A graph-based approach to investigating the influence of the landscape on population spread processes

Modelling of landscape connectivity is a key point in the study of the movement of populations within a given landscape. For studies focused on the preservation of biodiversity, graph-based methods provide an interesting framework to investigate the landscape influence on population spread processes. Such an approach is described here, based on the mapping of landscape categories in habitat patches, including a diachronic data set describing the population spread within the habitat patches. A minimum planar graph was built by computing spatial distances between all pairs of neighbouring patches. From this structure, two types of analysis are proposed: one focused on the links of the graph and consists in correlating spatial distances and gap indicators computed from the diachronic data. The other was based on the correlations between population data and connectivity metrics at the patch level. As an example, this approach was applied to the spread of the fossorial water vole on the Jura plateau (France), with annual population data covering eleven years from 1989 to 2000. Link analysis allowed to find an optimal set of resistance values used in the least-cost distances computations, and thus to build a relevant graph. From this graph, patch analysis displayed a cyclic correlation between a metric based on potential dispersal flux and the population density, outlining the strong role of landscape connectivity in the population spread. The present study clearly shows that landscape modelling and graph-based approach can produce parameters which are consistent with field observations and thus pave the way to simulating the effect of landscape modification on population dynamics.     

道路对林地景观连接度的影响——以巩义市为例

基于景观连接度原理,借用景观连接度指数,在地理信息系统支持下,探讨了巩义市山区林地景观在不同距离阈值下连接度的变化,定量分析了道路对林地景观连接度的影响。结果显示,随着景观距离阈值的增大,无论是否有道路,林地景观整体可能连通性指数值都表现为逐渐增大;对林地景观连接度起"非常高"和"高"作用的林地斑块数量都比较少,但占林地总面积比例较大,面积大的林地斑块在提高景观连接度中起的作用较大;道路的分割使得林地斑块重要值降低,就单一斑块而言,随着景观距离阈值的增大,分割成的小斑块的重要值降低程度在逐渐减小。     

Identifying potential conservation areas in the Cuitzeo Lake basin,Mexico by multitemporal analysis of landscape connectivity

Lake Cuitzeo basin is an important ecological area subjected to strong human pressure on forest covers that are key elements for the long-term support of biodiversity. We studied landscape connectivity changes for the years 1975, 1996, 2000, 2003 and 2008 to identify potential conservation areas in the basin. We modeled potential distributions of the Mexican bobcat (Lynx rufus escuinapae) and the ringtail (Bassariscus astutus) – two terrestrial mammal focal species with contrasting dispersal capacities – and we determined their habitat availability and suitability in the basin. We then identified their optimal habitat patches and produced landscape cumulative resistance maps, estimated least-cost paths (graph theory approach), and elaborated current flow maps (circuit theory approach). For evaluation of landscape connectivity, we applied an integral index of connectivity (IIC) to each study period, and determined individual habitat patch contribution to the overall landscape connectivity. The IIC index had very low values associated with reduced availability of focal species habitat. However, our study showed the conservation importance of the surface of optimal habitat patch areas. The combined application of a graph-based approach and current flow mapping were useful, and complementary both in terms of estimating potential dispersal corridors and identifying high probability dispersal areas. This indicated that landscape connectivity analysis is a useful tool for identification of potential conservation areas and for local landscape planning.     

Characterizing the importance of habitat patches in maintaining landscape connectivity for Tibetan antelope in the Altun Mountain National Nature Reserve,China

Connectivity of habitat patches is crucial for wildlife dispersal and survival, and identifying patches with high importance for maintaining connectivity can aid effective wildlife management. Knowledge of the habitat distribution of the Tibetan antelope in the Altun Mountain National Nature Reserve, which is essential for connectivity analysis, remains limited. We analyzed potential habitat distribution and priority patches using GIS-based habitat suitability modeling with three weighting factors and evaluated the connectivity of habitat patches under four dispersal distance scenarios. Patches with high habitat suitability covered 25.39 % of the total area, and these patches were selected for connectivity analysis as resource patches. Connectivity analysis indicated that, although the overall probability of connectivity (PC) showed an upward trend with increasing dispersal distance, the importance of each patch varied considerably under different dispersal distance scenarios. Transfer analysis of patch numbers between different importance levels revealed that the number of patches becoming less important was higher than the number of patches becoming more important when dispersal distance increased. In addition, nine patches covering 38.49 % of the suitable habitat area were identified as priority patches, in particular the patches near the Kardun and the Karchuka inspection stations. We also found that the habitat distribution of the Tibetan antelope obtained from the suitability model matched the population distribution determined by a field survey. Correlation analysis between patch area and the percentage of PC index value loss (dPC) revealed that the larger patches in this region were more likely to be important for maintaining connectivity.     

