Sustaining biodiversity conservation in human-modified landscapes in the Western Ghats: Remnant forests matter

Human-modified tropical landscapes under semi-natural or agro-ecosystems often harbor biodiversity of significant conservation value. In the Western Ghats of India, these ecosystems also provide connectivity between protected areas and other remnant forests. We investigated the conservation value of these landscapes and agro-ecosystems using results from 35 studies covering 14 taxonomic groups. Large, conspicuous taxonomic groups and tree-covered land-use types have received much focus in this area of research in the Western Ghats. We computed a response ratio defined as the log ratio of species richness in human land use to species richness in forest control site from 17 studies. In a meta-analysis, we investigated variation of this ratio across studies with respect to three variables: taxonomic group, the land-use type sampled and the extent of forest cover within the study landscape. Higher forest cover within the landscape emerged as a major positive influence on biodiversity in human-modified landscapes for vertebrates and vegetation while no patterns emerged for invertebrates. Our results suggest that loss of remnant forest patches from these landscapes is likely to reduce biodiversity within agro-ecosystems and exacerbate overall biodiversity loss across the Western Ghats. Conservation of these remnant forest patches through protection and restoration of habitat and connectivity to larger forest patches needs to be prioritized. In the densely populated Western Ghats, this can only be achieved by building partnerships with local land owners and stakeholders through innovative land-use policy and incentive schemes for conservation.     

Biodiversity conservation in human-modified Amazonian forest landscapes

Amazonia (sensu lato) is by far the largest tropical forest region, but has succumbed to the highest absolute rates of tropical deforestation and forest degradation, driven by rapid frontier expansion, road-building, and spontaneous or government-subsidized migration. The large area-through-time and paleo-climatic stability of Amazonian forests may help explain the high regional to local scale plant and animal species diversity of true forest specialists and high ecological sensitivity to contemporary land-use change. We describe the prevailing forms of anthropogenic disturbance that affect forest organisms in the context of the geographic and evolutionary background that has shaped the degree to which forest species may be resilient to environmental change. The fate of Amazonian biodiversity will partly depend upon the interaction between land-use and climate change, and the extent to which seasonally-dry forests can retain immunity against catastrophic recurrent wildfires. This review illustrates the importance of considering interactions between different forms of forest disturbance to develop effective conservation policy. We conclude with some considerations of the policy agenda necessary to protect forest cover and forest biodiversity at a meaningful scale across the Amazonian biome.     

Biodiversity conservation in human-modified landscapes of Mesoamerica: Past, present and future

Mesoamerica provides a unique context for biodiversity conservation in managed landscapes because of its geography, history of human intervention, and present conservation and development initiatives. The long and narrow form of the Mesoamerican landmass, and its division by a central mountain range, has served as both a bridge and a barrier. Conservation efforts in Mesoamerica are unique for the emphasis they place on regional connectivity through the Mesoamerican Biological Corridor and on biodiversity conservation in managed landscapes. The emphasis on conservation in agricultural systems has fostered innovations in payment for ecosystem services, and provides novel insights on the functional role that biodiversity plays in the provisioning of ecosystem services. The increasing rate of economic development in the region and the advent of new payment for ecosystem service schemes have provided new opportunities for forest regeneration and restoration. However, the small scale of private landholdings and the diversity of land uses featured in the region, while contributing to biodiversity conservation due to their structural and floristic complexity, present challenges for biodiversity monitoring and management.     

The challenge of maintaining Atlantic forest biodiversity: A multi-taxa conservation assessment of specialist and generalist species in an agro-forestry mosaic in southern Bahia

Recent developments have highlighted the importance of forest amount at large spatial scales and of matrix quality for ecological processes in remnants. These developments, in turn, suggest the potential for reducing biodiversity loss through the maintenance of a high percentage of forest combined with sensitive management of anthropogenic areas. We conducted a multi-taxa survey to evaluate the potential for biodiversity maintenance in an Atlantic forest landscape that presented a favorable context from a theoretical perspective (high proportion of mature forest partly surrounded by structurally complex matrices). We sampled ferns, butterflies, frogs, lizards, bats, small mammals and birds in interiors and edges of large and small mature forest remnants and two matrices (second-growth forests and shade cacao plantations), as well as trees in interiors of small and large remnants. By considering richness, abundance and composition of forest specialists and generalists, we investigated the biodiversity value of matrix habitats (comparing them with interiors of large remnants for all groups except tree), and evaluated area (for all groups) and edge effects (for all groups except trees) in mature forest remnants. Our results suggest that in landscapes comprising high amounts of mature forest and low contrasting matrices: (1) shade cacao plantations and second-growth forests harbor an appreciable number of forest specialists; (2) most forest specialist assemblages are not affected by area or edge effects, while most generalist assemblages proliferate at edges of small remnants. Nevertheless, differences in tree assemblages, especially among smaller trees, suggest that observed patterns are unlikely to be stable over time.     

