Continuum or discrete patch landscape models for savanna birds? Towards a pluralistic approach

Conceptualising landscapes as a mosaic of discrete habitat patches is fundamental to landscape ecology, metapopulation theory and conservation biology. An emerging question in ecology is: when is the discrete patch model more appropriate than alternative and conceptually appealing models such as the continuum model? There is limited empirical testing of the utility of alternative landscape models compared to the discrete patch model for a range of species. In this paper, we constructed three alternative sets of models for testing the effect of landscape structure on diversity and abundance of a suite of woodland birds in a savanna landscape of northern Australia: the null model (only site‐scale habitat variables, landscape context not important), the continuum model, and the discrete patch model. We utilised high‐spatial resolution satellite images to quantify spatial gradients in tree cover density (the continuum model), and to then aggregate the fine‐scale heterogeneity in tree cover into discrete patches of trees, with grass cover forming the “matrix” (the discrete patch‐model). We then evaluated the relative importance of the alternative models using generalised linear models and an information theoretic approach. We found that the importance of the models varied among species, with no single model dominant. Species that move between open grassy areas and woody shelter responded well to the continuum model, reflecting the importance of gradients in density of forage (grasses) and cover (trees), while the discrete model performed best for species that forage in all vegetation strata, and nest predominantly in dense woody vegetation. This finding supports a pluralistic approach, highlighting the need for adopting and testing more than one landscape model in savanna landscapes, and in other landscapes that do not have a well defined patch structure.     

基于GIS的上海市景观格局梯度分析

 定量分析景观格局及其变化对于监测和评价城市化的生态后果十分重要。该研究应用基于GIS的梯度分析与景观指数相结合的方法定量分析了上海市城市化的空间格局。沿一条自西向东64 km长6 km宽和另一条自南向北66 km长6 km宽的样带,应用移动窗口计算了一系列景观指数。结果表明,城市化的空间格局可以用梯度分析与景观指数来定量,不同的土地利用类型沿景观格局梯度确实表现出明显的“空间特征”。多项景观指数可准确地、定量地指示上海地区东西和南北发展轴线上城市化的前沿和城市景观梯度分异的特征。城市化的总体格局是随着城市化程度增加,景观在组分上更多样化,形状上更为复杂以及生态学上更破碎化。此外,该文显示的上海市城市化景观格局比较符合同心圆论的城市发展理论,研究结果支持斑块密度随城市化程度而增加,斑块大小和景观连接度则下降的假说,但与随人类干预程度增强,斑块形状变得更规则的假说不相符和,因此,需要进一步研究来证实该发现。     

基于模拟景观的城市森林景观格局指数选取

基于现实的沈阳城市森林景观,模拟了4个景观格局梯度,并选取了1个与之相应的现实景观格局梯度,分析了28个景观格局指数在各梯度对景观破碎化和景观斑块形状复杂性的反映,从而筛选出描述这两种景观格局特征的适宜景观格局指数.结果表明：斑块密度（PD）和平均斑块面积（AREA_MN）在城市森林景观破碎化方面表现出较规律的变化趋势,斑块密度随破碎化程度的增加而增加,平均斑块面积随破碎化程度的增加而减小;面积加权平均周长面积比（PARA_AM）在描述景观斑块形状复杂性方面与景观格局梯度相吻合,且随斑块形状复杂性的增加而增加,能够较为准确地描述景观斑块形状的复杂性.     

Characterizing the spatial distribution of giant pandas (Ailuropoda melanoleuca) in fragmented forest landscapes

