plant species occurrence in two different types of small marginal habitats were analysed, road verges and midfield islets. The study was conducted in two different agricultural landscapes in Sweden; one open modern agricultural landscape and one traditional rural landscape, and the results compared. Present and past land use, and landscape change was analysed using aerial photographs and old maps. There was a large grassland reduction more than 50 years ago in the modern landscape, when there still were quite a lot of grasslands left in the traditional landscape. Area and connectivity were more important for plant incidence in small remnant habitats in the modern landscape, compared to the less fragmented, traditional rural landscape. On the other hand there were more grassland specialists, 23% in the traditional landscape compared to 16%. Species richness became higher on midfield islet if grazing was re-introduced. The legacy of surrounding landscape remains in the species pool for a long time, atleast 50 years, even in small grassland fragments. Although small grassland remnants are more sensitive to fragmentation effects compared to larger grasslands, they still encompass a substantial part of the grassland species pool and may be valuable for reconstructing grassland management at a landscape scale.     

Long-term consequences of habitat fragmentation—highland birds in Oaxaca, Mexico

Studies of habitat fragmentation have been restricted primarily to anthropogenically-altered habitats, with most research conducted 60-90 years post-fragmentation. It is unclear whether patterns in older systems concur with results from these dynamic landscapes, and hence the long-term viability of populations inhabiting habitat fragments remains largely unexplored. I focused on resident birds in fragments of humid pine-oak forest in Oaxaca, southern Mexico, isolated over 5000 years ago by climate-change. Seventeen fragments, ranging from 2 ha to over 150,000 ha were sampled in 1997 and 1998 yielding 141 species, of which 60 residents were used for analysis. Avian assemblages exhibited a highly nested structure and, with several notable exceptions, assemblages of birds in low-richness fragments were predictable subsets of those in more diverse fragments. Patch-scale factors—area, shape, elevation, habitat diversity and fractal dimension of edge—all exerted strong univariate influence on avian richness but were so closely inter-related that none had a significant independent effect. Thus, larger fragments were more complex in shape, included higher peaks, supported more diverse forests, and contained higher diversities of resident species. In contrast, the landscape-scale index used—distance from nearest large fragment (>50,000 ha)—had little effect on richness. This was reinforced by species-level analyses—one species was significantly influenced by isolation, compared with 31 species that displayed significant minimum-area distributions, restricted to patches larger than a particular threshold value. In terms of autecology, vagility, relative abundance and elevational breadth were closely related to distribution—those species with greater mobility, higher abundances and broader elevational tolerances were consistently more widespread. I suggest that more abundant species were less prone to extinction initially, more vagile species were better dispersers and species with broader elevational tolerances more likely to be successful colonists. As with previous research from older landscapes, patch-scale factors were consistently found to be influential, with high quality fragments supporting diverse communities regardless of landscape context. This suggests that the influence of landscape-scale factors noted in younger, anthropogenically fragmented systems may be transitory, overwhelmed by patch-scale factors with time. Which patch attributes are most influential could not be resolved, however, indicating that even thousands of years after fragmentation, they affect diversity patterns in concert. Rather than differentiating effects of area from habitat heterogeneity and other patch-level factors, I advocate resource-based approaches to understand and manage diversity in habitat fragments.     

Use of remnant boreal forest habitats by saproxylic beetle assemblages in even-aged managed landscapes

Saproxylic Coleoptera are diverse insects that depend on dead wood in some or all of their life stages. In even-aged boreal forest management, remnant habitats left as strips and patches contain most of the dead wood available in managed landscapes and are expected to act as refuges for mature forest species during the regeneration phase. However, use of remnant habitats by the saproxylic fauna has rarely been investigated. Our objective was to characterize the saproxylic beetle assemblages using clearcuts and forest remnants in western Québec, Canada, and to explore the effects of forest remnant stand characteristics on saproxylic beetle assemblages. We sampled both beetle adults and larvae, using Lindgren funnels and snag dissection, in five habitat locations (clearcuts, forest interiors of large patches, edges of large patches, small patches and cut-block separators) from three distinct landscapes. Adult saproxylic beetles (all feeding guilds combined) had significantly higher species richness and catch rates in small patches compared to forest interiors of large patches; the phloeophagous/xylophagous group had significantly higher species richness only. Small patches, cut-block separators and edges of large patches also had the highest snag density and basal area, increasing habitat for many saproxylic beetles. No significant differences in density of saproxylic larvae were found between habitat patches, but snag dissection nevertheless suggests that snags in forest remnants are used by comparable densities of insects. Saproxylic beetles appear to readily use habitat remnants in even-aged managed landscapes suggesting that forest remnants can insure the local persistence of these species, at least in the timeframe investigated in our study.