基于景观连接度的土地整理生态效应评价

近年来,土地整理区的生物多样性保护和景观功能的维持受到广泛关注。本文选取贵州省道真县洛龙镇国家重点土地整理项目区,运用基于结构与功能相结合景观连接度模型(综合连接度(IIC)和连接度概率(PC)),计算了500、1000、2000、3000和4000m种群迁移扩散距离阈值情景下,土地整理前后各生境斑块对景观连接度的重要程度,识别重要斑块变化范围,从功能角度评价整理后的生态效应。结果表明:连接度指数能较好地识别土地整理前后各生境斑块的重要类别及其改变;其中,巨型斑块所占面积最大,分别占总面积的57.0%和57.2%;小型斑块数目整理前后分别占斑块总数的48.0%和48.9%;空间上,斑块的重要性分布也产生一定的变化。整理后生境斑块的连通性较整理前也有所下降,IIC和PC平均值分别为0.434和0.680,且随着距离阈值的增大,连接度逐渐增大。巨型斑块对生境斑块景观连通性的相对重要值最大,中型斑块和大型斑块次之,小型斑块最低。研究表明,土地整理过程中,大面积生境斑块和"踏脚石"斑块对维持土地整理生态效益具有重要作用。     

Forest pattern dynamics and landscape connectivity changes in the Manwan Basin after dam construction in the Lancang River,China

Regional habitat quality and landscape connectivity can be greatly impaired by dam construction and subsequent land-use changes. This study examined landscape pattern and connectivity changes in the Manwan Basin from 1974 to 2006 to quantify the effects of dam construction. Landscape pattern indices and kernel density estimation (KDE) were used to analyze forest pattern dynamics, while the probability of connectivity (PC) index was applied to evaluate landscape connectivity changes. The results showed that forest area decreased by nearly 20 %, while the number of forest patches increased more than sixfold from 1974 to 2006 in the Manwan Basin. During the study period, forest fragmentation mainly occurred in the western and northern parts of the Manwan Basin. Due to the changes in the forest landscape pattern, landscape connectivity decreased by an average of 54.74 %. In addition, we detected changes to key stepping-stone patches by calculating the percentage importance of each resource patch. The results showed that 300.38 km² of high-importance forest patches transformed to medium- and low-importance patches from 1974 to 2006.     

Network analysis to assess landscape connectivity trends: Application to European forests (1990–2000)

Landscape networks and ecosystems worldwide are undergoing changes that may impact in different ways relevant ecological processes such as gene flow, pollination, or wildlife dispersal. A myriad of indices have been developed to characterize landscape patterns, but not all of them are equally suited to evaluate temporal changes in landscape connectivity as is increasingly needed for biodiversity monitoring and operational indicator delivery. Relevant advancements in this direction have been recently proposed based on graph theoretical methods to analyze landscape network connectivity and on the measurement of habitat availability at the landscape scale. Building from these developments, we modify a recent index and present the equivalent connected area (ECA) index, defined as the size of a single patch (maximally connected) that would provide the same probability of connectivity than the actual habitat pattern in the landscape. The temporal changes in ECA can be directly compared with the changes in total habitat area. This allows for additional and straightforward insights on the degree to which the gains or losses in habitat amount can be beneficial or deleterious by affecting landscape elements that uphold connectivity in a wider landscape context. We provide a demonstrative example of application and interpretation of this index and approach to monitor changes in functional landscape connectivity. We focus on the trends in European forests at the province level in the period 1990–2000 from Corine land cover data, considering both changes in the forest spatial pattern and in the average permeability of the landscape matrix. The degree of connectivity was rather stable over most of the study area, with a slight overall increase in forest connectivity in Europe. However, a few countries and regions concentrated remarkably high changes in the analyzed period, particularly those with a low forest cover. The species traits also affected the responses to landscape pattern changes, which were more prominent for those species with limited dispersal abilities. We conclude discussing the potential of this approach for consistent indicator delivery, as well as the limitations and possibilities of application to a variety of situations, for which the required quantitative tools are freely available as open source projects.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Landscape dynamics and their effect on the functional connectivity of a Mediterranean landscape in Chile

