基于全域连通性识别气候变化风险下的生物多样性保护优先区——以京津冀为例

在保护优先区规划中,有必要考虑气候变化的潜在风险并关注物种在气候驱动下的扩散格局。基于未来生物气候数据、地形多样性数据以及土地利用数据,应用Omniscape算法,对21世纪中叶(2040-2061年)气候变化情景下京津冀地区陆生哺乳动物的扩散进行全域连通性建模并与当前情景对比分析,识别出生物多样性保护优先区。结果表明:区域尺度下,气候变化风险使得高连通性的区域逐渐从平原向山区转移,分布趋于集中;斑块尺度下,林缘连通性较高,而位于林地或草地边缘的耕地连通性低。在此基础上,共识别生物多样性保护优先区共51786 km²,其中涵养区(占56.4%)在当前和未来的连通状况均较为良好;优化区(占38.4%)应提升生境质量以满足未来连通性的更高需求;而修复区(占5.22%)面临的气候变化风险较高,亟需进行生态修复以免在未来出现连通性夹点。本研究通过评估京津冀地区两种情景下的全域连通格局,为生物多样性保护的气候适应性规划提供了科学依据。     

Prediction of habitat suitability,connectivity, and corridors in the future to conserve roe deer (Capreolus capreolus) as a locally endangered species in northern Iran

Roe deer is a protected species in Iran as its population and distribution in the country have considerably declined. Roe deer are threatened by several factors such as habitat fragmentation and road mortality, so studying their distribution and movement through the increasing habitat destruction and fragmentation is necessary. This will become increasingly important because climate change will transform the species’ future habitat and connectivity patterns. We evaluated the roe deer’s potential distribution range in northern Iran and, for the first time, developed connectivity models and designed corridors for the present and future to make better conservation plans. We collected 91 points indicating the presence of roe deer in the study region. After developing ensemble models using six species distribution algorithms, we defined high-ranked habitat cores using the concept of landscape suitability prioritization. From these, we designed connectivity and corridors in two time-frames with the help of least-cost paths and circuit theories to predict the potential movement throughout the study area. We estimated that the overall core habitats for roe deer in the present and future periods are, respectively, around 1200 km² and 2600 km², corresponding to 2 and 4 percent of the whole area. This suggests that the habitat core will expand in the future as a result of climate change. Similarly, the connectivity among the cores will strengthen. We also conclude that the temperature-driven and anthropogenic variables significantly affect the distribution of roe deer in northern Iran. It is necessary that conservationists and managers consider the designed corridors in the present study while planning conservation strategies.     

Predicting impacts of climate change on habitat connectivity of Kalopanax septemlobus in South Korea

Understanding the drivers of habitat distribution patterns and assessing habitat connectivity are crucial for conservation in the face of climate change. In this study, we examined a sparsely distributed tree species, Kalopanax septemlobus (Araliaceae), which has been heavily disturbed by human use in temperate forests of South Korea. We used maximum entropy distribution modeling (MaxEnt) to identify the climatic and topographic factors driving the distribution of the species. Then, we constructed habitat models under current and projected climate conditions for the year 2050 and evaluated changes in the extent and connectivity of the K. septemlobus habitat. Annual mean temperature and terrain slope were the two most important predictors of species distribution. Our models predicted the range shift of K. septemlobus toward higher elevations under medium-low and high emissions scenarios for 2050, with dramatic reductions in suitable habitat (51% and 85%, respectively). In addition, connectivity analysis indicated that climate change is expected to reduce future levels of habitat connectivity. Even under the Representative Construction Pathway (RCP) 4.5 medium-low warming scenario, the projected climate conditions will decrease habitat connectivity by 78%. Overall, suitable habitats for K. septemlobus populations will likely become more isolated depending on the severity of global warming. The approach presented here can be used to efficiently assess species and habitat vulnerability to climate change.     

Scattered trees: a complementary strategy for facilitating adaptive responses to climate change in modified landscapes?