Biodiversity conservation and livelihoods in human-dominated landscapes: Forest commons in South Asia

Strict protected areas are a critical component in global biodiversity conservation, but the future of biodiversity conservation may well depend upon the ability to experiment successfully with a range of institutional forms, including those that permit human use. Here, we focus on forest commons in human-dominated landscapes and their role in biodiversity conservation at the same time as they provide livelihood benefits to users. Using a dataset of 59 forest commons located in Bhutan, India, and Nepal, we estimated tree species richness from plot vegetation data collected in each forest, and drew on interview data to calculate a livelihoods index indicating the overall contribution of each forest to villager livelihoods for firewood, fodder, and timber. We found that tree species richness and livelihoods were positively and significantly correlated (rho = .41, p < 0.001, N = 59). This relationship held regardless of forest type or country, though significance varied somewhat across these two factors. Further, both benefits were similarly associated with several drivers of social-ecological change (e.g., occupational diversity of forest users, total number of users, and forest size), suggesting identification of potential synergies and complexes of causal mechanisms for future attention. Our analysis shows that forest commons in South Asia, explicitly managed to provide livelihoods for local populations, also provide biodiversity benefits. More broadly, our findings suggest that although strict protected areas are effective tools for biodiversity conservation, a singular focus on them risks ignoring other resource governance approaches that can fruitfully complement existing conservation regimes.     

Surrogate taxa for biodiversity in agricultural landscapes of eastern Austria

In an agricultural landscape in eastern Austria eight terrestrial organism groups were investigated as potential biodiversity indicators. We present a cross-taxon congruence assessment obtained at the landscape scale using two groups of plants (bryophytes and vascular plants), five groups of invertebrates (gastropods, spiders, orthopterans, carabid beetles and ants) and one vertebrate taxon (birds). We tested four different approaches: correlated species counts, surrogate measures of the overall species richness that was assessed, a multi-taxa (or shopping basket) approach and a simple complementarity algorithm. With few exceptions, pairwise correlations between taxa, correlations between one taxon and the species richness of the remaining groups, and correlations between a combination of the richness of two taxa and the remaining species richness were highly positive. Complementarity-derived priority sets of sampling sites using one taxon as a surrogate for the pooled species richness of all other taxa captured significantly more species than selecting areas randomly. As an essential first step in selecting useful biodiversity indicators, we demonstrate that species richness of vascular plants and birds showed the highest correlations with the overall species richness. In a multi-taxa approach and in complementarity site selection, each of the eight investigated taxa had the capability to capture a high percentage of the overall species richness.     

Drastic erosion in functional attributes of tree assemblages in Atlantic forest fragments of northeastern Brazil

The long-term dynamics of plant communities remain poorly understood in isolated tropical forest fragments. Here we test the hypothesis that tropical tree assemblages in both small forest fragments and along forest edges of very large fragments are functionally much more similar to stands of secondary growth (5-65-yr old) than to core primary forest patches. The study was carried out in a severely fragmented landscape of the Brazilian Atlantic forest. Nine functional attributes of tree assemblages were quantified by sampling all trees (DBH ? 10 cm) within 75 plots of 0.1 ha distributed in four forest habitats: small forest fragments (3.4-79.6 ha), forest edges, second-growth patches, and primary forest interior areas within a large forest fragment (3500 ha). These habitats were markedly different in terms of tree species richness, and in the proportion of pioneer, large-seeded, and emergent species. Age of second-growth stands explained between 31.4% and 88.2% of the variation in the functional attributes of tree assemblages in this habitat. As expected, most traits associated with forest edges and small forest fragments fell within the range shown by early (<25-yr old) and intermediate-aged secondary forest stands (25-45-yr old). In contrast to habitat type, tree assemblage attributes were not affected by vegetation type, soil type and the spatial location of plots. An ordination analysis documented a striking floristic drift in edge-affected habitats. Our results suggest that conservation policy guidelines will fail to protect aging, hyper-fragmented landscapes from drastic impoverishment if the remaining forest patches are heavily dominated by edge habitat.     

