Identifying priority areas for bioclimatic representation under climate change: a case study for Proteaceae in the Cape Floristic Region, South Africa

Biological reserves are established to protect natural resources and represent the diversity of environments found within a region. Unfortunately, many systems of protected areas do not proportionally capture the range of environmental conditions occupied by species and communities. Combinations of habitat loss and climate change may exacerbate these representational biases, and result in future distributions of environmental conditions that bare little resemblance to historic patterns. New protected areas need to be established to correct existing biases, and create conservation networks that remain representative despite climate change, habitat loss, and changes in species distributions. We demonstrate a new method to identify and prioritize habitat based on its value for improving bioclimatic representation. We assessed representation provided by existing protected areas for 301 Proteaceae species under historic and projected 2050 climate across the Cape Floristic Region in South Africa. The existing reserve system has relatively modest biases with respect to current species distributions and climate. However, if the system is not supplemented, protected areas in 2050 will capture an increasingly skewed sample of climatic conditions occupied by Proteaceae. These biases can be repaired through the systematic establishment of new protected areas, and many of the most valuable areas coincide with high priority ecosystem components and irreplaceable elements identified in the Cape Action for People and the Environmental conservation plan. Protecting these areas achieves nearly the best possible improvement in climatic representation while also meeting biodiversity representation goals.     

Uneven regional distribution of protected areas in Finland: Consequences for boreal forest bird populations

The significance of reserves in maintaining forest bird species of conservation concern (N = 36) was studied by large-scale quantitative line transect bird censuses in Finland, which stretches 1100 km through the boreal zone from the hemiboreal to the subarctic. Altogether 12 245 km of line transect was carried out in 1981-2004 in reserves covering 28 910 km². Bird census data in protected areas were combined into 100 km × 100 km squares. As a group the studied bird species were rather evenly distributed throughout the reserve network. The present reserve network is particularly significant for species having their highest densities in northern Finland, because large proportions of the populations of these species occurred in protected areas. However, over half of the studied species were concentrated in protected areas of southern and central Finland, and only small proportions of their populations were included in the present reserve network. In contrast to the whole species pool of conservation concern, the studied individual forest bird species were not distributed evenly throughout the whole reserve network. This suggests that for most individual species a regionally concentrated network is a preferred option, but for the whole species group the reserve network should clearly be regionally complementary and representative even in the boreal zone, where species have rather wide ranges. Thus, only some species and their habitats can be preserved in a spatially uneven reserve network in boreal forests.     

Efficiency and effectiveness in representative reserve design in Canada: The contribution of existing protected areas

To be effective, reserve networks should represent all target species in protected areas that are large enough to ensure species persistence. Given limited resources to set aside protected areas for biodiversity conservation, and competing land uses, a prime consideration for the design of reserve networks is efficiency (the maximum biodiversity represented in a minimum number of sites). However, to be effective, networks may sacrifice efficiency. We used reserve selection algorithms to determine whether collections of existing individual protected areas in Canada were efficient and/or effective in terms of representing the diversity of disturbance-sensitive mammals in Canada in comparison to (1) an optimal network of reserves, and (2) sites selected at random. Unlike previous studies, we restricted our analysis to individual protected areas that met a criterion for minimum reserve size, to address issues of representation and persistence simultaneously. We also tested for effectiveness and efficiency using historical and present-day data to see whether protected area efficiency and/or effectiveness varied over time. In general, existing protected areas did not effectively capture the full suite of mammalian species diversity, nor are most existing protected areas part of a near-optimal solution set. To be effective, Canada’s network of reserves will require at minimum 22 additional areas of >2700 km². This study shows that even when only those reserves large enough to be effective are considered, protected areas systems may not be representative, nor were they representative at the time of establishment.     

Relationship between distributions of threatened plants and protected areas in Britain

The establishment and maintenance of a system of protected areas is central to regional and global strategies for the conservation of biodiversity. The current global trend towards human population growth and widespread environmental degradation means that such areas are becoming increasingly isolated in fragmented habitat islands. In regions in which this process is well advanced a high proportion of species are thus predicted to have become restricted to protected areas. Here using uniquely detailed datasets for Britain, a region with close to the global level of percentage coverage by statutory protected areas, we determine the extent of restriction of Red List vascular plant species of conservation concern to these areas. On the basis of currently known distributions, overall our results strongly support the importance of a dual conservation strategy in Britain, in which protected areas are maintained with particular reference to those biodiversity features (such as many threatened plant species) that are highly dependent on them, and in which components of the wider landscape are also managed in such a way as to promote the abundance and distribution of such features with particular reference to those which are unlikely to persist in protected areas alone.     