1.  Facilitating adaptive responses of organisms in modified landscape will be essential to overcome the negative effects of climate change and its interaction with land use change. Without such action, many organisms will be prevented from achieving the predicted range shifts they need to survive.
2.  Scattered trees are a prominent feature of many modified landscapes, and could play an important role in facilitating climate change adaptation. They are keystone structures because of the disproportionally large ecological values and ecosystem services that they provide relative to the area they occupy in these landscapes. The provision of habitat and connectivity will be particularly relevant.
3.  Scattered trees are declining in modified landscapes due to elevated tree mortality and poor recruitment often associated with intensive land use. The continuing global decline of scattered trees will undermine the capacity of many organisms to adapt to climate change.
4.   Synthesis and applications. The sustainable management of scattered trees in modified landscapes could complement other strategies for facilitating climate change adaptation. They create continuous, though sparse, vegetation cover that permits multi-directional movements of biota across landscapes and ecological networks. They have the capacity to span ecosystems and climatic gradients that cannot be captured in formal reserves alone. The management of scattered trees should be an integral part of conservation objectives and agricultural activities in modified landscapes. Public investment, through mechanisms such as agri-environmental schemes, in rotational grazing, temporary set-asides, tree-planting and regulations that reduce clearing and early mortality among standing trees will improve the capacity of biota to adapt to climate change.     

Consequences of large-scale processes for the conservation of bird populations

1.  Detailed studies of population ecology are usually carried out in relatively restricted areas in which emigration and immigration play a role. We used a modelling approach to explore the population consequences of such dispersal and applied ideas from our simulations to the conservation of wild birds.
2.  Our spatial model incorporates empirically derived variation in breeding output between habitats, density dependence and dispersal. The outputs indicate that dispersal can have considerable consequences for population abundance and distribution. The abundance of a species within a patch can be markedly affected by the surrounding habitat matrix.
3.  Dispersal between habitats may result in lower population densities at the edge of good quality habitat blocks and could partially explain why some species are restricted to large habitat fragments.
4.  Habitat deterioration may not only lead to population declines within that habitat but also in adjacent habitats of good quality. This may confound studies attempting to diagnose population declines.
5.  Although mobile species have the advantages of colonizing sites within metapopulations, dispersal into poorer quality territories may markedly reduce total populations.
6.  There are two main approaches to conservation: one is to concentrate on establishing and maintaining protected areas, while the other involves conservation of the wider countryside. If dispersal is an important process then protecting only isolated areas may be insufficient to maintain the populations within them.     

The effect of local change in habitat quality on populations of migratory species

1. Habitat deterioration is a major problem world-wide as a result of processes such as change in land use, introduced species, human disturbance and exploitation of food supplies. Many studies have shown that habitat change can have considerable effect on the numbers of individuals using a site. For migratory species, however, the consequences for the total population cannot be deduced from local studies.
2. For a migratory species, the change in total population size Δ N , as a consequence of habitat change in the wintering area, can be calculated from Δ N  =  LM γ d '/( b ' +  d '), where γ is the expected proportional change in the number of birds using a site as a result of the habitat change, L is the area affected, M is the density of individuals using the site prior to habitat change, b ' is the strength of the per capita density-dependent breeding output, and d ' is the strength of the per capita density-dependent winter mortality. Similarly the consequences of habitat change in the breeding area can be calculated from Δ N  =  LM γ b '/( b ' +  d ').
3. The same approach can be used for predicting the consequences of improvements in habitat quality.
4. A worked example is given to illustrate how this approach could be used to predict the consequences for the total population of changes in the food supply of oystercatchers within one estuary.
5. There is a need for more measures of γ, the expected proportional change in the number of birds using a site as a result of various forms of habitat deterioration, and the strengths of density dependence.     

Anthropogenic impact on habitat connectivity: A multidimensional human footprint index evaluated in a highly biodiverse landscape of Mexico

Evaluating the cumulative effects of the human footprint on landscape connectivity is crucial for implementing policies for the appropriate management and conservation of landscapes. We present an adjusted multidimensional spatial human footprint index (SHFI) to analyze the effects of landscape transformation on the remnant habitat connectivity for 40 terrestrial mammal species representative of the Trans-Mexican Volcanic System in Michoacán (TMVS_Mich), in western central Mexico. We adjusted the SHFI by adding fragmentation and habitat loss to its original three components: land use intensity, time of human landscape intervention, and biophysical vulnerability. The adjusted SHFI was applied to four scenarios: one grouping all species and three grouping several species by habitat spatial requirements. Using the SHFI as a dispersal resistance surface and applying a circuit theory based approach, we analyzed the effects of cumulative human impact on habitat connectivity in the different scenarios. For evaluating the relationship between habitat loss and connectivity, we applied graph theory-based equivalent connected area (ECA) index. Results show over 60% of the TMVS_Mich has high SHFI values, considerably lowering current flow for all species. Nevertheless, the effect on connectivity of human impact is higher for species with limited dispersal capacity (100–500 m). Our approach provides a new form of evaluating human impact on habitat connectivity that can be applied to different scales and landscapes. Furthermore, the approach is useful for guiding discussions and implementing future biodiversity conservation initiatives that promote landscape connectivity as an adaptive strategy for climate change.     