Biased seed rain in forest edges: Evidence from the Brazilian Atlantic forest

In this paper, we test the hypothesis that seed rain in forest edges differs from that in forest interior in terms of seed abundance, species richness, seed size, dispersal mode, and manipulation by vertebrates. The study was carried out in the Coimbra forest (3500 ha), the largest fragment currently found in the Atlantic forest of northeast Brazil. We assessed seed rain during a 1-year period by using a 1500-m long transect (with 100 sampling units) for each habitat. Seeds were categorized according to size, dispersal mode, and vertebrate manipulation. A total of 76,207 seeds belonging to 146 species were collected during the whole study and in average edge received more seeds and species per unit of area during particular periods of time. However, the analysis of all seeds and species recorded in both habitats during the 1-year period revealed that forest interior received a significant higher percentage of medium, large and very large seeds (21.8%) and species (82.6%) in comparison to edge (13.5% of seeds and 57.5% of species). The contribution of large and very large seed species dispersed by vertebrates was also lower in the edge forest as it represented 13.04% of all vertebrate-dispersed species recorded in this habitat vs. 31.5 % in the forest interior. Finally, seeds handled by vertebrates accounted for 5.9% of all seeds in forest edge, and reached 11% in the forest interior. The biased seed rain documented in the Coimbra forest raises the possibility that the creation of forest edges may alter some attributes of seed rain, particularly its content of large-seeded plants and of those dispersed by vertebrates.     

农业景观生物多样性功能和保护对策

研究表明,农业集约化生产和农业景观均质化导致农业景观生物多样性降低,致使生物多样性相关野生资源保护、自然授粉、害虫调节、水土涵养等生态系统服务功能受损,进而影响了农业可持续发展。本文对国内外农业景观生物多样性及其生态系统服务功能研究和实践进行了综述。分别探讨了农业景观在生物多样性保护、授粉服务、调节服务、水土涵养等多方面生态服务功能中发挥的作用及其生物多样性保护的意义,总结了欧美在农业景观生物多样性保护所采取的实践措施,指出中国虽然在生物多样性保护方面做出很多努力,但却忽略了对农业景观的保护,近些年的城市化和集约化发展都进一步加剧了农业景观生物多样性的丧失,亟需借鉴欧美国家经验提出我国的农业景观生物多样性保护策略。我们认为保护和提高农业景观生物多样性的生态集约化对维护农作物产量,改善品种具有重要的作用。农业景观生物多样性保护需要从农田生态系统和农业景观两个尺度上开展农业景观综合管理,提高农田作物和景观植物多样性和异质性,恢复和提高生物多样性及其生态服务功能。最后,针对我国农业可持续发展面临的问题和需求,建议从政策和法规、监测和评估、工程技术研发、技术集成示范、生态补贴制度、培训和推广等方面,开展农业景观生物多样性保护。     

Designing protected areas to conserve riverine biodiversity: Lessons from a hypothetical redesign of the Kruger National Park

The process of designing protected areas to represent all ecosystems in an area adequately is becoming increasingly sophisticated. To date freshwater aquatic ecosystems have seldom been considered in this process. How much of a difference does it make when they are considered as well?This study examined the conservation of riverine biodiversity within 17 assessment units contained by the catchment areas of five perennial rivers flowing through Kruger National Park and two seasonal rivers that are largely contained within this park. Physical river types, fish species and invertebrate families or genera were used as surrogates of riverine biodiversity. Conservation planning software was used to select an optimal set of planning units to represent and maintain riverine biodiversity.The current spatial configuration of Kruger National Park, largely an accident of history, is particularly poor when assessed against the objective of conserving riverine biodiversity. Several alternative layouts are examined. These options are theoretical since there is little current opportunity to reassign land uses in the region. This study shows that substantially improved layouts for both riverine and terrestrial biodiversity are possible, under the constraint of the same total area under protection. The study also shows that even these optimal layouts are only partially successful in efforts to conserve fully representative samples of riverine biodiversity. Because of the longitudinal connectivity of rivers, conservation strategies that extend beyond protected areas are essential. Explicit conservation visions, targets and strategies need to be included in integrated water resource management plans.Based on the results of this study, nine recommendations are provided for increasing the effectiveness of current and future protected areas in conserving riverine biodiversity. These are to use systematic conservation planning to make biodiversity benefits explicit; mend the disconnect between terrestrial and freshwater conservation; use multiple surrogates wherever possible; be strategic about the collection and management of primary data; strive for maximum hydrologic connectivity; resist development pressure; foster good relationships across park fences; where relevant, pursue multi-national cooperation at the basin scale; and engage the value debate and resolve awareness and capacity constraints.     