Aim To examine the effects of forest fragmentation on the distribution of the entire wild giant panda (Ailuropoda melanoleuca) population, and to propose a modelling approach for monitoring the spatial distribution and habitat of pandas at the landscape scale using Moderate Resolution Imaging Spectro‐radiometer (MODIS) enhanced vegetation index (EVI) time‐series data. Location Five mountain ranges in south‐western China (Qinling, Minshan, Qionglai, Xiangling and Liangshan). Methods Giant panda pseudo‐absence data were generated from data on panda occurrences obtained from the third national giant panda survey. To quantify the fragmentation of forests, 26 fragmentation metrics were derived from 16‐day composite MODIS 250‐m EVI multi‐temporal data and eight of these metrics were selected following factor analysis. The differences between panda presence and panda absence were examined by applying significance testing. A forward stepwise logistic regression was then applied to explore the relationship between panda distribution and forest fragmentation. Results Forest patch size, edge density and patch aggregation were found to have significant roles in determining the distribution of pandas. Patches of dense forest occupied by giant pandas were significantly larger, closer together and more contiguous than patches where giant pandas were not recorded. Forest fragmentation is least in the Qinling Mountains, while the Xiangling and Liangshan regions have most fragmentation. Using the selected landscape metrics, the logistic regression model predicted the distribution of giant pandas with an overall accuracy of 72.5% (κ = 0.45). However, when a knowledge‐based control for elevation and slope was applied to the regression, the overall accuracy of the model improved to 77.6% (κ = 0.55). Main conclusions Giant pandas appear sensitive to patch size and isolation effects associated with fragmentation of dense forest, implying that the design of effective conservation areas for wild giant pandas must include large and dense forest patches that are adjacent to other similar patches. The approach developed here is applicable for analysing the spatial distribution of the giant panda from multi‐temporal MODIS 250‐m EVI data and landscape metrics at the landscape scale.     

Fragmentation in the Legal Amazon,Brazil: Can landscape metrics indicate agricultural policy differences?

Large-scale land conversion for agriculture in Brazilian Amazonia is occurring at persistently high rates. Basin-wide net land use and land cover changes imply substantially different situations between distinct regions and states due to different agricultural policies. This research used eight landscape metrics to quantify and investigate the spatial patterns of cattle pasture and cropland throughout the states of Pará, Mato Grosso, Rondônia, and Amazonas. These metrics were patch density (DEN), mean patch size (MPS), largest patch index (LPI), mean edge density (MED), mean twist number (TWI), corrected perimeter-to-area ratio (CPA), fractal dimension (FDI), and fragmentation index (FRG). A total of 1852 patches were analyzed, originating from 86 samples in 71 different plots, covering a total of 177,500 km² throughout all four states.Principal component analysis showed a partial overlap in the spatial pattern of agricultural patches between all states. The largest percentage of variance was explained by patch area metrics, which can be related to the different approaches in agricultural policies, but no clear division between the states was identified in this dimension. The metrics quantifying patch shape were de facto independent of deforestation area, and related to the second principal component axis. Although some overlap in this dimension was present as well, these metrics proved a possible measure for discerning the patterns of agriculture attached to a certain state. Different land use policies are hypothesized to lead to more heterogeneity in landscape patterns in an early stage, yet the increasing influence of both cropland and pasture agriculture eventually leads to more uniform landscapes in which spatial differences gradually disappear.     

Evaluating the effectiveness of landscape metrics in quantifying spatial patterns

The effectiveness of landscape metrics in quantifying spatial patterns is fundamental to metrics assessment. Setting 36 simulated landscapes as sample space and focusing on 23 widely used landscape metrics, their effectiveness in quantifying the complexity of such spatial pattern components as number of patch types, area ratio of patch types and patch aggregation level, were analyzed with the application of the multivariate linear regression analysis method. The results showed that all the metrics were effective in quantifying a certain component of spatial patterns, and proved that what the metrics quantified were not a single component but the complexity of several components of spatial patterns. The study also showed a distinct inconsistency between the performances of landscape metrics in simulated landscapes and the real urban landscape of Shenzhen, China. It was suggested that the inconsistency resulted from the difference of the correlation among spatial pattern components between simulated and real landscapes. After considering the very difference, the changes of all 23 landscape metrics against changing of number of patch types in simulated landscapes were consistent with those in the real landscape. The phenomenon was deduced as the sign effect of spatial pattern components on landscape metrics, which was of great significance to the proper use of landscape metrics.     

