Edge responses are different in edges under natural versus anthropogenic influence: a meta‐analysis using ground beetles

Most edges are anthropogenic in origin, but are distinguishable by their maintaining processes (natural vs. continued anthropogenic interventions: forestry, agriculture, urbanization). We hypothesized that the dissimilar edge histories will be reflected in the diversity and assemblage composition of inhabitants. Testing this “history‐based edge effect” hypothesis, we evaluated published information on a common insect group, ground beetles (Coleoptera: Carabidae) in forest edges. A meta‐analysis showed that the diversity‐enhancing properties of edges significantly differed according to their history. Forest edges maintained by natural processes had significantly higher species richness than their interiors, while edges with continued anthropogenic influence did not. The filter function of edges was also essentially different depending on their history. For forest specialist species, edges maintained by natural processes were penetrable, allowing these species to move right through the edges, while edges still under anthropogenic interventions were impenetrable, preventing the dispersal of forest specialists out of the forest. For species inhabiting the surrounding matrix (open‐habitat and generalist species), edges created by forestry activities were penetrable, and such species also invaded the forest interior. However, natural forest edges constituted a barrier and prevented the invasion of matrix species into the forest interior. Preserving and protecting all edges maintained by natural processes, and preventing anthropogenic changes to their structure, composition, and characteristics are key factors to sustain biodiversity in forests. Moreover, the increasing presence of anthropogenic edges in a landscape is to be avoided, as they contribute to the loss of biodiversity. Simultaneously, edges under continued anthropogenic disturbance should be restored by increasing habitat heterogeneity.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

How important are large vs. small forest remnants for the conservation of the woodland flora in an urban context?

Aim To test whether, in an urban area, small forest fragments are more important than large ones, the present study aimed at firstly exploring the relationship between plant functional groups or individual species and urban woodlot characteristics such as patch area and isolation, and secondly investigating whether equal‐sized combinations of islands tend to differ in species richness. Location The city of Brussels. Methods We considered the relationship between size, species richness and plant functional groups among one very large (1666 ha) and 11 rather small (2–123 ha) woodlots. Results The largest woodlot harboured species missing in the smaller ones. The species‐area relationship plotted for these wooded patches fitted the semilog model very well. Twenty‐three species had a significantly higher frequency in the main forest. Only six species had a significantly higher frequency in smaller woodlots. The occurrence of species groups with high conservation value (e.g. ancient forest species, rare species) was higher in large patches. However, a SLOSS analysis showed that habitat subdivision appeared to be associated with increased species counts. A marginally nonsignificant effect of distance to the main forest became significant when matrix species were removed from the patch samples. Conclusions Although overall the data showed a higher conservation value for large woods, some plant functional groups (e.g. woodland species vs. ancient forest species) responded differently to fragmentation. This illustrates that, for conservation strategies, studies considering the biotic characteristics of remnants should focus on the species number of particular plant functional groups, especially those with high conservation value. Furthermore, matrix species should be removed from the analysis in order not to mask underlying patch size and distance effects.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

The importance of considering both taxonomic and habitat guild approaches in small mammal research

Studies on the effects of habitat fragmentation on small mammals often lead to confounding results as they only consider taxonomic groups in their analysis and neglect functional diversity of the communities. Here we describe the structure and composition of small mammal communities at 22 sites, ranging from 41 to 7035 ha, in a hyper‐fragmented landscape of an Amazonia‐Cerrado ecotone. Also, in considering a taxonomic and habitat guild approach, we report the effects of habitat structures and patch spatial attributes on richness, abundance and species composition. Small mammal richness reported in southern Amazonia (N = 23 species) is greater than most previous studies in the tropics. All rare small mammals captured in this study were forest interior species. Richness of forest interior species was positively related to larger patches, as shown by the species–area relationship. However, 52% of the small mammal species were in forest fragments smaller than 50 ha, highlighting the importance of preserving both large and small forest fragments in a landscape with accelerated habitat reduction. Richness of edge‐tolerant species was not associated with the tested variables, yet edge‐tolerant species were more abundant in degraded environments. Marsupials were positively associated with vertical habitat structures, while rodents were more strongly related to a ground‐level habitat structure. The landscape studied is extremely variable and has contributed to the difficulty in detecting clear patterns, particularly when considering only one approach. Because of the complementary outputs when analysing either taxonomic groups or habitat guilds, we recommend the use of multi‐taxa studies of different guilds to assist decision makers in designing conservation strategies and appropriate management of small mammal populations.     