Land use and cover changes have been identified as a major factor contributing to shape landscape structure and biodiversity patterns, particulary in areas with a long history of human occupation and habitat fragmentation, such as the Mediterranean landscapes. However, the existing studies on landscape change indicators for Mediterranean areas have mostly focused in Europe, while for other Mediterranean zones, and especially for South America, there is a serious lack of knowledge concerning the impact of landscape dynamics on ecological processes. Further research on this topic is urgently needed, given the high biodiversity levels and the rapidly increasing rates of human modification in the Mediterranean landscapes of South America. For this purpose, we investigated the dynamics of a landscape in the semiarid region of the Mediterranean zone of Chile, and measured the effect of those dynamics on functional connectivity, during a period of about four decades (1975–2011). Landscape connectivity indicators were extracted from a series of Landsat images. The Equivalent Connnected Area index (ECA) was used as indicator of connectivity trends, and was evaluated for three representative distances of seed dispersal in the study area (150 m, 500 m and 1000 m). In addition, the patches that most contribute to maintain the present connectivity, and their roles as connectivity providers, were identified through a set of commensurable indicators: betweenness centrality and the fractions (intra, flux connector) of the Integral Index of Connectivity. We found that these indicators were useful to detect and summarize a number of previously unreported trends in these Mediterranean landscapes. First, population growth and economic development were compatible with an increase in functional connectivity for forest habitats, mainly because the abandonment of marginal agricultural lands and their subsequent conversion to espinals (Acacia caven) triggered vegetation succession towards secondary forests. Second, increased forest connectivity was not associated to a decrease in the characteristic heterogeneity of Mediterranean landscapes. Third, many patches of espinal, despite being commonly regarded as of poor conservation value, were crucial to promote connectivity by acting as stepping stones among other patches with higher habitat quality. The approach here presented provides a combined assessment of landscape structure, function and change that should be valuable and applicable to deliver operational indicators in dynamic landscapes in South America and other Mediterranean regions.     

Forest species in an agricultural landscape in the Netherlands: Effects of habitat fragmentation

Abstract. For 312 forest patches on sandy soils in the Netherlands, effects of fragmentation are studied of forest habitat in the past on the present occurrence of forest plant species. Using regression techniques, the numbers of forest edge, interior, zoochorous and anemochorous species, as well as occurrence of 24 individual species were related to patch area and connectivity measures. Connectivity was defined as the amount of forest habitat around patches within three zones up to 1000 m. Plant categories were distinguished by habitat type and dispersal mechanism. The results showed that number of total species and number of species of all habitat and dispersal categories increased with area. The occurrence of ten individually studied species were also positively related to area. Most of them were interior species. The number of zoochorous species increased with increasing connectivity. Also occurrence of ten individually studied species were affected by connectivity. Interior zoochorous species showed the highest percentage of affected species. The relationship of interior, animal-dispersed plants to connectivity can be explained by the limited distances covered by their dispersal agents (forest birds and ants) in a non-forest habitat. Also, some anemochorous plants appeared to be affected by connectivity, especially those with heavy seeds and potentially short distance dispersal. As not all species within a certain dispersal or habitat category react similar to area or isolation, it is suggested that differences in underlying processes of fragmentation such as local extinction and colonization need more focus.     

Landscape connectivity and the role of small habitat patches as stepping stones: an assessment of the grassland biome in South America

Connectivity losses lead to a reduction of the amount of habitat resources that can be reached and used by species, and hence to a decline in the ranges and abundance of multiple taxa. Despite the recognized important role of small habitat patches for many species inhabiting fragmented landscapes, their potential contribution as stepping stones for maintaining overall landscape connectivity has received less attention. Using connectivity metrics based on a graph-theoretic approach we (i) quantified the connectivity of grassland patches in a sector of the Pampa region in Argentina, using a range of dispersal distances (from 100 to 10,000 m) representative of the scale of dispersal of different species; (ii) identified the most relevant patches for maintaining overall connectivity; and (iii) studied the importance of small patches (defined for different area thresholds of 5, 20, and 50 ha) as connectivity providers in the landscape. Although grassland patches were in general poorly connected at all distances, some of them were critical for overall connectivity and were found to play different crucial roles in the patch network. The location of small patches in the grassland network allowed them to function as stepping stones, yielding significant connectivity gains for species that move large distances (>5000 m) for the three area thresholds considered. Thus, under the spatial pattern of the studied landscape, species that move long distances would benefit from stepping stones, while less mobile organisms would benefit from, and mostly rely on the largest patches. We recommend that future management activities should (i) aim at preserving the grassland patches with the highest potential as stepping stones to promote landscape-level connectivity; and (ii) pay more attention to the conservation of key small patches, particularly given that usually they are those more vulnerable to land clearing for agriculture.     