Altitude effects on spatial components of vascular plant diversity in a subarctic mountain tundra

Environmental gradients are caused by gradual changes in abiotic factors, which affect species abundances and distributions, and are important for the spatial distribution of biodiversity. One prominent environmental gradient is the altitude gradient. Understanding ecological processes associated with altitude gradients may help us to understand the possible effects climate change could have on species communities. We quantified vegetation cover, species richness, species evenness, beta diversity, and spatial patterns of community structure of vascular plants along altitude gradients in a subarctic mountain tundra in northern Sweden. Vascular plant cover and plant species richness showed unimodal relationships with altitude. However, species evenness did not change with altitude, suggesting that no individual species became dominant when species richness declined. Beta diversity also showed a unimodal relationship with altitude, but only for an intermediate spatial scale of 1 km. A lack of relationships with altitude for either patch or landscape scales suggests that any altitude effects on plant spatial heterogeneity occurred on scales larger than individual patches but were not effective across the whole landscape. We observed both nested and modular patterns of community structures, but only the modular patterns corresponded with altitude. Our observations point to biotic regulations of plant communities at high altitudes, but we found both scale dependencies and inconsistent magnitude of the effects of altitude on different diversity components. We urge for further studies evaluating how different factors influence plant communities in high altitude and high latitude environments, as well as studies identifying scale and context dependencies in any such influences.     

Effects of spatial heterogeneity on butterfly species richness in Rocky Mountain National Park, CO, USA

We investigated butterfly responses to plot-level characteristics (plant species richness, vegetation height, and range in NDVI [normalized difference vegetation index]) and spatial heterogeneity in topography and landscape patterns (composition and configuration) at multiple spatial scales. Stratified random sampling was used to collect data on butterfly species richness from seventy-six 20 × 50 m plots. The plant species richness and average vegetation height data were collected from 76 modified-Whittaker plots overlaid on 76 butterfly plots. Spatial heterogeneity around sample plots was quantified by measuring topographic variables and landscape metrics at eight spatial extents (radii of 300, 600 to 2,400 m). The number of butterfly species recorded was strongly positively correlated with plant species richness, proportion of shrubland and mean patch size of shrubland. Patterns in butterfly species richness were negatively correlated with other variables including mean patch size, average vegetation height, elevation, and range in NDVI. The best predictive model selected using Akaike’s Information Criterion corrected for small sample size (AIC_c), explained 62% of the variation in butterfly species richness at the 2,100 m spatial extent. Average vegetation height and mean patch size were among the best predictors of butterfly species richness. The models that included plot-level information and topographic variables explained relatively less variation in butterfly species richness, and were improved significantly after including landscape metrics. Our results suggest that spatial heterogeneity greatly influences patterns in butterfly species richness, and that it should be explicitly considered in conservation and management actions.     

The spatial distribution of birds and carabid beetles in pine plantation forests: the role of landscape composition and structure

Aim To evaluate the joint and independent effects of spatial location, landscape composition and landscape structure on the distribution patterns of bird and carabid beetle assemblages in a mosaic landscape dominated by pine plantation forests. Location A continuous 3000‐ha landscape mosaic with native maritime pine Pinus pinaster plantations of different ages, deciduous woodlands and open habitats, located in the Landes de Gascogne forest of south‐western France. Methods We sampled breeding birds by 20‐min point counts and carabid beetles by pitfall trapping using a systematic grid sampling of 200 points every 400 m over the whole landscape. Explanatory variables were composed of three data sets derived from GIS habitat mapping: (1) spatial variables (polynomial terms of geographical coordinates of samples), (2) landscape composition as the percentage cover of the six main habitats, and (3) landscape structure metrics including indices of fragmentation and spatial heterogeneity. We used canonical correspondence analysis with variance partitioning to evaluate the joint and independent effects of the three sets of variables on the ordination of species assemblages. Moran's I correlograms and Mantel tests were used to assess for spatial structure in species distribution and relationships with separate landscape attributes. Results Landscape composition was the main factor explaining the distribution patterns of birds and carabids at the mesoscale of 400 × 400 m. Independent effects of spatial variables and landscape structure were still significant for bird assemblages once landscape composition was controlled for, but not for carabid assemblages. Spatial distributions of birds and carabids were primarily influenced by the amount of heathlands, young pine plantations, herbaceous firebreaks and deciduous woodlands. Deciduous woodland species had positive responses to edge density, while open habitat species were positively associated with mean patch area. Main conclusions Forest birds were favoured by an increase in deciduous woodland cover and landscape heterogeneity, but there was no evidence for a similar effect on carabid beetles. Fragmentation of open habitats negatively affected both early‐successional birds and carabids, specialist species being restricted to large heathlands and young plantations. Several birds of conservation concern were associated with mosaics of woodlands and grasslands, especially meadows and firebreaks. Conserving biodiversity in mosaic plantation landscapes could be achieved by the maintenance of a significant amount of early‐successional habitats and deciduous woodland patches within a conifer plantation matrix.     