Assemblage-level responses of phyllostomid bats to tropical forest fragmentation: land-bridge islands as a model system

Aim Working within a system of high structural contrast between fragments and the surrounding matrix, we assessed patterns of species loss and changes in species composition of phyllostomid bats on artificial land‐bridge islands relative to mainland assemblages, and evaluated the responses of bats to forest edges. We further examined the relative influence of local‐scale characteristics (e.g. vegetation structure, island area) versus landscape attributes (e.g. forest cover, patch density) and the importance of spatial scale in determining phyllostomid species richness and composition on islands. Location Islands in Gatún Lake and adjacent mainland peninsulas in the Barro Colorado Nature Monument, Panama. Methods Bats were sampled over a 2‐year period on 11 islands as well as at forest‐edge and interior sites on adjacent mainland, resulting in > 8400 captures. Results The islands harboured a less diverse and structurally simplified phyllostomid bat fauna. Islands far from the mainland were especially species‐poor. This decline in species richness was associated with compositional shifts towards assemblages strongly dominated by frugivores with good dispersal abilities. Members of other ensembles, most importantly gleaning animalivores, were much less common or absent. Although overall species composition was not significantly altered, species richness at continuous forest‐edge sites was significantly lower compared with that at interior sites. Distance from the mainland and amount of forest cover in the landscape were the best predictors of species richness and assemblage composition. Responses were scale‐dependent. At the local scale, species richness was independent of island area but was correlated positively with distance from the mainland. In contrast, area effects became more important at larger spatial scales, suggesting that many species use multiple fragments. Main conclusions Our results underline the conservation value of small habitat remnants, which, even when embedded in a hostile matrix, can support a relatively diverse bat fauna, provided that there is a low degree of patch isolation and spatial proximity to larger tracts of continuous forest. Although the results at the assemblage level were inconclusive, we demonstrate that certain bat species and ensembles, particularly gleaning animalivores, exhibit high edge‐sensitivity. Our results point to habitat loss rather than changes in landscape configuration as the main process after isolation underlying phyllostomid bat responses, suggesting that conservation efforts should focus on habitat preservation instead of trying to minimize fragmentation per se at the expense of habitat amount.     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.     

Composition and distribution of ground-dwelling beetles among oak fragments and surrounding pine plantations in a temperate forest of North China

In this study, we compared ground-dwelling beetle assemblages （Coleoptera） from a range of different oak fragments and surrounding conifer plantations to evaluate effects of forest size and surrounding matrix habitat in a temperate forest of north China. During 2000, beetles were sampled via pitfall traps within two large oak fragments （ca. 2.0-4.0 ha）, two small oak fragments （ca. 0.2-0.4 ha） and two surrounding matrices dom- inated by pine plantations （〉4 ha） in two sites of different aspects. Overall, no significantly negative effects from forest patch size and the surrounding matrix habitat were detected in total species number and abundance of ground-dwelling beetles. However, compared with small oak patches or pine plantations, more species were associated with an affinity for at least one large oak patch of the two aspects. Multivariate regression trees showed that the habitat type better determined the beetle assemblage structure than patch size and aspect, indicating a strong impact of the surrounding matrix. Linear mixed models indicated that species richness and abundance of all ground-dwelling beetles or beetle families showed different responses to the selected environmental variables. Our results suggest that more disturbed sites are significantly poorer in oak forest specialists, which are usually more abundant in large oak fragments and decrease in abundance or disappear in small fragments and surrounding matrix habitats. Thus, it is necessary to preserve a minimum size of forest patch to create conditions characteristic for forest interior, rather than the more difficult task of increasing habitat connectivity.     