Gene flow and functional connectivity in the natterjack toad

Functional connectivity is a key factor for the persistence of many specialist species in fragmented landscapes. However, connectivity estimates have rarely been validated by the observation of dispersal movements. In this study, we estimated functional connectivity of a real landscape by modelling dispersal for the endangered natterjack toad (Bufo calamita) using cost distance. Cost distance allows the evaluation of 'effective distances', which are distances corrected for the costs involved in moving between habitat patches in spatially explicit landscapes. We parameterized cost-distance models using the results of our previous experimental investigation of natterjack's movement behaviour. These model predictions (connectivity estimates from the GIS study) were then confronted to genetic-based dispersal rates between natterjack populations in the same landscape using Mantel tests. Dispersal rates between the populations were inferred from variation at six microsatellite loci. Based on these results, we conclude that matrix structure has a strong effect on dispersal rates. Moreover, we found that cost distances generated by habitat preferences explained dispersal rates better than did the Euclidian distances, or the connectivity estimate based on patch-specific resistances (patch viscosity). This study is a clear example of how landscape genetics can validate operational functional connectivity estimates.     

Effects of landscape matrix on population connectivity of an arboreal mammal,Petaurus breviceps

Ongoing habitat loss and fragmentation is considered a threat to biodiversity as it can create small, isolated populations that are at increased risk of extinction. Tree‐dependent species are predicted to be highly sensitive to forest and woodland loss and fragmentation, but few studies have tested the influence of different types of landscape matrix on gene flow and population structure of arboreal species. Here, we examine the effects of landscape matrix on population structure of the sugar glider (Petaurus breviceps) in a fragmented landscape in southeastern South Australia. We collected 250 individuals across 12 native Eucalyptus forest remnants surrounded by cleared agricultural land or exotic Pinus radiata plantations and a large continuous eucalypt forest. Fifteen microsatellite loci were genotyped and analyzed to infer levels of population differentiation and dispersal. Genetic differentiation among most forest patches was evident. We found evidence for female philopatry and restricted dispersal distances for females relative to males, suggesting there is male‐biased dispersal. Among the environmental variables, spatial variables including geographic location, minimum distance to neighboring patch, and degree of isolation were the most important in explaining genetic variation. The permeability of a cleared agricultural matrix to dispersing gliders was significantly higher than that of a pine matrix, with the gliders dispersing shorter distances across the latter. Our results added to previous findings for other species of restricted dispersal and connectivity due to habitat fragmentation in the same region, providing valuable information for the development of strategies to improve the connectivity of populations in the future.     

Influence of environmental and spatial variables on regional distribution of forest plant species in a fragmented and changing landscape

During the past several centuries, forests in Europe and large parts of North America have been subject to extensive forest clearance. The last several decades, however, numerous new forest patches have been established onto former agricultural land. As a result, the present forest area often consists of a mixture of small forest patches of different age, area, habitat quality and connectivity embedded within a hostile agricultural landscape. In these patchy landscapes, distribution patterns of plant species may be affected by both regional and local factors, although the relative importance of both is still poorly understood. In this study, we investigated distribution patterns of 113 forest plant species in a fragmented landscape. Species abundances at the regional scale conformed to a clearly unimodal abundance distribution which we believe to be related to 1) environmental heterogeneity due to succession and 2) inequality in migration rates. Patch incidence was significantly related to life form, which in turn was correlated to seed mass and dispersal mechanism. Multiple logistic regressions showed that presence/absence of 59 species studied was significantly affected by patch connectivity, patch area and age for 35, 30 and 34 species, respectively. The results of this study indicate that distribution patterns of forest plant species are influenced by both local and regional factors. Moreover, they also demonstrate that next to spatial aspects of fragmentation, temporal patterns of landscape change may have far-reaching effects on presence/absence patterns of plant species and therefore should be incorporated in studies dealing with regional population structures of plants.     

