Neutral communities in fragmented landscapes

Habitat fragmentation displays a crucial role in conservation biology. Despite this, little is known about the detailed ecological consequences of habitat fragmentation due to the scarce number of controlled experimental surveys. The species–area relationship, a fundamental concept in ecology, requires the understanding of the fragmentation effects in a long term perspective, which turns this task even harder. Here we address the spatial patterns of species distribution in fragmented landscapes, assuming a neutral community model. We study the species area relationship and how its shape changes as the landscape becomes more fragmented. Recent investigations, based on extensive computer simulation, have contributed to establish some definite conclusions in the study of non‐fragmented landscapes: the existence of a three‐regime or two‐regime scenario for the species–area relationship, the emergence of a power‐law regime at intermediate scales and the augment of the species–area exponent z with the speciation rate. Despite the recent efforts, some other questions remain, such as the dependence of z in the whole range of the speciation rate. Questions like these are currently debated but generalizations cannot be drawn. This is the first paper, to our knowledge, that uses the coalescence method and neutral theory to examine biodiversity on more complex spatial structures. Our simulation results corroborate that the fragmentation plays a crucial role in shaping the species–area relationship, by determining the existence and extension of the power‐law regime associated with small and intermediate areas. On the other hand, when individuals are allowed to disperse over longer distances the species–area relationship now displays the classic triphasic pattern, and the intermediate regime, which is well described by a power‐law, is established even for highly fragmented landscapes.     

Using prioritisation tools to strategically restore vegetation communities in fragmented agricultural landscapes

Restoring native habitats in heavily cleared and fragmented areas such as agricultural landscapes is important to maintain and increase remaining native floral and faunal communities. Identifying priority vegetation types for restoration – as well as the parcels of land where this restoration could take place at a landscape scale – may assist in strategically protecting these biodiversity assets. To prioritise the restoration of terrestrial habitats around an ecologically and culturally significant Ramsar‐listed wetland in South Australia, we used the spatial prioritisation tool Marxan. Originally designed for prioritising the protection of reserve areas, Marxan can also be used to identify parcels of land for restoration purposes. We tested how Marxan prioritised the restoration of four distinct vegetation types around the Coorong and Lower Lakes region of South Australia using the inverse of habitat remnancy as a cost and soil type and distance to ecologically significant bird species as a conservation feature. By prioritising restoration activities around certain landscape features, such as remnant areas, our results indicate that we would be able to strategically restore parcels of native habitat that would maximise biodiversity outcomes. This study highlights the need for robust input data, such as priority vegetation types and bird species associated with these habitats, to ensure informative modelling outputs. It also suggests that other measures, such as the cost of different land types, should be included in future restoration planning. Finally, we illustrate how prioritisation tools such as Marxan can be used by natural resource managers to restore areas within fragmented agricultural landscapes.     

Yellow‐throated miners Manorina flavigula homogenize bird communities across intact and fragmented landscapes

In Australia, the role of noisy miners Manorina melanocephala in biotic homogenization of the avifauna has been well established in modified landscapes, and is listed as a threatening process under national conservation legislation. However, less is known about the effect of the congeneric and more widely distributed yellow‐throated miner, M. flavigula. In this paper we investigate the relative roles of habitat loss and increased dominance by the yellow‐throated miner in avian homogenization and species functional group decline. We examined bird community data collected from 368 woodland sites across three bioregions. For each site there was a local and a landscape scale measure of remnant vegetation cover. We used both multivariate and regression analysis to test the relative influence of yellow‐throated miner abundance and vegetation on bird community composition. There was clear compositional change and homogenization of the avifauna where yellow‐throated miners were present and vegetation cover was low. The abundance of 40 bird species was predicted by combinations of vegetation cover or yellow‐throated miner abundance, and 31 of these regressions included the term yellow‐throated miner. Of these, there was a negative relationship with 23 species, and 19 of these were insectivores or nectarivores. We postulate that the combination of clearing and yellow‐throated miner abundance can interact to disrupt the ecological function of woodlands, by the depletion of insect‐ and nectar‐feeding species and the disturbance to mixed feeding flocks. We propose future research objectives that include a continental‐scale analysis of the determinants of yellow‐throated miner overabundance, the numerical and geographical thresholds of their potential impacts, and the ecological consequences on both avifauna and the woodlands they inhabit.     