Environmental filtering is the main assembly rule of ground beetles in the forest and its edge but not in the adjacent grassland

In a fragmented landscape,transitional zones between neighboring habitats are common,and our understanding of community organizational forces across such habitats is important.Edge studies are numerous,but the majority of them utilize information on species richness and abundance.Abundance and taxonomic diversity,however,provide little information on the functioning and phylogeny of the co-existing species.Combining the evaluation of their functional and phylogenetic relationships,we aimed to assess whether ground beetle assemblages are deterministically or stochastically structured along grassland-forest gradients.Our results showed different community assembly rules on opposite sides of the forest edge.In the grassland,co-occurring species were functionally and phylogenetically not different from the random null model,indicating a random assembly process.Contrary to this,at the forest edge and the interior,co-occurring species showed functional and phylogenetic clustering,thus environmental filtering was the likely process structuring carabid assemblages.Community assembly in the grassland was considerably affected by asymmetrical species flows (spillover)across the forest edge:more forest species penetrated into the grassland than open-habitat and generalist species entered into the forest.This asymmetrical species flow underlines the importance of the filter function of forest edges.As unfavorable,human-induced changes to the structure,composition and characteristics of forest edges may alter their filter function,edges have to be specifically considered during conservation management.     

Island biogeography of temporary wetland carabid beetle communities

Aim The study tests if island biogeography is applicable to invertebrate communities of habitat islands in the agricultural landscape that are not fragments of formerly larger habitats. Location Thirty temporary wetlands in the agricultural landscape of northeast Germany. Methods The composition and species richness of carabid beetle communities was analysed. Habitat area, isolation, the density of temporary wetlands in the landscape, land‐use intensity and the maximum duration of flooding were recorded as independent variables. Overall species richness and wetland species richness were studied in independent regression analyses. The community composition was analysed by means of a Canonical Correspondence Analysis (CCA). A partial CCA was used to analyse the effect of the distance to the edge of the field after removing impacts of other independent variables. Results The area of the habitats and various measures of isolation (mean distances = 81–240 m) did not influence species richness or wetland species richness. The community composition was mainly determined by the land‐use intensity, habitat area did not have significant effects, and the distance to the edge of the field was the only effective isolation parameter. Short‐winged species were more often affected by the distance to the edge of the field than full‐winged species. Main conclusion There is evidence that the distances between the wetlands do not provide an effective barrier to the species dispersal and, therefore, metapopulation structures including subpopulations of multiple temporary wetlands might counteract local area effects on subpopulations. Short‐winged species, however, might be more affected by isolation than full‐winged species. As carabid beetle community structure in most early successional habitats is similar, these results may be representative of many agricultural landscape habitats. Nature conservancy concepts that aim to increase habitat area and habitat connectivity have successfully been applied to fragmented late‐successional habitats. The present study indicates that such concepts do not necessarily result in higher diversity or larger populations in early successional habitats.     

Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

Effects of forest fragmentation on species richness and composition of ground beetles (Coleoptera: Carabidae and Brachinidae) in urban landscapes

To clarify the effects of forest fragmentation in urban landscapes on the abundance, species richness, dominance, and species composition of ground beetles (Coleoptera: Carabidae and Brachinidae), we compared the beetles collected in 12 pitfall traps from April to July and from September to November between three continuous suburban forests and eight isolated urban forests (0.06–1.02 ha), most of which were in the precincts of shrines and temples in Hanshin District, Honshu, Japan. A total of 28 species and 4178 individuals of ground beetles were collected. Segregation of urban forests from continuous suburban forests has changed the species composition and resulted in the loss of some large‐sized forest species and the addition of some non‐forest species. Simpson's index of dominance (λ) also increased in the urban forests. The richness of forest species markedly decreased with the reduction in forest area but not with the distance from continuous forests, although the species richness of non‐forest species did not change with them. Also, species composition changed only with forest area. These findings indicate that continuous forests do not necessary serve as a “mainland” for urban forest species and that every urban habitat, however small in size, acts as a temporary reservoir of species. In comparison with populations of small‐sized species, populations of large‐sized forest species appeared to decline more readily during forest fragmentation.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

Forest patch size and isolation as drivers of bird species richness in Maputaland,Mozambique