Patch quality and connectivity influence spatial dynamics in a dune wolfspider

The spatial population dynamics of the wolfspider Pardosa monticola, inhabiting patchily distributed grasslands in the Flemish coastal dunes of Belgium and Northern France were investigated with incidence function models using field survey data from 1998 and 2000. Vegetation height and patch size were related to habitat quality. Mark-recapture experiments revealed maximum cursorial dispersal distances of 280 m for moss dunes and 185 m for higher dune grassland. Higher shrub vegetation appeared to be dispersal barriers. These habitat-dependant cursorial distances and the theoretically estimated ballooning distance were included with patch distances into a connectivity index for both dispersal modes. Forward multiple regression indicated that patch occurrence was influenced by habitat quality and ballooning connectivity. Habitat quality and cursorial connectivity explained patterns in short-term colonisation. Extinction appeared to be stochastic and not related to habitat quality and connectivity. Genetic differentiation and variability was low. The discrepancy between the estimated low dispersal capacity and the indirect estimate of gene flow ( F(ST)) indicates that historical population dynamics and/or historical ballooning dispersal influence the genetic structure in this species.     

Functional connectivity as an indicator for patch occupancy in grassland specialists

Habitat specialists living in metapopulations are sensitive to habitat fragmentation. In most studies, the effects of fragmentation on such species are analyzed based on Euclidean inter-patch distances. This approach, however, ignores the role of the landscape matrix. Recently, therefore, functional distances that account for the composition of the landscape surrounding the habitat patches have been used more frequently as indicators for patch occupancy. However, the performance of functional and non-functional connectivity measures in predicting patch occupancy of such species has never been compared in a multi-species approach.Here we evaluate the effect of habitat connectivity on the patch occupancy of 13 habitat specialists from three different insect orders (Auchenorrhyncha, Lepidoptera, Orthoptera) in fragmented calcareous grasslands. In order to calculate functional distances we used four different sets of resistance values and rankings. We then modelled species’ occurrence using both Euclidean and functional (based on least-cost modelling) inter-patch distances as predictors.We found that functional connectivity measures provided better results than the non-functional approach. However, a functional connectivity measure that was based on very coarse land-cover data performed even better than connectivity measures that were based on much more detailed land-use data.In order to take into account possible effects of the landscape matrix on patch occupancy by habitat specialists, future metapopulation studies should use functional rather than Euclidean distances whenever possible. For practical applications, we recommend a ‘simple approach’ which requires only coarse land-cover data and in our study performed better than all other functional connectivity measures, even more complex ones.     

Movements of the flying squirrel Pteromys volans in corridors and in matrix habitat

Elements of the landscape, such as patches of preferred habitat, matrix between patches, and corridors linking patches, differ as movement habitat for animals. To understand how landscape structure influences the movement and thus, population dynamics of animals, clear empirical knowledge on patterns of movement is needed. The Siberian flying squirrel inhabits spruce-dominated boreal forests from Finland to eastern Siberia. Numbers of flying squirrels have declined severely in Finland in past decades, probably due to modern forestry. We studied the movement of radio-collared adult flying squirrels in preferred (spruce forest) and in matrix habitat (open areas and other habitats with trees) in Finland 1997–2000, and determined whether the woodland strips connecting patches of preferred habitat could function as ecological corridors for flying squirrels.
Flying squirrels used woodland strips for inter-patch movements, but also used matrix with trees and were able to cross narrow open gaps. Males moved longer total distances and crossed edges more often than females. Males used matrix habitats for movement between spruce patches, and moved faster and more directly in the matrix than in the spruce forest. Females seldom changed spruce patches, but instead used the matrix for foraging. For both sexes probability of leaving the spruce forest patch correlated negatively with the size of the patch, but the type of connection the patch had to other patches did not affect the leaving probability. Due to efficient movement abilities of the flying squirrel and forest-dominated landscape structure of southern Finland, we suggest that conservation acts for maintaining viable populations of flying squirrels should focus on the quality of managed forest and the area of suitable breeding habitat (i.e. on habitat loss), but not necessarily on ecological corridors.     