Assessing the impacts of fragmentation on plant communities in New Zealand: scaling from survey plots to landscapes

Aim Few studies have attempted to assess the overall impact of fragmentation at the landscape scale. We quantify the impacts of fragmentation on plant diversity by assessing patterns of community composition in relation to a range of fragmentation measures. Location The investigation was undertaken in two regions of New Zealand – a relatively unfragmented area of lowland rain forest in south Westland and a highly fragmented montane forest on the eastern slopes of the Southern Alps. Methods We calculated an index of community similarity (Bray–Curtis) between forest plots we regarded as potentially affected by fragmentation and control forest plots located deep inside continuous forest areas. Using a multiple nonlinear regression technique that incorporates spatial autocorrelation effects, we analysed plant community composition in relation to measures of fragmentation at the patch and landscape levels. From the resulting regression equation, we predicted community composition for every forest pixel on land‐cover maps of the study areas and used these maps to calculate a landscape‐level estimate of compositional change, which we term ‘BioFrag’. BioFrag has a value of one if fragmentation has no detectable effect on communities within a landscape, and tends towards zero if fragmentation has a strong effect. Results We detected a weak, but significant, impact of fragmentation metrics operating at both the patch and landscape levels. Observed values of BioFrag ranged from 0.68 to 0.90, suggesting that patterns of fragmentation have medium to weak impacts on forest plant communities in New Zealand. BioFrag values varied in meaningful ways among landscapes and between the ground‐cover and tree and shrub communities. Main conclusions BioFrag advances methods that describe spatial patterns of forest cover by incorporating the exact spatial patterns of observed species responses to fragmentation operating at multiple spatial scales. BioFrag can be applied to any landscape and ecological community across the globe and represents a significant step towards developing a biologically relevant, landscape‐scale index of habitat fragmentation.     

中性景观模型与真实景观的一致性

采用RULE和SimMap中性景观模型,使其形成模拟景观的图幅大小、景观内各类型数目以及各类型之间的比例等项与真实景观相同,通过不同景观指标对中性模型系列的反应,看其在多大程度上代替真实景观．研究发现,中性景观模型在斑块数、斑块周长、聚集度、蔓延度以及孔隙度等指标所反映的格局特征方面,能很好地代替真实景观,而在校正斑块周长面积比、分维数以及边界分布均匀度等指标所反映的格局特征方面,并不能很好地代替真实景观,说明中性景观模型只能在一定的范围内可以代替真实景观。而不能完全取而代之.     

景观指数之间的相关分析

应用辽宁省1997~1998年的TM 5影像数据,编制了景观类型图,以78个县市区为单位,分割成78个景观,共计算39个景观格局指数,对它们进行了相关分析。总面积是最基本的景观指数,它决定景观总边界长度、斑块数、类型密度等基本指数,同时与多个指数有显著的相关关系(相关系数绝对值大于0.75)。形状指数的独立性强,极少数指数与其它指数有显著的相关关系;多样性指数和蔓延度指数之间信息重复量最多,都表示景观的异质性,但多样性指数以面积百分比表示景观异质性,而蔓延度指数以类型之间相邻边界的百分比表示景观异质性。研究发现,如果两个指数之间存在显著的相关关系,而由它们两个构成的指数与它们之间没有显著的相关关系。如果指数平均值之间存在显著的相关关系,则它们的变异系数之间不存在显著的相关关系。景观指数间的相关系数不仅与景观格局本身有关,还与空间尺度,分类系统、计算公式及其参数、计算单元和生态学意义关系密切。指数之间影响因子的相同之处越多,它们之间存在显著相关关系的概率越大。     

景观空间异质性及城市化关联——以江苏省沿江地区为例

空间异质性是揭示地理现象空间分异规律的重要标志.基于高分辨率卫星遥感影像数据,利用景观生态指数和统计学方法,构建景观空间异质性综合模型,定量测度了江苏省沿江地区景观空间异质性特征及城市化关联,并基于二者的相关性将研究区域划分为6个异质性梯度区,提出了应对城市化干扰的景观空间协调对策.结果表明:景观空间异质性综合模型是对景观斑块结构、形态、数量3个组分特征的反映,是揭示景观连通性、稳定性和破碎化的重要依据;景观空间异质性与3个城市化参数之间存在显著的正相关,城镇空间扩张模式和城镇人口的集聚强度对景观空间异质性影响较大;景观空间异质性梯度分异与区域开发强度、开发时序存在显著的耦合关系;同时,面对城市化的干扰,应该实施保护生态景观空间、减少建设空间与其它景观的接触概率和实施差别化的城市化战略等政策协调城市化发展与景观生态保护之间的矛盾.     

