Avifaunal responses to habitat fragmentation in the threatened littoral forests of south-eastern Madagascar

Aim Madagascar's lowland forests are both rich in endemic taxa and considered to be seriously threatened by deforestation and habitat fragmentation. However, very little is known about how these processes affect biodiversity on the island. Herein, we examine how forest bird communities and functional groups have been affected by fragmentation at both patch and landscape scales, by determining relationships between species richness and individual species abundance and patch and landscape mosaic metrics. Location Littoral forest remnants within south‐eastern Madagascar. Methods We sampled 30 littoral forest remnants in south‐eastern Madagascar, within a landscape mosaic dominated by Erica spp. heathland. We quantified bird community composition within remnants of differing size, shape and isolation, by conducting point counts in November–December in 2001 and October–November 2002. Each remnant was characterized by measures of remnant area, remnant shape, isolation, and surrounding landscape complexity. We used step‐wise regression to test the relationship between bird species richness and landscape structural elements, after correcting for sampling effort. Relationships between bird species abundances and the landscape variables were investigated with Canonical Correspondence Analysis and binomial logistic regression modelling. Results Bird species richness and forest‐dependent bird species richness were significantly (P < 0.01) explained by remnant area but not by any measure of isolation or landscape complexity. The majority of forest‐dependent species had significant relationships with remnant area. Minimum area requirements for area‐sensitive species ranged from 15 to 150 ha, with the majority of species having area requirements > 30 ha. Surprisingly, there was no relationship between bird body size and minimum area requirement. Forest‐dependent canopy insectivorous species and large canopy frugivorous species were the most sensitive functional groups, with > 90% species sensitivity within each group. The distribution of four forest‐dependent species also appeared to be related to remnant shape where remnant area was < 100 ha. Main conclusions The majority of forest‐dependent species, including many that are considered widespread and common, were found to have significant relationships with fragment size, indicating that they are sensitive to processes associated with habitat loss and fragmentation. As deforestation and habitat fragmentation remain serious problems on the island, it follows that many forest‐dependent bird species will decline in abundance and become locally extinct. At the regional scale, we urge that large (> 200 ha) blocks of littoral forest are awarded protected status to preserve their unique bird community.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

The geometry of habitat fragmentation: Effects of species distribution patterns on extinction risk due to habitat conversion

Land‐use changes, which cause loss, degradation, and fragmentation of natural habitats, are important anthropogenic drivers of biodiversity change. However, there is an ongoing debate about how fragmentation per se affects biodiversity in a given amount of habitat. Here, we illustrate why it is important to distinguish two different aspects of fragmentation to resolve this debate: (a) geometric fragmentation effects, which exclusively arise from the spatial distributions of species and habitat fragments, and (b) demographic fragmentation effects due to reduced fragment sizes, and/or changes in fragment isolation, edge effects, or species interactions. While most empirical studies are primarily interested in quantifying demographic fragmentation effects, geometric effects are typically invoked as post hoc explanations of biodiversity responses to fragmentation per se. Here, we present an approach to quantify geometric fragmentation effects on species survival and extinction probabilities. We illustrate this approach using spatial simulations where we systematically varied the initial abundances and distribution patterns (i.e., random, aggregated, or regular) of species as well as habitat amount and fragmentation per se. As expected, we found no geometric fragmentation effects when species were randomly distributed. However, when species were aggregated, we found positive effects of fragmentation per se on survival probability for a large range of scenarios. For regular species distributions, we found weakly negative geometric effects. These findings are independent of the ecological mechanisms which generate nonrandom species distributions. Our study helps to reconcile seemingly contradictory results of previous fragmentation studies. Since intraspecific aggregation is a ubiquitous pattern in nature, our findings imply widespread positive geometric fragmentation effects. This expectation is supported by many studies that find positive effects of fragmentation per se on species occurrences and diversity after controlling for habitat amount. We outline how to disentangle geometric and demographic fragmentation effects, which is critical for predicting the response of biodiversity to landscape change.     