Grassland connectivity explains entomophilous plant species assemblages in an agricultural landscape of the Pampa Region,Argentina

The Pampa grassland of Argentina is one of the most highly threatened biomes in the world. A high proportion of the original grassland cover has been transformed into land for agriculture or degraded. In the southern part of the region, fragmented semi‐natural grasslands over exposed rock still persist and connectivity between them is assumed to be crucial for maintaining viable populations. We quantified overall connectivity of grassland patches in a sector of the Southern Pampa region, and investigated the degree to which landscape connectivity explains entomophilous plant species assemblages in a subset of patches. We characterized each of the 301 patches in the landscape by their degree of intra‐patch and inter‐patch connectivity based on graph theory, and considering threshold dispersal distances from 100 to 1000 m. We surveyed entomophilous plant species in 39 grassland patches and classified the species in three categories (annual herbs, perennial herbs and shrubs) considering their different growth form and longevity. The influence of connectivity variables on entomophilous plant species assemblages variation was explored using Canonical Correspondence Analysis. Although grassland patches were poorly connected at all threshold distances, some of them were found to be critical for global connectivity. Connectivity significantly explained total, annual‐biennial and shrub assemblages for all threshold dispersal distances (6–13% of total variation). Variation in annual species assemblages was associated with intra‐patch and inter‐patch connectivity at short distance (100 m), while variation in shrub species assemblages was explained by intra‐patch and inter‐patch connectivity for distances between 100 m and 1000 m. This study evidenced the low connectivity of the study system, allowed the identification of critical areas for conservation, and provided valuable information to develop management strategies in increasingly human‐dominated landscapes.     

Dispersal potential of spores and asexual propagules in the epixylic hepatic Anastrophyllum hellerianum

Dispersal ability is of great importance for plants, which commonly occupy spatially and temporally limited substrate patches. Mixed reproductive strategies with abundant diaspore production are favoured in a heterogeneous landscape to ensure successful colonisation at different distances. In bryophytes, long-distance dispersal has been thought to take place primarily by spores, while asexual propagules are important in local dispersal and in the maintenance of colonies. In the present study, we investigated the dispersal potential of two equally sized propagules, sexually formed spores and asexually produced gemmae in the dioecious, epixylic hepatic, Anastrophyllum hellerianum, which inhabits spatially and temporally limited substrate patches. We trapped propagules at different distances (0–10 m) and directions from the source colonies in two experiments: one in a natural habitat within a forest and another involving an artificial set-up in an open habitat. Spore dispersal showed only slight distance dependence both in the open and the forest habitats, presumably as a consequence of wind affecting the dispersal pattern. Gemma dispersal was more strongly distance-dependent in the open habitat than in the forest sites. Considerably more gemmae were deposited during rainy than dry periods, possibly because of the effect of rain drops on gemma release. However, weather conditions had no effect on the dispersal patterns of spores or gemmae. In A. hellerianum, the combination of occasional spore production and practically continuous, massive gemma production facilitates dispersal both on local scale and over long distances. Unlike previously assumed, not only spores but also the asexual propagules may contribute to long-distance dispersal, thus allowing considerable gene flow at the landscape level.     

Relationships between spatial configuration of tropical forest patches and woody plant diversity in northeastern Puerto Rico

The destruction and fragmentation of tropical forests are major sources of global biodiversity loss. A better understanding of anthropogenically altered landscapes and their relationships with species diversity and composition is needed in order to protect biodiversity in these environments. The spatial patterns of a landscape may control the ecological processes that shape species diversity and composition. However, there is little information about how plant diversity varies with the spatial configuration of forest patches especially in fragmented tropical habitats. The northeastern part of Puerto Rico provides the opportunity to study the relationships between species richness and composition of woody plants (shrubs and trees) and spatial variables [i.e., patch area and shape, patch isolation, connectivity, and distance to the Luquillo Experimental Forest (LEF)] in tropical forest patches that have regenerated from pasturelands. The spatial data were obtained from aerial color photographs from year 2000. Each photo interpretation was digitized into a GIS package, and 12 forest patches (24–34 years old) were selected within a study area of 28 km². The woody plant species composition of the patches was determined by a systematic floristic survey. The species diversity (Shannon index) and species richness of woody plants correlated positively with the area and the shape of the forest patch. Larger patches, and patches with more habitat edge or convolution, provided conditions for a higher diversity of woody plants. Moreover, the distance of the forest patches to the LEF, which is a source of propagules, correlated negatively with species richness. Plant species composition was also related to patch size and shape and distance to the LEF. These results indicate that there is a link between landscape structure and species diversity and composition and that patches that have similar area, shape, and distance to the LEF provide similar conditions for the existence of a particular plant community. In addition, forest patches that were closer together had more similarity in woody plant species composition than patches that were farther apart, suggesting that seed dispersal for some species is limited at the scale of 10 km.