Aim To examine the response of forest‐dependent and generalist bird assemblages and feeding guilds to patch characteristics of forest fragments in the context of island biogeography and metapopulation theories in an area that supports high levels of species diversity and endemism. Location Dune, riverine, sand and swamp forest fragments in southern Mozambique’s Maputaland. Methods We recorded 20 forest‐dependent and 69 generalist bird species at 220 survey points in 30 forest fragments that ranged in size from 5 to 7432 ha and that were 100 to 5100 m apart. We used linear regressions to relate the number of species to forest fragment size, isolation, perimeter, fractal dimension, shape and core area for frugivores, insectivores, granivores, nectarivores, carnivores and omnivores separately for forest‐dependent and generalist bird assemblages. We tested for modality in occupancy frequency distributions of species in forest fragments and used the binary matrix temperature calculator to determine whether the assemblages had a nested structure separately for forest‐dependent and generalist bird species. Results The number of forest‐dependent and generalist bird species categorized into the feeding guilds varied independently from forest patch parameter values. Occupancy models showed a random distribution for forest‐dependent birds, and generalists had a unimodal distribution. Both the forest‐dependent and generalist bird assemblages had a non‐nested structure. The presence of eight rare forest‐dependent and four endemic bird species in 22 of the 30 fragments contributed to the non‐nested structure of these bird assemblages. Main conclusions Fragmented forests did not induce the expected responses of the forest bird assemblages in that the number of species was not explained by patch characteristics, nor could we find evidence for metapopulation dynamics. Non‐nestedness may be caused by the few rare and endemic species occupying forest fragments with a wide range of patch characteristics. Knowledge of the response of species to forest fragmentation can benefit conservation planning catering for rare and endemic species. Our study suggests that all forest fragments in the region may have to be included in a conservation network to complement existing landscape‐based plans to protect the region.     

Red deer (<Emphasis Type="Italic">Cervus elaphus</Emphasis> Linnaeus, 1758) increasing density effects on species assemblage of ground beetles (Coleoptera: Carabidae) in Alpine forests

Red deer (Cervus elaphus) increasing density may trigger several modifications at the ecosystem level. Positive and negative impact of red deer on the invertebrate fauna is known in northern Europe, whereas it is currently unknown in the Alps. In this paper, we tested the impact of red deer density on ground beetles’ (Coleoptera: Carabidae) richness and species distribution in forests of the central-eastern Italian Alps (Stelvio National Park). Carabid beetles were sampled using pitfall traps in nine forest sites with different density of red deer. Carabid species richness was related to elevation, habitat type and red deer density in summer and winter using generalised linear mixed models and canonical correspondence analysis. The high winter red deer density affected positively the carabid species richness. Overgrazing seemed to increase presence of generalist species of dried and unshaded soils and decreasing of specialised species typical of well-preserved Rhododendro-Vaccinietum cover. Accordingly, the presence of patches with different red deer density seems to contribute to maintaining high levels of carabid beetles’ richness, albeit overgrazing needs to be monitored because it is likely to cause a local decrease of specialised forest species.     

Edge effect of a pine plantation reduces dry grassland invertebrate species richness

Natural steppes in European agricultural landscapes are characterized by high biotic richness but are subject to fragmentation and associated edge effects. Edge effects on species richness were investigated at an ecotone from a pine plantation to a short-grass steppe in Eastern Austria for eleven invertebrate taxa differentiated into habitat guilds based on known live-history strategies of individual species (grassland species, forest species, generalist species), including Red-Listed and non-threatened grassland species. The large size of the studied grassland site provided an opportunity to test edge effects in the absence of confounding factors and to a gradient length of 208 m into the grassland habitat along a clear-cut border to a pine plantation. All sampling was done by pitfall trapping. Species richness of habitat guilds, but not total richness, was effectively explained by biotic variables reflecting the influence of shading in particular (i.e. soil temperature sums). Total species richness showed a bimodal response pattern, with increases towards the habitat edge and interior grassland habitat. Habitat guilds showed diverging responses to distances from the edge, but no saturation in species richness, with a continuum of edge effects across the entire distance of the grassland samples. Our findings contrast those of previous investigations based on samples taken from smaller patches and across shorter distances from the edge. Methodological and conservation implications are discussed.     