Functional biotic homogenization of bird communities in disturbed landscapes

Aim  Worldwide, functional homogenization is now considered to be one of the most prominent forms of biotic impoverishment induced by current global changes. Yet this process has hardly been quantified on a large scale through simple indices, and the connection between landscape disturbance and functional homogenization has hardly been established. Here we test whether changes in land use and landscape fragmentation are associated with functional homogenization of bird communities at a national scale.
Location  France.
Methods  We estimated functional homogenization of a community as the average specialization of the species present in that community. We studied the spatial variation of this community specialization index (CSI) using 1028 replicates from the French Breeding Bird Survey along spatial gradients of landscape fragmentation and recent landscape disturbance, measured independently, and accounting for spatial autocorrelation.
Results  The CSI was very sensitive to both measures of environmental degradation: on average, 23% of the difference in the CSI values between two sample sites was attributed to the difference in fragmentation and the disturbance between sites. This negative correlation between CSI and sources of landscape degradation was consistent over various habitats and biogeographical zones.
Main conclusions  We demonstrate that the functional homogenization of bird communities is strongly positively correlated to landscape disturbance and fragmentation. We suggest that the CSI is particularly effective for measuring functional homogenization on both local and global scales for any sort of organism and with abundance or presence–absence data.     

Predation leads to assembly rules in fragmented fish communities

Diamond [Assembly of species communities. In: Ecology and Evolution of Communities (eds Cody, M.L. & Diamond, J.M.). Harvard University Press, Cambridge, pp. 342–444] proposed that resource competition leads to checkerboard-like distributions of competing species. This proposal prompted research that revealed checkerboard patterns within a wide range of communities, but the mechanisms that generate such patterns are still poorly understood. Here we present whole-lake natural experiments and analyses of species–environment relationships in small coastal lake fish communities that were fragmented when land uplift isolated these lakes from the Baltic Sea, showing that a combination of predation and habitat suitability generated checkerboard distributions. Checkerboard patterns developed because two piscivores, northern pike and Eurasian perch, caused the extinction of several prey species in deep lakes. Conversely, low oxygen levels in shallow lakes caused extinction of the piscivores, and these areas served as a refuge for tolerant prey species. Based on these findings, we suggest that habitat suitability and biotic interactions should be viewed simultaneously in null models of assembly rules.     

Spatial dynamics of Microtus vole populations in continuous and fragmented agricultural landscapes

Small mammal populations often exhibit large-scale spatial synchrony, which is purportedly caused by stochastic weather-related environmental perturbations, predation or dispersal. To elucidate the relative synchronizing effects of environmental perturbations from those of dispersal movements of small mammalian prey or their predators, we investigated the spatial dynamics of Microtus vole populations in two differently structured landscapes which experience similar patterns of weather and climatic conditions. Vole and predator abundances were monitored for three years on 28 agricultural field sites arranged into two 120-km-long transect lines in western Finland. Sites on one transect were interconnected by continuous agricultural farmland (continuous landscape), while sites on the other were isolated from one another to a varying degree by mainly forests (fragmented landscape). Vole populations exhibited large-scale (>120 km) spatial synchrony in fluctuations, which did not differ in degree between the landscapes or decline with increasing distance between trapping sites. However, spatial variation in vole population growth rates was higher in the fragmented than in the continuous landscape. Although vole-eating predators were more numerous in the continuous agricultural landscape than in the fragmented, our results suggest that predators do not exert a great influence on the degree of spatial synchrony of vole population fluctuations, but they may contribute to bringing out-of-phase prey patches towards a regional density level. The spatial dynamics of vole populations were similar in both fragmented and continuous landscapes despite inter-landscape differences in both predator abundance and possibilities of vole dispersal. This implies that the primary source of synchronization lies in a common weather-related environment.     

Using historical ecology to understand patterns of biodiversity in fragmented agricultural landscapes

Aim To enhance current attempts to understand biodiversity patterns by using an historical ecology approach to highlight the over‐riding influence of land‐use history in creating past, current and future patterns of biodiversity in fragmented agricultural landscapes. Methods We develop an integrative conceptual framework for understanding spatial and temporal variations in landscape patterns in fragmented agricultural landscapes by presenting five postulates (hypotheses) which highlight the important role of historical, anthropogenic disturbance regimes. We then illustrate each of these postulates with examples drawn from fragmented woodlands in agricultural areas of south‐eastern Australia, and discuss these findings in an international context. Location examples are drawn from agricultural areas in south‐eastern Australia. Results We conclude that there is limited potential to refine our understanding of patterns of biodiversity in human‐modified landscapes based on traditional concepts of island biogeography, or simple assumptions of ongoing destruction and degradation. Instead, we propose that in agricultural landscapes that were largely cleared over a century ago: (1) present‐day remnant vegetation patterns are not accidental, but are logically arrayed due to historic land‐use decisions, (2) historic anthropogenic disturbances have a major influence on current ecosystem conditions and diversity patterns, and (3) the condition of remnant ecosystems is not necessarily deteriorating rapidly. Main conclusions An historical ecology approach can enhance our understanding of why different species and ecosystem states occur where they do, and can explain internal variations in ecological conditions within remnant ecosystems, too often casually attributed to the ‘mess of history’. This framework emphasizes temporal changes (both past and future) in biotic patterns and processes in fragmented agricultural landscapes. Integration of spatially and temporally explicit historical land‐use information into ecological studies can prove extremely useful to test hypotheses of the effects of changes in landscape processes, and to enhance future research, restoration and conservation management activities.     