Determinants of local spider (Araneidae and Thomisidae) species richness on a regional scale: climate and altitude vs. habitat structure

Abstract.  1. The habitat heterogeneity hypothesis states that the more complex the habitat, the higher the species richness. The present study analyzes the effect of local factors on regional spider ( Araneidae and Thomisidae ) richness. The main objective is to disentangle the relative importance of habitat structure and other environmental variables.
2. Fifteen territorial units of 1 km² were sampled to obtain reliable and comparable inventories of the two spider families. Richness values were modelled using general regression models and a set of climate, topographic and vegetation structure variables. Pure and joint effects were computed via variation partitioning.
3. The results highlight the great importance of vegetation complexity, especially of grass and sub-shrub cover, in determining spider species richness.
4. The maximum temperature is the only climate variable significantly related to species richness, although its effect is combined with that of spatial and vegetation structure variables.
5. These results support the habitat heterogeneity hypothesis, and highlight the importance of taking vegetation complexity into account when managing habitats and where spider conservation is desired.     

Population genetics suggests effectiveness of habitat connectivity measures for the European tree frog in Switzerland

1.  Governmental authorities in many countries financially support the implementation of habitat connectivity measures to enhance the exchange of individuals among fragmented populations. The evaluation of the effectiveness of such measures is crucial for future management directions and can be accomplished by using genetic methods.
2.  We retraced the population history of the European tree frog in two Swiss river valleys (Reuss and Thur), performed comprehensive population sampling to infer the genetic structure at 11 microsatellite markers, and used first-generation migrant assignment tests to evaluate the contemporary exchange of individuals.
3.  Compared with the Thur valley, the Reuss valley has lost almost double the number of breeding sites and exhibited a more pronounced genetic grouping. However, similar numbers of contemporary migrants were detected in both valleys. In the Reuss valley, 81% of the migration events occurred within the identified genetic groups, whereas in the Thur valley migration patterns were diffuse.
4.  Our results show that the connectivity measures implemented in the Reuss valley facilitated effective tree frog migration among breeding sites within distances up to 4 km. Nevertheless, the Reuss valley exhibited high genetic differentiation, which reflected the impact of barriers to tree frog movement such as the River Reuss. By contrast in the Thur valley, a larger number of breeding sites have been preserved and high admixture indicated exchange of individuals at distances up to 16 km.
5.   Synthesis and applications . We show that genetic methods can substantiate the effectiveness of connectivity measures taken in conservation management at the landscape scale. We urge responsible authorities from both river valleys to continue implementing connectivity measures and to create a dense network of breeding sites, as spatial gaps of 8 km are rarely traversed by tree frogs.     

Alpine glacial relict species losing out to climate change: The case of the fragmented mountain hare population (Lepus timidus) in the Alps

Alpine and Arctic species are considered to be particularly vulnerable to climate change, which is expected to cause habitat loss, fragmentation and—ultimately—extinction of cold‐adapted species. However, the impact of climate change on glacial relict populations is not well understood, and specific recommendations for adaptive conservation management are lacking. We focused on the mountain hare (Lepus timidus) as a model species and modelled species distribution in combination with patch and landscape‐based connectivity metrics. They were derived from graph‐theory models to quantify changes in species distribution and to estimate the current and future importance of habitat patches for overall population connectivity. Models were calibrated based on 1,046 locations of species presence distributed across three biogeographic regions in the Swiss Alps and extrapolated according to two IPCC scenarios of climate change (RCP 4.5 & 8.5), each represented by three downscaled global climate models. The models predicted an average habitat loss of 35% (22%–55%) by 2100, mainly due to an increase in temperature during the reproductive season. An increase in habitat fragmentation was reflected in a 43% decrease in patch size, a 17% increase in the number of habitat patches and a 34% increase in inter‐patch distance. However, the predicted changes in habitat availability and connectivity varied considerably between biogeographic regions: Whereas the greatest habitat losses with an increase in inter‐patch distance were predicted at the southern and northern edges of the species’ Alpine distribution, the greatest increase in patch number and decrease in patch size is expected in the central Swiss Alps. Finally, both the number of isolated habitat patches and the number of patches crucial for maintaining the habitat network increased under the different variants of climate change. Focusing conservation action on the central Swiss Alps may help mitigate the predicted effects of climate change on population connectivity.     