Establishment of protected areas in different ecoregions, ecosystems, and diversity hotspots under successive political systems

Protected areas are valuable tools for nature conservation but the effectiveness of reserve networks must be monitored continuously. Knowing the history of the establishment of protected areas can help to improve future conservation. We explore how different ecoregions, ecosystems and diversity hotspots have been incorporated in protected areas in Estonia during the last century. We found that the average rate of establishment of protected areas has been surprisingly constant despite profound changes in political systems. However, establishment of protected areas has varied regionally; an agricultural region in south-east Estonia has seen less protection, partly due to lower biodiversity. Wetland ecosystems were initially more placed under protection, whereas recently semi-natural grasslands have gained more attention. In contrast, farmland is under-protected. Surprisingly, biodiversity hotspots were no more protected than the national average. We consider how the development of a protected area network has been influenced by individual persons and public opinion under successive political systems. We suggest that simultaneous gap analysis of ecoregions, ecosystems and diversity hotspots provides a more complete picture than examining a single aspect. Therefore this study can be used as a model for other regions.     

Assessing the performance of the existing and proposed network of marine protected areas to conserve marine biodiversity in Chile

The growing concern about the profound influence of human activities on marine ecosystems has been the driving force behind the creation of marine reserves in the last few decades. With almost 4200 km of coastline, Chile has not been the exception to this trend. A set of conservation priority sites has recently been proposed by the Chilean government to expand the current marine reserve network. In this study, we used the most comprehensive information currently available on the distribution of 2513 marine species in Chile to assess the efficiency of the existing system of marine protected areas (MPA) and the conservation priority sites identified by the government. Additionally, we evaluated the vulnerability of the reserve network selected with respect to threatening human activities. Our results show that both the existing protected areas and the proposed priority sites are relatively effective at protecting Chilean marine biodiversity. However, the majority of the species that are not represented within the existing or projected MPA network have very restricted distributions and are, therefore, of high conservation concern. To cover all species requires a network of 35 MPAs (46% of the total number of planning units). Many of the sites identified as irreplaceable present conflict with one or more human activities, particularly in the central region of the country. This study emphasizes the need for a systematic conservation planning approach to maximize the representation of species and prioritize those areas where conflicts between marine biodiversity conservation and human activities may occur.     

Expanding protected areas beyond their terrestrial comfort zone: Identifying spatial options for river conservation

There has been very little consideration of freshwater ecosystems in identifying and designing protected areas. Recent studies suggest that protected areas hold enormous potential to conserve freshwater biodiversity if augmented with appropriate planning and management strategies. Recognizing this need, South Africa’s relevant government authority commissioned a spatial assessment to inform their national protected area expansion strategy. This study presents the freshwater component of the spatial assessment, aimed at identifying focus areas for expanding the national protected area system for the benefit of river biodiversity. Conservation objectives to guide the assessment aimed to improve representation of river biodiversity pattern and processes in both new and existing protected areas. Data to address these objectives were collated in a Geographic Information System (GIS) and a conservation planning algorithm was used as a means of integrating the multiple objectives in a spatially efficient manner. Representation of biodiversity pattern was based on achieving conservation targets for 222 river types and 47 freshwater fish endemic to South Africa. Options were also identified for representing coarse-scale biodiversity processes associated with free-flowing rivers and catchment-estuarine linkages. River reaches that, with only minor expansion of existing protected area boundaries, could be fully incorporated into the national protected area system were also identified. Based on this study, generic recommendations are made on how to locate, design and manage protected areas for river biodiversity: use appropriate planning units, incorporate both biodiversity pattern and process, improve planning and management of individual protected areas, incorporate a mixture of protection strategies, and embed planning into an ongoing research and implementation process.     