Contrasting effects of habitat quantity and quality on moth communities in fragmented landscapes

Habitat loss is commonly identified as a major threat to the loss of global biodiversity. In this study, we expand on our previous work by addressing the question of how lepidopteran species richness and composition vary among remnants of North American eastern deciduous forest located within agricultural or pastoral landscapes. Specifically, we tested the relative roles of habitat quantity (measured as stand area and percent forest in the greater landscape) and habitat quality (measured as tree species diversity) as determinants of moth species richness. We sampled >19 000 individuals comprising 493 moth species from 21 forest sites in two forested ecoregions. In the unglaciated Western Allegheny Plateau, the species richness of moths with woody host plants diminished as forest stand size and percent forest in the landscape decreased, but the total species richness and abundance of moths were unaffected by stand size, percent forest in the landscape, or tree species diversity. In contrast, the overall species richness and abundance of moths in the glaciated North Central Tillplain were affected primarily by tree species diversity and secondarily by forest size. Higher tree species diversity may reduce species loss from smaller forest stands, suggesting that small, diverse forests can support comparable numbers of species to those in less diverse, large stands. Smaller forests, however, contained a disproportionate number of moth species that possess larvae known to feed on herbaceous vegetation. Thus, although woody plant feeding moths are lost from forests with changes in stand area, new species appear capable of recolonizing smaller fragments from the surrounding habitat matrix. Our study further suggests that when species replacement occurs, local patch size and habitat quality may be more important than landscape context in determining the community structure of forest Lepidoptera.     

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.     

破碎化次生林斑块面积及栖息地质量对繁殖鸟类群落结构的影响

于1998～2000年夏季。运用GPS定位系统确定了12块面积范围为6．5～112．8hm。的彼此隔离的森林斑块。比较了斑块面积和栖息地质量对繁殖鸟类群落结构的影响。结果表明：不同面积斑块中繁殖鸟类的群落结构有所差异。各斑块所容纳的繁殖鸟类的物种数从4种到26种不等。鸟类物种数随着斑块面积的增大而增多。不同鸟类对斑块面积的反应并不相同,耐边缘种偏爱面积较小的斑块。而非边缘种偏爱在大面积的斑块中繁殖。斑块栖息地质量也是影响鸟类群落结构的重要因素,质量好的斑块包容的鸟类物种较多。鸟类物种丰富度与斑块质量的相关性(R2=0．67)小于与斑块面积的相关性(R2=0．88)。各斑块中的鸟类群落结构在不同年份间比较相似。栖息地破碎化不但缩小了栖息地面积。同时也不同程度地降低了栖息地的质量。从而消极地影响着鸟类群落结构的稳定性和鸟类的物种多样性。     

Metapopulation extinction in fragmented landscapes: using bacteria and protozoa communities as model ecosystems

Extinction is notoriously difficult to study because of the long timescales involved and the difficulty in ascertaining that extinction has actually occurred. The effect of habitat subdivision, or fragmentation, on extinction risk is even harder to study, as it requires copious replication of habitat patches on large spatial scales and control of area effects between treatments. I used simple small-scale communities of bacteria and protozoa to study extinction in response to habitat loss and habitat fragmentation. I studied several different community configurations, each with three trophic levels. Unlike most metapopulation studies (experimental as well as theoretical), which have tended to deal with inherently unstable species interactions, I deliberately used community configurations that were persistent in large stock cultures. I recorded the time to extinction of the top predator in single habitat patches of different sizes and in fragmented systems with different degrees of subdivision but the same amount of available habitat. Habitat loss reduced the time to extinction of isolated populations. Fragmented systems went extinct sooner than corresponding unfragmented (continuous) systems of the same overall size. Unfragmented populations persisted longer than fragmented systems (metapopulations) with or without dispersal corridors between subpopulations. In fact, fragmented systems where the fragments were linked by dispersal corridors went extinctly significantly sooner than those where subpopulations were completely isolated from each other. If these results extend to more "natural" systems, it suggests a need for caution in management programs that emphasize widespread establishment of wildlife corridors in fragmented landscapes.     