A Framework to Optimize Biodiversity Restoration Efforts Based on Habitat Amount and Landscape Connectivity

The effectiveness of ecological restoration actions toward biodiversity conservation depends on both local and landscape constraints. Extensive information on local constraints is already available, but few studies consider the landscape context when planning restoration actions. We propose a multiscale framework based on the landscape attributes of habitat amount and connectivity to infer landscape resilience and to set priority areas for restoration. Landscapes with intermediate habitat amount and where connectivity remains sufficiently high to favor recolonization were considered to be intermediately resilient, with high possibilities of restoration effectiveness and thus were designated as priority areas for restoration actions. The proposed method consists of three steps: (1) quantifying habitat amount and connectivity; (2) using landscape ecology theory to identify intermediate resilience landscapes based on habitat amount, percolation theory, and landscape connectivity; and (3) ranking landscapes according to their importance as corridors or bottlenecks for biological flows on a broader scale, based on a graph theory approach. We present a case study for the Brazilian Atlantic Forest (approximately 150 million hectares) in order to demonstrate the proposed method. For the Atlantic Forest, landscapes that present high restoration effectiveness represent only 10% of the region, but contain approximately 15 million hectares that could be targeted for restoration actions (an area similar to today's remaining forest extent). The proposed method represents a practical way to both plan restoration actions and optimize biodiversity conservation efforts by focusing on landscapes that would result in greater conservation benefits .     

Seagrass patch size affects fish responses to edges

1.  Patch area and proximity of patch edge can influence ecological processes across patchy landscapes and may interact with each other. Different patch sizes have different amounts of core habitat, potentially affecting animal abundances at the edge and middle of patches. In this study, we tested if edge effects varied with patch size.
2.  Fish were sampled in 10 various-sized seagrass patches (114–5934 m²) using a small (0·5 m²) push net in three positions within each patch: the seagrass edge, 2 m into a patch and in the middle of a patch.
3.  The two most common species showed an interaction between patch size and the edge–interior difference in abundance. In the smallest patches, pipefish ( Stigmatopora nigra ) were at similar densities at the edge and interior, but with increasing patch size, the density at the edge habitat increased. For gobies ( Nesogobius maccullochi ), the pattern was exactly the opposite.
4.  This is the first example from a marine system of how patch size can influence the magnitude and pattern of edge effects.
5.  Both patch area and edge effects need to be considered in the development of conservation and management strategies for seagrass habitats.     

Predicting the effects of climate change on population connectivity and genetic diversity of an imperiled freshwater mussel,Cumberlandia monodonta (Bivalvia: Margaritiferidae), in riverine systems

In the face of global climate change, organisms may respond to temperature increases by shifting their ranges poleward or to higher altitudes. However, the direction of range shifts in riverine systems is less clear. Because rivers are dendritic networks, there is only one dispersal route from any given location to another. Thus, range shifts are only possible if branches are connected by suitable habitat, and stream‐dwelling organisms can disperse through these branches. We used Cumberlandia monodonta (Bivalvia: Unionoida: Margaritiferidae) as a model species to investigate the effects of climate change on population connectivity because a majority of contemporary populations are panmictic. We combined ecological niche models (ENMs) with population genetic simulations to investigate the effects of climate change on population connectivity and genetic diversity of C. monodonta. The ENMs were constructed using bioclimatic and landscape data to project shifts in suitable habitat under future climate scenarios. We then used forward‐time simulations to project potential changes in genetic diversity and population connectivity based on these range shifts. ENM results under current conditions indicated long stretches of highly suitable habitat in rivers where C. monodonta persists; populations in the upper Mississippi River remain connected by suitable habitat that does not impede gene flow. Future climate scenarios projected northward and headwater‐ward range contraction and drastic declines in habitat suitability for most extant populations throughout the Mississippi River Basin. Simulations indicated that climate change would greatly reduce genetic diversity and connectivity across populations. Results suggest that a single, large population of C. monodonta will become further fragmented into smaller populations, each of which will be isolated and begin to differentiate genetically. Because C. monodonta is a widely distributed species and purely aquatic, our results suggest that persistence and connectivity of stream‐dwelling organisms will be significantly altered in response to future climate change.     