Just passing through: Global change and the conservation of biodiversity in protected areas

Climate and land-use changes are expected to cause many species to shift into or beyond the boundaries of protected areas, leading to large turnover in species composition. Here, we tested whether long-established protected areas in Canada were more robust to such climate change impacts than areas with no formal protection by measuring changes in modeled butterfly species distributions (n = 139) within them. We used a recently established distribution modeling technique, Maxent, to model butterfly species’ distributions in two epochs (1900-1930 and 1960-1990). We compared rates of butterfly species richness and composition change within protected areas against distributions of randomly selected, ecologically similar, but non-protected, areas. Change in species richness and composition within protected areas were, for the most part, the same as changes observed among random areas outside protected area boundaries. These results suggest that existing protected area networks in Canada have provided little buffer against the impacts of climate change on butterfly species richness, possibly because land-use change surrounding long-standing protected areas has not been substantial enough to elevate the habitat protection afforded by these protected areas relative to other areas. Although protected areas are unarguably beneficial in conserving biological diversity, their capacity to maintain habitat appears insufficient to prevent broader-scale climate changes from sweeping species beyond their boundaries.     

Trends associated with residential development in riparian breeding bird habitat along the Snake River in Jackson Hole, WY, USA: Implications for conservation planning

Throughout North America, bird population declines may be attributable to loss of habitat on the breeding grounds. Human land uses such as residential development, greatly impact landscapes and biota, particularly breeding birds. Riparian areas, which are essential to breeding birds, are particularly vulnerable to development pressures. In North America, most studies have focused on eastern birds and it is unclear how western bird assemblages respond to the effects of residential development. We sampled bird community parameters and habitat variables at three spatial scales (microhabitat, macrohabitat, and landscape) along a residential development gradient within the Snake River riparian corridor in Jackson Hole, WY, USA. Principal component analysis and multiple linear regression statistical tests were used to determine the effects of housing densities on avian community parameters, guilds, individual species distributions, and environmental variables. Landscape-level features were most affected by residential development and trends associated with increasing housing densities, such as anthropogenic habitat fragmentation primarily structured local bird communities. Overall species richness and diversity declined with increasing residential development. Neotropical migrant species were most negatively impacted and consistently declined in proportional representation on forested plots as residential development densities increased. Food generalists, ground gleaners, and avian nest predators all increased with increasing residential development. Brood parasites, on the other hand, did not increase with increasing housing densities and their distribution may reflect the availability of nest host species. These results suggest that residential development within riparian habitats may be exerting a strong negative influence on western bird communities and at high densities may lead to a depauperate avian biota. The results of our study indicate that preserving forest structure may benefit certain species but will not likely conserve as diverse and rich an avian assemblage as minimizing fragmentation associated with residential development.     

Hierarchical systematic conservation planning at the national level: Identifying national biodiversity hotspots using abiotic factors in Russia

National conservation planning should operate with measures of biodiversity similar to those applied globally in order to harmonize national and international conservation strategies. Here we suggest quantitative measures which enable two criteria of the global biodiversity hotspots to be applied on a national level for 74 large countries, and show how these measures can be applied to map national biodiversity hotspots. The plant endemism criteria of global hotspots are captured by quantitative measures of endemism, which are approximately scale-independent and can be corrected to account for a country’s environmental conditions and conservation priorities. The flexible land use criteria for national biodiversity hotspots are defined from percentage of natural vegetation remaining in the global hotspots. The minimum-area-required approach is applied to define the borders of national biodiversity hotspots using data on vascular plants species richness. We show how national biodiversity hotspots can be mapped from the species-energy relationship for vascular plants using climate, topographical and land use data when spatial pattern of species richness is not known. This methodology to map national biodiversity hotspots from abiotic factors is applied to Russia as a case study. Three Russian biodiversity hotspots, North Caucasus, South Siberia and Far East were identified. The resulting hotspot maps cover national-scale environmental gradients across Russia and although they are also identified by Russian experts their actual geographical locations were hitherto unspecified. The large-scale national hotspots, identified for Russia, can be used for further fine scale and more detailed conservation planning.     