Managing rock outcrops to improve biodiversity conservation in Australian agricultural landscapes

Summary  Rocky outcrops are prominent geological features in agricultural landscapes worldwide. Reptiles are a major component of these habitats and some species are restricted to, and more abundant on, rocky outcrops than in remnant vegetation. Rock outcrops are important to reptiles because they provide resources that are often limited in the surrounding landscape (e.g. micro-gradients in climatic conditions, basking- and retreat-sites). However, there is a knowledge gap in the literature addressing the conservation value of small, rocky outcrops. Management may be necessary to reverse habitat degradation in these systems. We identify four key areas of management that need to be addressed to improve outcrop habitat values and enhance biodiversity conservation in agricultural landscapes. Specific actions involve: (i) protecting outcrops from processes that cause damage to rock microhabitat, (ii) monitoring and managing changes in vegetation structure to maintain thermally suitable environments, (iii) applying integrated pest animal control and (iv) improving matrix management to enhance inselberg function and landscape connectivity. Further research is required to evaluate the efficacy of different management regimes on outcrop biota. We hope this paper will provide the stimulus for land managers to incorporate rocky outcrops in future biodiversity conservation programmes.     

Noncrop features and heterogeneity mediate overwintering bird diversity in agricultural landscapes of southwest China

Farmland birds are of conservation concerns around the world. In China, conservation management has focused primarily on natural habitats, whereas little attention has been given to agricultural landscapes. Although agricultural land use is intensive in China, environmental heterogeneity can be highly variable in some regions due to variations in crop and noncrop elements within a landscape. We examined how noncrop heterogeneity, crop heterogeneity, and noncrop features (noncrop vegetation and water body such as open water) influenced species richness and abundance of all birds as well as three functional groups (woodland species, agricultural land species, and agricultural wetland species) in the paddy‐dominated landscapes of Erhai water basin situated in northwest Yunnan, China. Birds, crop, and noncrop vegetation surveys in twenty 1 km × 1 km landscape plots were conducted during the winter season (from 2014 to 2015). The results revealed that bird community compositions were best explained by amounts of noncrop vegetation and compositional heterogeneity of noncrop habitat (Shannon–Wiener index). Both variables also had a positive effect on richness and abundance of woodland species. Richness of agricultural wetland species increased with increasing areas of water bodies within the landscape plot. Richness of total species was also greater in the landscapes characterized by larger areas of water bodies, high proportion of noncrop vegetation, high compositional heterogeneity of noncrop habitat, or small field patches (high crop configurational heterogeneity). Crop compositional heterogeneity did not show significant effects neither on the whole community (all birds) nor on any of the three functional groups considered. These findings suggest that total bird diversity and some functional groups, especially woodland species, would benefit from increases in the proportion of noncrop features such as woody vegetation and water bodies as well as compositional heterogeneity of noncrop features within landscape.     

The conservation value of cacao agroforestry for bird functional diversity in tropical agricultural landscapes

Cacao agroforestry have been considered as biodiversity‐friendly farming practices by maintaining habitats for a high diversity of species in tropical landscapes. However, little information is available to evaluate whether this agrosystem can maintain functional diversity, given that agricultural changes can affect the functional components, but not the taxonomic one (e.g., species richness). Thus, considering functional traits improve the understanding of the agricultural impacts on biodiversity. Here, we measured functional diversity (functional richness‐FD, functional evenness‐FEve, and functional divergence‐Rao) and taxonomic diversity (species richness and Simpson index) to evaluate changes of bird diversity in cacao agroforestry in comparison with nearby mature forests (old‐growth forests) in the Brazilian Atlantic Forest. We used data from two landscapes with constraining areas of mature forest (49% Una and 4.8% Ilhéus) and cacao agroforestry cover (6% and 82%, respectively). To remove any bias of species richness and to evaluate assembly processes (functional overdispersion or clustering), all functional indices were adjusted using null models. Our analyses considered the entire community, as well as separately for forest specialists, habitat generalists, and birds that contribute to seed dispersal (frugivores/granivores) or invertebrate removal (insectivores). Our findings showed that small cacao agroforestry in the forested landscape sustains functional diversity (FD and FEve) as diverse as nearby forests when considering the entire community, forest specialist, and habitat generalists. However, we observed declines for frugivores/granivores and insectivores (FD and Rao). These responses of bird communities differed from those observed by taxonomic diversity, suggesting that even species‐rich communities in agroforestry may capture lower functional diversity. Furthermore, communities in both landscapes showed either functional clustering or neutral processes as the main driver of functional assembly. Functional clustering may indicate that local conditions and resources were changed or lost, while neutral assemblies may reveal high functional redundancy at the landscape scale. In Ilhéus, the neutral assembly predominance suggests an effect of functional homogenization between habitats. Thus, the conservation value of cacao agroforestry to harbor species‐rich communities and ecosystem functions relies on smallholder production with reduced farm management in a forested landscape. Finally, we emphasize that seed dispersers and insectivores should be the priority conservation targets in cacao systems.     