How do species respond to climate change along an elevation gradient? A case study of the grey‐headed robin (Heteromyias albispecularis)

Climate is predicted to change rapidly in the current century, which may lead to shifts of species' ranges, reduced populations and extinctions. Predicting the responses of species abundance to climate change can provide valuable information to quantify climate change impacts and inform their management and conservation, but most studies have been limited to changes in habitat area due to a lack of abundance data. Here, we use generalized linear model and Bayesian information criteria to develop a predictive model based on the abundance of the grey‐headed robin (GHR) and the data of climatic environmental variables. The model is validated by leave‐one‐out cross‐validation and equivalence tests. The responses of GHR abundance, population size and habitat area by elevation are predicted under the current climate and 15 climate change scenarios. The model predicts that when temperature increases, abundance of GHR displays a positive response at high elevation, but a negative response at low elevation. High precipitation at the higher elevations is a limiting factor to GHR and any reduction in precipitation at high elevation creates a more suitable environment, leading to an increase in abundance of GHR, whereas changes in precipitation have little impact at low elevation. The loss of habitat is much more than would otherwise be assumed in response to climate change. Temperature increase is the predominant factor leading to habitat loss, whereas changes in precipitation play a secondary role. When climate changes, the species not only loses part of its habitat but also suffers a loss in its population size in the remaining habitat. Population size declines more than the habitat area under all considered climate change scenarios, which implies that the species might become extinct long before the complete loss of its habitat. This study suggests that some species might experience much more severe impacts from climate change than predicted from models of habitat area alone. Management policies based on predictions of habitat area decline using occurrence data need to be re‐evaluated and alternative measures need to be developed to conserve species in the face of rapid climate change.     

A method for spatial freshwater conservation prioritization

1. Freshwater ecosystems are amongst the most threatened and poorly protected globally. They continue to be degraded through habitat loss, pollution and invading species and conservation measures are urgently needed to halt declining trends in their biodiversity and integrity.
2. During the past decade a suite of decision support tools and computational approaches have been developed for efficient and targeted conservation action in terrestrial or marine ecosystems. These methods may be poorly suited for planning in freshwater systems because connectivity in terrestrial and marine systems is typically modelled in a way unsuitable for rivers, where connectivity has a strong directional component.
3. We modify the conservation prioritization method and software, zonation , to account for connectivity in a manner better suited to freshwater ecosystems. Prioritization was performed using subcatchment/catchment-based planning units and connectivity was modified to have directional upstream and downstream components consistent with the ecology of our target species.
4. We demonstrate this modified method for rivers and streams in the southern North Island of New Zealand. Data included predicted occupancy from boosted regression tree models of species distributions for 18 fish species. The study area covered 2.1 million hectares and included 394 first- to fourth order catchment or subcatchment planning units.
5. Realistic modelling of connectivity had a major influence on the areas proposed for conservation. If connectivity was ignored, recommended conservation areas were very fragmented. By contrast, when connectivity was modelled, high priority conservation targets consisted of entire river basins or headwater subcatchments.
6. The proposed method serves as a starting point for the implementation of reserve selection methods in river ecosystems.     

Incorporating climate change adaptation into national conservation assessments

The Convention on Biological Diversity requires that member nations establish protected area networks that are representative of the country's biodiversity. The identification of priority sites to achieve outstanding representation targets is typically accomplished through formal conservation assessments. However, representation in conservation assessments or gap analyses has largely been interpreted based on a static view of biodiversity. In a rapidly changing climate, the speed of changes in biodiversity distribution and abundance is causing us to rethink the viability of this approach. Here we describe three explicit strategies for climate change adaptation as part of national conservation assessments: conserving the geophysical stage, identifying and protecting climate refugia, and promoting cross‐environment connectivity. We demonstrate how these three approaches were integrated into a national terrestrial conservation assessment for Papua New Guinea, one of the most biodiverse countries on earth. Protected areas identified based on representing geophysical diversity were able to capture over 90% of the diversity in vegetation communities, suggesting they could help protect representative biodiversity regardless of changes in the distribution of species and communities. By including climate change refugia as part of the national conservation assessment, it was possible to substantially reduce the amount of environmental change expected to be experienced within protected areas, without increasing the overall cost of the protected area network. Explicitly considering environmental heterogeneity between adjacent areas resulted in protected area networks with over 40% more internal environmental connectivity. These three climate change adaptation strategies represent defensible ways to guide national conservation priority given the uncertainty that currently exists in our ability to predict climate changes and their impacts. Importantly, they are also consistent with data and expertise typically available during national conservation assessments, including in developing nations. This means that in the vast majority of countries, these strategies could be implemented immediately.     