广州市主城区树冠覆盖景观格局梯度

城市树冠覆盖是城市森林在小尺度上的景观表达,是衡量城市森林生态功能服务质量的重要指标.通过在ArcGIS9.2中对广州市中心城区的高分辨率航片进行目视解译,对形成的树冠覆盖专题图分析并生成栅格数据,利用Fragstats3.3软件分别选择标准方法和移动窗口方法分析研究区景观格局特征.其中基于移动窗口法形成了一系列基于所选择格局特征的连续栅格表面,每一个栅格单元代表的是设定的移动窗口半径尺度下景观类型的格局特征.分析表明,研究区景观格局呈现明显的空间异质性梯度特征.基于样带区尺度效应分析的结果表明,粒度3-5m和移动窗口半径400-600m可适合于研究区的景观格局梯度分析,选择的景观指数能够形成较为平滑的栅格表面.基于移动窗口的分析结果以连续变量图的形式对景观格局空间异质性进行可视化表达,能够为小尺度上的城市森林景观格局优化提供重要的参考.     

基于移动窗口法的岷江干旱河谷景观格局梯度分析

以岷江干旱河谷为研究区,基于GIS技术和移动窗口法对其景观格局梯度变化进行分析,以期为区域的景观格局优化和管理提供支持。根据研究区的形状特征,分别沿干流和支流设置4条样带;选取景观水平下的景观指数,利用FRAGSTATS3.3软件分别采取标准法和移动窗口法获得不同尺度下的景观指数值;综合利用景观指数粒度效应分析、区域面积信息守恒评价方法和景观指数幅度效应评价曲线确定研究区景观格局梯度分析移动窗口尺寸,并计算了此窗口尺度下4条样带上的景观指数,得到沿样带方向的景观梯度格局。结果表明:岷江干旱河谷的景观基质是灌木林地,面积占73.82%。有林地和草地的景观异质性低,居民地和耕地斑块破碎化程度较大。确定了50m的栅格大小,250m的移动窗口尺寸为研究区景观格局梯度分析的窗口尺度;4条样带上各景观指数均随景观类型变化出现不同幅度的上下波动特征,梯度特征明显。处于景观类型过渡地带的区域,景观多样性和异质性增加,破碎化程度高;干旱河谷景观格局梯度变化主要受地形、水热因子,以及堤坝建设和土地开发利用影响。研究作为一种有益的尝试,更精细地分析了研究区的景观格局,能够为河谷地区景观格局量化分析提供参考,但也存在一定不足,需要在今后工作中继续深入研究。     

Empirical assessment of the relation between ecological connectivity and land complexity based on information-theoretic metrics

Habitat fragmentation and connectivity loss pose significant threats to biodiversity at both local and landscape levels. Strategies to increase ecological connectivity and preserve strong connectivity are important for dealing with the potential threat of habitat degradation. Various metrics have been used to measure (i.e., quantify) landscape composition and configuration in landscape ecology. However, their relationship with ecological connectivity must be understood to interpret landscape patterns comprehensively. In the present study, correlations between ecological connectivity and land complexity are examined based on information-theory metrics. Two primary questions are explored: (1) to what extent are landscape mosaic measures of entropy correlated with ecological connectivity, with landscape gradient-based measures, and with each other? (2) are landscape gradient-based entropy measures correlated with ecological connectivity more than discrete entropy measures? Results show that all information theoretic metrics are statistically significant (p < 0.05) for modelling ecological connectivity. Among categorically-based indices, the relationship between ECI and joint entropy was the most significant, while a generalized additive model indicated that Boltzmann entropy could predict the ecological connectivity index, explaining ∼60% of the variance. Therefore, configurational entropy can be used for improving ecological connectivity models.     