Species specific connectivity in reserve-network design using graphs

Systematic conservation planning applications based solely on the presence/absence of a large number of species are not sufficient to guarantee their persistence in highly fragmented landscapes. Recent developments have thus incorporated much desired spatial design considerations, and reserve-network connectivity has received increased attention. Nonetheless, connectivity is often determined without regard to species-specific responses to habitat fragmentation. But species differ in their dispersal ability and habitat requirements, making proximate priority areas necessary for some species, while undesirable for others. We present a novel approach that incorporates species-specific connectivity needs in reserve-network design. Importantly, our method differs from previous approaches in that connectivity is not part of the objective function, but part of the constraints, thus avoiding typical undesirable trade-off that may result in high connectivity for some species but null connectivity for others. We use graphs to describe the dispersal pattern of each species and our goal is to identify minimum sets of reserves with connected sites for each of the species. This is not a trivial problem and we present three algorithms, one heuristic and two integer cutting algorithms that guarantee optimality, based on different 0-1 linear programming formulations. Applications to simulated data show that one of the algorithms that guarantee optimality is superior to the other, although both have limited application due to the number of sites and species they can manage. Remarkably, the heuristic can obtain very satisfactory solutions in short computational times, surpassing the limitations of the exact algorithms.     

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.     

Systematic reserve design as a dynamic process: F-TRAC and the Florida Forever program

Systematic conservation planning has become an important tool for increasing the efficiency of conservation decisions, but many planning efforts result in static plans that may lose relevance over time. We developed a process whereby planning is integrated into the decision-making process and updated every six months in response to conservation actions. The Florida Forever program is a 10-year, $3 billion land acquisition program expected to acquire approximately 1.25 million acres (607,000 ha) for conservation in Florida through the year 2010. With limited funding and duration, the program needs to be able to efficiently protect the most natural resources for a fixed cost, a situation well suited to a systematic reserve design approach. To inform this program, we conducted an assessment of natural resource conservation needs and developed the Florida Forever Tool for Efficient Resource Acquisition and Conservation (F-TRAC), a systematic reserve design analysis based on a simulated annealing site selection algorithm using Marxan software. The analysis considered conservation needs for a variety of natural resources including species, natural communities, high quality watersheds, wetlands, and sustainable forestry. Each 6-month analysis identifies an efficient portfolio of sites for resource protection, given the amount of land area likely remaining to be acquired by the Florida Forever program. The Spring 2004 model portfolio had a cost threshold of 206,308 ha, met conservation targets for 18 of 32 resource conservation features, and identified approximately 116,000 ha outside of current land acquisition projects. This study also demonstrates the use of reserve design results to evaluate existing and proposed land acquisition projects and inform decision makers; and the evaluation of acquisition trends and program success based on potential achievements as indicated by reserve design analyses.     

Forest degradation deepens around and within protected areas in East Asia

Forest degradation in protected areas has been monitored around the world with remote sensing data, but degradation processes undetectable by widely used satellite sensors have been largely overlooked. Increased human pressures and socioeconomic development make forest protection more challenging, particularly for forest ecosystems that lie across national borders because of the differences in national socioeconomic policies and conditions within them. Here with Landsat data, Google Earth images, and field observations, we show that, in two adjacent biosphere reserves across the border of China and North Korea, over one half of primary forest landscapes have been deteriorated by exploitive uses, including seed harvesting and systematic logging. The combined effects of detectable and hidden degradation processes have further damaged forest ecosystems in the core areas in the two biosphere reserves, threatening sustainable biodiversity conservation in the region. It is urgent to develop cross-border collaborative conservation strategies that can help combat both detectable and hidden degradation processes at a regional scale.     

Three decades of deforestation in southwest Sumatra: Have protected areas halted forest loss and logging, and promoted re-growth?