Biodiversity in agricultural landscapes: different non-crop habitats increase diversity of ground-dwelling beetles (Coleoptera) but support different communities

Agricultural intensification poses a major threat to the conservation of biodiversity and associated ecosystem services. Since non-crop habitats are regarded as important refuges for farmland biodiversity, various greening measures have been proposed to halt biodiversity loss. However, the effectiveness of these measures for biodiversity conservation is still under debate. Therefore, we here compared ground-dwelling beetle (Coleoptera) assemblages of different non-crop habitats (field margins, set-aside fields sown with wildflowers, and permanent grassland fallows) and wheat fields within an intensively used agricultural landscape in western Germany. Taxonomic diversity of Carabidae, Staphylinidae and other coleopteran families and their conservation value were higher in all non-crop habitats than on wheat fields. Surprisingly, though, different types of non-crop habitats did not differ in species richness or the number of threatened species. Thus, field margins and sown wildflower fields were as effective in promoting beetle diversity as grassland fallows. However, different non-crop habitats supported different species assemblages, and several species, in particular especially large ones, were restricted to grassland fallows. These results suggest that different greening measures are effective in promoting the biodiversity of beetles, and that permanent grassland fallows are essential for nature conservation. The fact that habitat types harbored different assemblages stresses the need to combine a variety of greening measures to yield the highest benefit for biodiversity.

     

Roles of fragmented and logged forests for bird communities in industrial Acacia mangium plantations in Indonesia

Industrial timber plantations severely impact biodiversity in Southeast Asia. Forest fragments survive within plantations, but their conservation value in highly deforested landscapes in Southeast Asia is poorly understood. In this study, we compared bird assemblages in acacia plantations and fragmented forests in South Sumatra to evaluate each habitat’s potential conservation value. To clarify the impact of habitat change, we also analyzed the response of feeding guild composition. Five habitat types were studied: large logged forest (LLF), burnt logged forest (BLF), remnant logged forest (RLF), 4-year-old acacia plantation (AP4), and 1-year-old acacia plantation (AP1). Estimated species richness (Chao 2) was highest in LLF then AP4 and BLF, while AP1 and RLF had lower estimated species richness. Community composition was roughly divided into two groups by non-metric multidimensional scaling ordination: acacia plantation and logged forest. Sallying substrate-gleaning insectivores, such as drongos, broadbills, and some flycatchers, were restricted to LLF, whereas acacia plantation hosted many terrestrial frugivores, such as doves. Although fragmented forests in our study site lacked several common tropical forest species, these fragments provide an important habitat for some sallying and terrestrial insectivores. A network of small riparian remnant forests could be a complementary habitat for some species, while the conservation value of burnt forest might be low. In conclusion, the highly fragmented forests in plantations are suboptimal habitats for birds but are still very important, because large primary forest blocks have been nearly lost in the surrounding landscape.     

Variation in plant species composition in and adjacent to 64 ponds in SE Norwegian agricultural landscapes

All vascular plants, classified by life and growth form into aquatic species (hydrophytes and helophytes) and terrestrial species of pond margins, were inventoried in 64 SE Norwegian agricultural landscape ponds and their adjacent margins for which also 56 explanatory variables were recorded. Gradients in species composition, found separately for aquatic and terrestrial species by parallel DCA and GNMDS ordinations, were interpreted by correlation and geostatistical analyses.     

Variation in plant species richness in and adjacent to 64 ponds in SE Norwegian agricultural landscapes

All vascular plants, classified by life and growth form into six groups, four groups of hydrophytes (lemnids, nymphaeids, elodeids and isoetids), helophytes, and terrestrial species of pond margins, were inventoried in 64 SE Norwegian agricultural landscape ponds and their adjacent margins. The study sites varied considerably with respect to species richness; 0–4 for each hydrophyte group, 0–9 for helophytes, and 13–77 for terrestrial species. A total of 56 explanatory variables were recorded for each pond and adjacent margin to explain the observed richness variability.