Using landscape metrics to analyze micro-scale soil erosion processes

Methods of recording soil erosion using photographs exist but they are not commonly considered in scientific studies. Digital images may hold an expressive amount of information that can be extracted quickly in different manners. The investigation of several metrics that were initially developed for landscape ecology analysis constitutes one method. In this study we applied a method of landscape metrics to quantify the spatial configuration of surface micro-topography and erosion-related features, in order to generate a possible complementary tool for environmental management. In a 3.7 m wide and 9.7 m long soil box used during a rainfall simulation study, digital images were systematically acquired in four instances: (a) when the soil was dry; (b) after a short duration rain for initial wetting; (c) after the first erosive rain; and (d) after the 2nd erosive rain. Thirteen locations were established in the box and digital photos were taken at these locations with the camera positioned at the same orthogonal distance from the soil surface under the same ambient light intensity. Digital photos were converted into bimodal images and seven landscape metrics were analyzed: percentage of land, number of patches, density of patches, largest patch index, edge density, shape index, and fractal dimension. Digital images were an appropriate tool because they can generate data very quickly. The landscape metrics were sensitive to changes in soil surface micro-morphology especially after the 1st erosive rain event, indicating significant erosional feature development between the initial wetting and first erosive rainfall. The method is considered suitable for spatial patterns of soil micro-topography evolution from rainfall events that bear similarity to landscape scale pattern evolution from eco-hydrological processes. Although much more study is needed for calibrating the landscape metrics at the micro-scale, this study is a step forward in demonstrating the advantages of the method.     

Towards a core set of landscape metrics for biodiversity assessments: A case study from Dadia National Park,Greece

Spatial heterogeneity has an important influence on a wide range of ecological patterns and processes, and many landscape metrics in GIS environment are used to facilitate the investigation of the relation between landscape structure and biodiversity. Data reduction analyses have been applied to tackle the problem of highly correlated indices, but valid landscape predictors for fine scale Mediterranean forest-mosaics are still missing. Therefore, we analyzed the landscape structure of Dadia National Park, Greece, a Mediterranean forest landscape of high biodiversity, characterized by pine, oak and mixed woodland. By distinguishing nine land cover classes, 119 variables were computed and factor analysis was applied to detect the statistical dimensions of landscape structure and to define a core set of representative metrics. At landscape level, diversity of habitats, fragmentation and patch shape and at class level dominance of mixed forest and the gradient from one pure forest type to another turned out to be the crucial factors across three different scales. Mapping the encountered dimensions and the representative metrics, we detected that the pattern of landscape structure in Dadia National Park was related to dominating habitat types, land use, and level of protection. The evaluated set of metrics will be useful in establishing a landscape monitoring program, to detect the local drivers of biodiversity, and to improve management decisions in Dadia NP and similar mosaic-landscapes.     

Untangling Natural Seascape Variation from Marine Reserve Effects Using a Landscape Approach

Distinguishing management effects from the inherent variability in a system is a key consideration in assessing reserve efficacy. Here, we demonstrate how seascape heterogeneity, defined as the spatial configuration and composition of coral reef habitats, can mask our ability to discern reserve effects. We then test the application of a landscape approach, utilizing advances in benthic habitat mapping and GIS techniques, to quantify this heterogeneity and alleviate the confounding influence during reserve assessment. Seascape metrics were quantified at multiple spatial scales using a combination of spatial image analysis and in situ surveys at 87 patch reef sites in Glover''s Reef Lagoon, Belize, within and outside a marine reserve enforced since 1998. Patch reef sites were then clustered into classes sharing similar seascape attributes using metrics that correlated significantly to observed variations in both fish and coral communities. When the efficacy of the marine reserve was assessed without including landscape attributes, no reserve effects were detected in the diversity and abundance of fish and coral communities, despite 10 years of management protection. However, grouping sites based on landscape attributes revealed significant reserve effects between site classes. Fish had higher total biomass (1.5×) and commercially important biomass (1.75×) inside the reserve and coral cover was 1.8 times greater inside the reserve, though direction and degree of response varied by seascape class. Our findings show that the application of a landscape classification approach vastly improves our ability to evaluate the efficacy of marine reserves by controlling for confounding effects of seascape heterogeneity and suggests that landscape heterogeneity should be considered in future reserve design.