Much of the forest cover in southern Sumatra, Indonesia has been cleared since the early 1970s, but accurate estimates of the scales and rates of loss are lacking. This study combined high-quality remote sensing applications and extensive field surveys, both to provide an accurate picture of deforestation patterns across an area of 1.17 million ha in southwest Sumatra and to assess whether southwest Sumatra’s Bukit Barisan Selatan National Park (BBSNP) has halted forest loss and logging, and promoted re-growth, since its creation in 1984. Of the single large (692,850 ha) contiguous area of forest standing across our study area in 1972, nearly half (344,409 ha) has been cleared from 1972 to 2002, at an average rate per original forest cover of 1.69% y⁻¹. In Gunung Raya Wildlife Sanctuary (GRWS) and Hydrological Reserves (HR), forests have shrunk by 28,696 ha and 113,105 ha, at an average rate of 2.74% y⁻¹ and 2.13% y⁻¹, respectively. In contrast, forests in BBSNP have reduced four times more slowly than those in GRWS and HR, and have shrunk by 57,344 ha, at an average rate of 0.64% y⁻¹. Nevertheless, the forests within BBSNP were cleared almost as rapidly during the post-establishment, as during the pre-establishment, period (0.65% y⁻¹ and 0.63% y⁻¹, respectively) despite the introduction of protection measures during the post-establishment period, following the government’s pledge to expand and protect Indonesia’s network of Protected Areas (PAs) at the 1982 Bali World Parks Congress. While these protection measures failed to slow down rates of forest loss caused by agricultural encroachments they reduced large-scale mechanised logging by a factor of 4.2 and stabilized some 8610 ha of agricultural encroachments, enabling forest re-growth.     

Comparing static versus dynamic protected areas in the Québec boreal forest

Conservation planning is often based on static mapping of species’ ranges or habitat distributions. Succession and disturbance alter, however, habitat quality and quantity through time especially under global climate and land use change scenarios; hence, static protected areas may not ensure habitat persistence and species survival. Here, we examined the relative merits of static and dynamic (floating) protected areas for the conservation of American marten (Martes americana) habitat in a dynamic boreal forest of Québec (Canada). Forest dynamics were modeled using a spatially-explicit landscape disturbance model and protected areas were selected based on the quality and compactness of marten home ranges using MARXAN. Static protected areas were fixed in space during 200 year simulations of boreal forest dynamics, while dynamic protected areas were re-located every 50 years to track dynamic habitat. Dynamic protected areas supported more high quality home ranges through time than static protected areas. The locations of dynamic protected areas were constrained, however, by the highly fragmented forest patterns created through logging and fire in unprotected areas. Our findings emphasize the often-overlooked point that if dynamic conservation planning is to be successful in the long term, the landscape matrix quality surrounding protected areas must be managed in such a way that options remain when it comes to re-planning.     

The role of private conservation areas in biodiversity representation and target achievement within the Little Karoo region, South Africa

It is becoming increasingly difficult to manage and expand statutory conservation areas (i.e., parks and formally protected areas). Therefore, alternative opportunities for land conservation merit closer attention. This paper examines the extent to which privately owned conservation areas contribute to biodiversity representation. Gap analyses were performed for a large semi-arid region in South Africa with a comprehensive database of private conservation areas. The distribution of private conservation areas was compared to statutory conservation areas using several landscape characteristics: biome and vegetation variant, elevation class, ecological process area, total area, and threat status (endangerment). Conservation target achievement for the vegetation variants was also assessed, as was the degree to which private conservation areas complemented statutory conservation areas by representing different landscape characteristics. The number of targets achieved nearly tripled if private conservation areas were considered in addition to statutory conservation areas. Further, private conservation areas significantly complemented statutory conservation areas in the types of biomes, elevation classes, and threat status classes conserved. Private conservation areas were especially important in conserving lower elevation habitat, and by association, endangered vegetation. This particular relationship is expected to be common worldwide. Our results indicate that private lands conservation deserves an increased allocation of resources for both research and implementation.     

Evaluating hedgerow corridors for the conservation of native forest herb diversity

The maintenance of connecting habitats such as hedgerows in production landscapes could become increasingly critical as species migration is expected to accelerate with climate change. Species of particular conservation interest that could benefit from connecting habitats especially in agroecosystems are native forest herbs. It is still unclear, however, which hedgerow habitats have the best potential of supporting diverse forest herb communities, making it hard to target particular structures for conservation. Our objective was to identify the local and landscape characteristics of hedgerows that could help predict their potential at maximizing the richness, abundance, and diversity of native forest herbs of temperate deciduous forests of North-East America. We used multiple regression, Moran’I correlograms, and conditional autoregressive models to assess the effect of landscape and historical variables (hedgerow age, amount of adjacent forest cover, connection to forests and other hedgerows) and of local variables (hedgerow width, canopy cover, cover of other species) on the response variables. The results show an increase in forest herb diversity and abundance in hedgerows with time and with increasing nearby forest cover. Local factors such as greater width and reduced cover of other species also relate to a greater abundance and diversity of forest herbs in linear habitats. The fact that forest herb communities can reassemble in hedgerow corridors with time implies that there could be long-term benefits in maintaining and even creating linear habitats where there is a pool of dispersing forest herbs.     