Prioritisation of conservation areas in the Western Ghats, India

Areas of high conservation value were identified in the Western Ghats using a systematic conservation planning approach. Surrogates were chosen and assessed for effectiveness on the basis of spatial congruence using Pearson’s correlations and Mantel’s tests. The surrogates were, threatened and endemic plant and vertebrate species, unfragmented forest areas, dry forests, sub-regionally rare vegetation types, and a remotely sensed surrogate for unique evergreen ecosystems. At the scale of this analysis, amphibian richness was most highly correlated with overall threatened and endemic species richness, whereas mammals, especially wide-ranging species, were better at capturing overall animal and habitat diversity. There was a significant relationship between a remote sensing based habitat surrogate and endemic tree diversity and composition. None of the taxa or habitats served as a complete surrogate for the others. Sites were prioritised on the basis of their irreplaceability value using all five surrogates. Two alternative reserve networks are presented, one with minimal representation of surrogates, and the second with 3 occurrences of each species and 25% of each habitat type. These networks cover 8% and 29% of the region respectively. Seventy percent of the completely irreplaceable sites are outside the current protected area network. While the existing protected area network meets the minimal representation target for 88% of the species chosen in this study and all of the habitat surrogates, it is not representative with regard to amphibians, endemic tree species and small mammals. Much of the prioritised unprotected area is under reserve forests and can thus be incorporated into a wider network of conservation areas.     

Plant species richness in the Chaco Serrano Woodland from central Argentina: Ecological traits and habitat fragmentation effects

The Chaco Serrano Woodland from central Argentina has been dramatically reduced during the past 30 years, and is currently confined to several isolates of different size. In this study, we evaluated the effects of forest size, isolation and edge formation on plant species richness. Furthermore, we tested whether plants species with particular ecological traits were differentially affected by habitat fragmentation. Habitat area showed the highest explanatory value for plant species richness in stepwise multiple regressions. The effect of area was most pronounced for rare species, suggesting that large forests are necessary to preserve species with low local or regional abundance. Differences between edge and interior of Chaco Serrano were more pronounced for native and shrub species richness. The analysis of individual species cover revealed that native and biotically pollinated plants were less abundant in woodland edges. Our results showed that forest transformation into smaller remnants has lead to an impoverishment of plant communities, with particular subsets of species defined by ecological traits (rarity, origin and pollination mode) being more susceptible.     

Small mammals in agricultural landscapes of Prince Edward Island (Canada): Effects of habitat characteristics at three different spatial scales

We examined the influence of habitat characteristics at the microhabitat, macrohabitat, and landscape spatial scales on small mammals occurring in 12 forest patches within four agricultural landscapes of Prince Edward Island (Canada). Landscape features were important determinants of small mammal variables at all levels, but especially at the community level, whereas microhabitat characteristics tended to influence small mammals at the population level. Macrohabitat characteristics had only minor effects on small mammals occurring in our study sites. Species richness was most strongly influenced by patch area, reaching a threshold at forest patches of roughly 8-10 ha. The proportions of both forest and hedgerow cover within 400 m from the study site were also significant determinants of small mammals species diversity, possibly reflecting their ability to perceive suitable habitats, forage in areas outside the forest patches, and/or disperse in agricultural landscapes. At least one small mammal species (Napaeozapus insignis) benefitted from the presence of agricultural fields at distances up to 1000 m. Tamias striatus benefitted from the presence of hedgerow cover within 400 m from forest patches, possibly allowing them to move between forest patches. Clearly, the maintenance of forest patches of 8-10 ha and of forest cover within 400 m from them is fundamental for the conservation of small mammals inhabiting agricultural landscapes on the Island. Conservation strategies should also consider the establishment of more effective regulations to prevent and/or reduce hedgerow removal on Prince Edward Island.     