Understorey plant species richness and composition in metropolitan forest archipelagos: effects of forest size, adjacent land use and distance to the edge

Aim To address the relative role of adjacent land use, distance to forest edge, forest size and their interactions on understorey plant species richness and composition in perimetropolitan forests. Location The metropolitan area of Barcelona, north‐eastern Spain. Methods Twenty sampling sites were distributed in two forest size‐categories: small forest patches (8–90 ha) and large forest areas (> 18,000 ha). For each forest‐size category, five sites were placed adjacent to crops and five sites adjacent to urban areas. Vascular plant species were recorded and human frequentation was scored visually in 210 10 × 10 m plots placed at 10, 50 and 100 m from the forest edge, and additionally at 500 m in large forest areas. Plant species were grouped according to their ecology and rarity categories. A nonmetric multidimensional scaling (NMS) ordination was carried out to detect patterns of variation in species assemblage, and to explore the relationships between these patterns and the richness of the species groups and the studied factors. Factorial anovas were used to test the significance of the studied factors on the richness of species groups. Relationships between human frequentation and the studied variables were assessed through contingency tables. Results Forest‐size category was the main factor affecting synanthropic species (i.e. those thriving in man‐made or man‐disturbed habitats). Synanthropic species richness decreased with increasing distance from the forest edge and, when forests were adjacent to crops, it was higher in small forest patches than in large forest areas. Richness of rare forest species was lower in small forest patches than in large forest areas when forests were adjacent to urban areas. Richness of common forest species and of all forest species together were higher close to the forest edge than far from it when forests were adjacent to urban areas. Forests adjacent to urban areas were more likely to experience high human frequentation, particularly in those plots nearest to the forest edge. Main conclusions Forest‐size category and adjacent land use were the most important factors determining species richness and composition. The preservation of large forests adjacent to crops in peri‐urban areas is recommended, because they are less frequented by humans, are better buffered against the percolation of nonforest species and could favour the persistence of rare forest species.     

Bryophyte Species Diversity in Secondary Forests Dominated by the Introduced Species Spathodea campanulata Beauv. in Puerto Rico

The introduced tree species Spathodea campanulata (Bignoniaceae) forms novel forests in Puerto Rico, these having emerged after the abandonment of fields in the mid‐20th century and resulting in forests with a new species composition. We assessed bryophyte species richness in these novel forests and sought correlations with geological substrate, past land use, forest edge and patch area, forest structure, elevation, microhabitat diversity, tree species richness, and microclimatic conditions. Transects were established (edge and forest interior) in nine moist forest patches dominated by Spathodea in north‐central Puerto Rico. These Spathodea forest patches ranged from 0.6 to 9 ha. ANOVA, Chi‐square, correlation, and cluster analyses were used in data analyses. We found 57 bryophyte species. There was a significant difference in bryophyte richness among patches. Those on karst exhibited highest bryophyte richness due to microhabitat diversity, past land use, and shorter hydroperiods. Alluvial sites scored lowest in bryophyte species richness, and forest structure was important for bryophyte communities on these sites. Significant differences in temperature, relative humidity, and light intensity were observed between edge and forest interior. These appeared important for establishing bryophyte species cover but not richness and composition. Microhabitat diversity, patch area, and forest age were more related to bryophyte species richness than elevation, exposed edge, and tree species richness, regardless of geologic substrate. Collectively, Spathodea patches were similar to mature forests on the Island with respect to bryophyte species richness and composition. Novel Spathodea forests have conservation value due to their habitat suitability for bryophyte communities.     

Dung beetle (Coleoptera: Scarabaeinae) persistence in Amazonian forest fragments and adjacent pastures: biogeographic implications for alpha and beta diversity

The aim of this study is to analyze the effects of habitat loss and forest replacement by cattle pasture on the alpha and beta diversity, abundance, biomass and species composition of dung beetles with different dispersal ability. Dung beetles were captured in 19 forest fragments and neighbouring pastures. Forest fragment area ranged from 3.7 to 4825 ha and in this study were grouped into four categories: small, medium, large and control forest. A total of 35,048 dung beetles representing 101 species were collected. Forest fragments had the highest richness with 81 species, followed by pasture with 58 species. Replacement of forest by pasture reduced species richness; however, due to the proximity and connectivity of these areas with Cerrado patches, pastures also had high species richness, but species composition was independent of adjacent fragments. Small fragments had lower abundance and species richness than our other habitat categories, even pastures. Our results highlight that proximity and connectivity with Cerrado areas influenced the patterns of alpha and beta diversity of dung beetles in fragments and pastures. We highlight that the ability to cross the pasture matrix is a strong adaptive trait for species living in human-modified landscapes. Consequently, species with these abilities are less susceptible to the effects of forest fragmentation and local extinction. Our results reinforce the importance of considering the biogeographic location and distribution pattern of species in forest fragmentation studies.