气候速度影响下中国周边海域鲨鱼保护优先区识别

鲨鱼在气候变化和人类活动等因素的影响下面临着种群衰退的风险,开展鲨鱼保护优先区研究是鲨鱼保护行动的重要工作.将气候速度引入鲨鱼保护优先区的识别过程,旨在阐明中国周边海域鲨鱼现状保护成效和保护空缺,并预测气候速度影响下的鲨鱼保护优先区空间格局及其变化趋势.以集成物种分布模型模拟的146种鲨鱼栖息地作为保护对象,以2015年至2100年两种气候变化情景下的气候速度作为保护的机会成本,基于系统保护规划理论模拟现状和未来情景下的鲨鱼保护优先区选址方案.研究结果表明：（1）长江口以南至台湾海峡和北部湾近岸海域为鲨鱼多样性分布的主要区域,台湾海峡区域亦为珍稀濒危物种的重要分布区;（2）在两种气候情景下,南海中南部将面临较高的气候变化风险,而长江口以南至珠江口的近岸海域气候速度均相对较低,提示了这些区域或能成为气候变化影响下的生物避难所;（3）现有保护区仅保护了1.33%的海域和不到4%的鲨鱼物种,尚存在较大的保护空缺.当保护海域比例提升至10%时,可覆盖绝大多数鲨鱼物种.而当比例提升至30%时,珍稀濒危物种的栖息地将得到有效保护;（4）气候变化影响下保护优先区选址将发生不同程度的变化,尤其是在中国南海区域,如在保护规划时兼顾气候速度,可在满足相似保护目标的前提下减少保护优先区内25%以上的气候压力,以使其具有较强的应对气候变化潜力。     

祁连山国家公园青海片区雪豹栖息地适宜性评价

雪豹是全球生物多样性保护的旗舰物种,精准评估雪豹栖息地质量可为其种群保护提供科学参考。本文选取地形、气候、植被类型和人为干扰等环境因子,采用景观连接度和MaxEnt模型两种方法,对祁连山国家公园青海片区雪豹栖息地质量进行评价。结果表明：高适宜栖息地主要分布在研究区的西部和中部,适宜面积占青海片区总面积的71.8%～77.5%,两种方法预测的适宜栖息地重叠区为8 979.1 km2,占青海片区总面积的56.7%;39.8%～43.3%的适宜栖息地分布在一般控制区,56.7%～60.2%的适宜栖息地分布在核心保护区;疏勒河源区适宜栖息地面积最多,石羊河源区适宜栖息地面积最少,黑河源区适宜栖息地面积占比最高,约为77.1%～91.8%。适宜栖息地面积整体呈现自东向西递增的趋势。一般控制区分布有较大面积的雪豹栖息地,人类活动将是一般控制区雪豹栖息地的潜在威胁。本研究对优化雪豹就地保护措施提供科学借鉴与参考。     

Hydrological connectivity drives patterns of macroinvertebrate biodiversity in floodplain rivers of the Australian wet /dry tropics

1. Floodplain rivers in Australia's wet/dry tropics are regarded as being among the most ecologically intact and bio-diverse lotic ecosystems in the world, yet there have been relatively few community-based studies of their aquatic fauna.
2. To investigate relationships between hydrological connectivity and biodiversity in the region, macroinvertebrates were collected from sites within two contrasting floodplain rivers, the 'tropical' Gregory River and 'dryland' Flinders River systems, during the dry season and analysed at various spatial scales. A subset of sites was re-sampled in the following dry season to explore temporal variation. The fauna consisted of 124 morphotaxa, dominated by gatherers and the Insecta.
3. As predicted, hydrological connectivity (the lotic or lentic status of waterbodies) had a major influence on macroinvertebrate assemblage composition and diversity, both in space and time. Assemblages from waterbodies with similar connection histories were most alike, and beta-diversity between assemblages was greatest between lotic and lentic waterbodies, tending to increase with increasing spatial separation.
4. At smaller spatial scales, a number of within-waterbody, habitat and water quality characteristics were important for explaining variation (61%) in the taxonomic organization of assemblages, and characteristics associated with primary productivity and habitat diversity were important for explaining variation (45%) in the functional organization of assemblages. However, much of the small-scale environmental variation across the study region appeared to be related to broad-scale variation in hydrological connectivity, which had both direct and indirect effects on macroinvertebrate assemblages.
5. Conservation of the biodiversity in Australia's wet/dry tropics may depend on conserving the natural variation in hydrological connectivity and the unregulated flow of floodplain rivers.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.