Plant resource conservation planning in protected natural areas: an example from the Canary Islands, Spain

An approach to an overall management planning for the conservation of endangered species within protected natural areas is presented, based mainly on the development of methods for the diagnosis of the conservation status and the identification of critical life cycle stages (phenophases) of target species. This model includes other aspects inherent in the overall planning of a protected area so as to match its technical, human and economic resources in terms of research and conservation. We provide examples of some island endemics that are included in the “Recovery Plans for the Endangered Plants in the Canary Islands National Parks”. The use of demography and genetics is particularly emphasised. We propose reintroductions as one of the most important tools for the management and conservation of endangered plants and the use of “system dynamics” tools as an alternative to traditional population matrix analysis. We provide examples of the application of system dynamics to population dynamics studies and to the design of management actions.     

The performance of procedures for selecting conservation areas: waterbirds in the UK

In recent years, much attention has been directed to the ways in which the most important areas for inclusion in reserve networks can be identified, and the most effective ways in which existing networks can be expanded. In contrast, rather little attention has been paid to the present and likely future performance of the approaches that have actually been employed. Using Wetland Bird Survey data, the effectiveness of the current Special Protection Area (SPA) network in the UK was assessed by comparing annual counts for 17 species of migratory waterbird on SPAs with the numbers supported by hypothetical site networks selected by five alternative site-selection methods. These analyses suggest that focusing on complementarity between component sites rather than applying criteria to each site individually improved the level of representation for each species, conserved a greater percentage of the national total across the 17 species and was robust over time.     

Habitat fragmentation and the desiccation of forest canopies: A case study from eastern Amazonia

Across the deforestation frontier in eastern Amazonia, we examined the relationship between edge-related forest desiccation and deforestation patterns using remote-sensing techniques. Canopy-moisture levels were estimated over an eight-year period in three study sites that encompassed ∼100,000 km² in area. We found four main effects of deforestation on dry-season canopy desiccation. First, intact forests showed no detectable change in canopy water content whereas forests adjacent to clearings showed significant water loss. Second, the distance to which edge-related desiccation penetrated into forest interiors varied among landscapes with differing forest loss and fragmentation. In moderately fragmented landscapes (with 65% and 51% remaining forest cover), canopy desiccation extended 1-1.5 km into forest interiors, whereas in heavily fragmented landscapes (20% forest cover) desiccation penetrated up to 2.7 km into forests. Third, the magnitude of edge-related desiccation varied among landscapes with differing fragmentation. Moderately fragmented landscapes exhibited a greater magnitude of change in canopy-water loss over the first 1 km from an edge than did heavily fragmented landscapes. Finally, forest desiccation penetrated further into forests over time in the moderately fragmented landscapes, but not in the heavily fragmented landscape, where edge-related desiccation had evidently ‘saturated’ remaining forests. We conclude that protracted dry seasons will have far more serious effects on fragmented than intact rainforests, with the former becoming highly vulnerable to destructive fires. With ∼30,000 km of new forest edge being created annually in Brazilian Amazonia, these finding have serious implications for forest conservation.     

Assessing landscapes: a case study of tree and shrub diversity in the seasonally dry tropical forests of Oaxaca, Mexico and southern Honduras

Tropical dry forests, with their distinct and economically important diversity, are acknowledged conservation priorities because of alarming rates of forest conversion. Whilst it is realised that terrestrial conservation requires an understanding of landscape level patterns of diversity, forests are rarely assessed accordingly. Here we demonstrate that, in the case of the seasonally dry tropical forests of the Pacific watershed of Mesoamerica, landscape level assessment of woody diversity can inform decision making relevant to both between-landscape and within-landscape prioritisation. We report floristic surveys of dry forest landscapes in Oaxaca, Mexico and southern Honduras. It is noted that these forests are floristically similar to other seasonally dry tropical forests in the neotropics. By calculation of Genetic Heat Indices, a relative measure of the concentration of restricted range species in a sample, we determine that the conservation of the tree diversity of the coastal lowlands of Oaxaca should be prioritised over that of southern Honduras. The current conservation status of forested areas in Oaxaca is briefly described. We suggest that the greater degree of anthropogenic disturbance in southern Honduras may explain the relative lack of restricted range species there. We argue that some forest fallows can act as analogues of mature forest and therefore landscape elements other than mature forest need to be included in forest conservation assessments. We conclude that diversity sampling of any forest type should not be limited to mature forests, but extended to other elements of forested landscapes.