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.     

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.     

The buffer effect and the selection of protected areas for waterbirds in Britain

The buffer effect predicts that where the reproductive success and survivorship of a species vary between potential habitats, sites will be sequentially filled according to a preference hierarchy. Once favoured sites reach saturation, numbers on the less-suitable/poorer quality sites will show a greater rate of increase compared with those on favoured sites. Supporting evidence for a buffer effect is readily available in the literature for many species, although this is generally restricted to small-scale analyses. In this paper we test for a buffer effect on a national scale for 19 species of waterbirds regularly wintering in the UK for which populations have increased nationally. The results provide little support for the effect, with only four species showing significant negative correlations. Nonetheless, a number of factors are likely to confound the occurrence/identification of a buffer effect for these species, including site area and data limitations. By contrast, for the majority of these 19 species, those sites where initial population totals are largest are also those with the fastest rates of population increase. Encouragingly, these sites are, therefore, more likely to be classified and managed as Special Protection Areas (SPAs) or Ramsar sites using the current numerical criteria for identifying such conservation areas.     

The role of social aggregations and protected areas in killer whale conservation: The mixed blessing of critical habitat

Protected areas are often proposed as tools for conserving endangered populations of marine megafauna. Our study area includes a voluntary no-entry reserve embedded within wider critical habitat for Threatened ‘northern resident’ killer whales under Canada’s Species at Risk Act. Our study quantified the reserve’s importance to whales by assessing habitat preference in a behavioural context, and population-level implications of that preference given threats from human activities, such as oil spills resulting from shipping traffic. We recorded summertime activities of whales from 1995 to 2002. Whales were observed on 397 of 530 (74.9%) days. Whales showed strong preference for the reserve over adjacent waters, and used it preferentially for feeding and beach-rubbing. While the area comprises ∼0.001% of the whales’ range, an overall average of 6.5% of the population was present each day. Frequently, >50% of this small population was aggregated in the restricted and heavily trafficked waterway of Johnstone Strait. Using the Potential Biological Removal equation, we calculated potential annual mortality limits (ML) of 2.2 animals. Mean group size in the area exceeded ML on 55.8% of days overall, and 98.8% of days when conditioning on whale presence. The whales’ high reliance on a trivial fraction of their range means that opportunities are routine for one stochastic, catastrophic event to cause population decline. On 20 August 2007, a barge loaded with ∼10,000 L of diesel sank in the area, exposing 25% of the population. This underscores the importance of identifying critical habitat for threatened populations, and ensuring meaningful protection.     

Effects of forest stand type on oribatid mite (Acari: Oribatida) assemblages in a southwestern Quebec forest

Despite the ubiquity of oribatid mites in soil and litter systems, and their importance in decomposition and nutrient cycling processes, little is known of the factors underlying the composition of their assemblages. Our objective was to address this by determining how oribatid assemblage composition changes by forest stand type. This work was done in and near a hardwood forest in southwestern Quebec, Canada. We sampled mites by collecting 1 L of litter and 170 cm³ of soil from four sites in each of four distinct habitat types: American beech stands, sugar maple stands, mixed deciduous stands and mixed conifer plantations. Samples were collected in July and September 2005, and June 2006, and over 6500 oribatid mites were collected and identified to species. Abundance and species richness differed between forest types: for abundance conifer>beech>maple>mixed deciduous while for species richness beech and conifer>maple>mixed deciduous. Ordination analyses revealed that conifer plantations and beech stands supported distinct assemblages, while there were some overlap in the assemblages found in maple stands and mixed deciduous stands. These data support the importance of aboveground plant communities in affecting the composition of oribatid assemblages even at local scales and provide insight into additional impacts that may be caused by shifts in plant species ranges due to global changes.