Role of movement,interremnant dispersal and edge effects in determining sensitivity to habitat fragmentation in two forest‐dependent rodents

Abstract Habitat fragmentation and disturbance affect patterns of habitat use, animal movement and spatial behaviour and might have significant effects upon population dynamics and trends, and ultimately population persistence. Previous studies have suggested that the ability to disperse between remnants and a positive or neutral response to edges should be associated with species capable of persisting in remnant habitat. Using both radiotracking and trapping data, movement patterns, dispersal and response to habitat edges of Rattus fuscipes were examined within forests, corridors, remnants and pastures in south‐east Queensland, Australia. Rattus fuscipes has previously been shown to be robust to the effects of habitat fragmentation; however, contrary to expectations, R. fuscipes was found to be sensitive to edges, and no evidence of interremnant dispersal was detected, despite interremnant distances that were substantially smaller than the distances R. fuscipes was found to move in continuous habitat. Using only trapping data, the same factors were examined in relation to Melomys cervinipes, a species sensitive to fragmentation. Melomys cervinipes was found to utilize edge habitat, but no evidence of interremnant dispersal was detected, although the capacity to detect such movement was limited by low abundance in remnants where M. cervinipes was extant, and the species absence from many remnants. Movement patterns, interremnant dispersal capacity, and sensitivity to edges did not prove to be good predictors of these species responses to habitat fragmentation. Alternative explanations, such as population fluctuation and the capacity for rapid population growth in remnants for these two species, and the influence habitat quality has on these parameters should be investigated.     

Quantifying the activity levels and behavioural responses of butterfly species to habitat boundaries

1. The ability of species' to undergo climate‐driven range shifts across fragmented landscapes depends on their dispersal ability as well as the structure of the landscape. For species' range shifts to occur, individuals must first leave suitable habitat to seek new habitat; this is likely to depend on the rate of movement of individuals within habitat and the likelihood that a boundary is crossed, once it is encountered. For three species of butterfly with contrasting histories of recent range expansion, we examined the propensity of individuals to move within a habitat and their responses to habitat boundaries. 2. We quantified the extent to which Plebejus argus (Linnaeus) (a declining habitat specialist), Aricia agestis (Schiffermuller) (an expanding generalist) and Polymmatus icarus (Rottemburg) (a geographically ubiquitous generalist) crossed habitat boundaries into unsuitable habitat and moved within suitable habitat. The observed movement was then related to individual and environmental conditions. 3. Species differed in their activity levels in accordance within their recent distribution patterns (P. icarus > A. agestis > P. argus). Our results for P. argus suggest that movement may be motivated by nectar‐seeking, and that males generally move more than females. All three species tended to avoid crossing habitat boundaries; however the proportion of individuals crossing habitat boundaries did not differ significantly among species. 4. We conclude that levels of activity within a habitat, which will affect the frequency with which individuals encounter habitat boundaries, rather than behavioural responses to the boundaries, may be important drivers of distribution change.     

Differential and delayed response of two ant species to habitat fragmentation via the introduction of a pine matrix

1. Ants are a ubiquitous and crucial component of Australian Eucalyptus forests, but responses to long‐term habitat fragmentation remain poorly understood. Two ant species were followed across a 21‐year history of pine plantation establishment and maturation in a southeast Australian Eucalyptus forest. 2. At Wog Wog in southeastern Australia, a Native Eucalyptus forest was clear‐cut to make way for plantation establishment and 12 remnant patches of forest were left intact and subsequently surrounded by a pine matrix. Pitfall traps were placed in the continuous native forest, remnant Eucalyptus patches, and the pine matrix between fragments, and were stratified based on proximity to remnant patch edges and habitat type. Two ant species are focused on that represent the only remaining data for the early years of the experiment. 3. While Leptomyrmex erythrocephalus (Fabricius), the rarer of the two species, was not affected by fragmentation in the short term, 21 years after fragmentation, it was less likely to occur in both the mature pine matrix and fragments than in continuous forest controls. Aphaenogaster longiceps (Smith F.) was equally likely to occur in the fragments, continuous forest, and pine matrix early in the experiment but by year 21 post‐fragmentation was less likely to occur in the pine matrix than fragments or controls. 4. Importantly, we only detected negative impacts of fragmentation on ant occurrence as the pine plantation matrix matured and isolated ant populations on fragments. 5. It is concluded that changes in matrix suitability and specific habitat characteristics influence ant persistence in Eucalyptus fragments.     

Avian responses to forest fragmentation during the breeding and non-breeding seasons

Forest fragmentation represents a threat to several bird species worldwide. Several factors can change across seasons (e.g. bird perception of the landscape, weather conditions, biotic interactions), which can modify the response of bird populations to forest fragmentation. However, most studies have been conducted only during the breeding season. Here we assessed the relationship between forest fragmentation (patch area and patch isolation) with population abundances of resident species during both the breeding and the non-breeding seasons. Bird population abundances (all species in the community, subsets of forest and habitat generalist species and for individual species) were estimated across a gradient of area-isolation in a semi-arid forest in Cordoba, Argentina. Population abundance of the overall avian community and of the subset of forest species declined with patch area reduction independently of the season. By contrast, the subset of habitat generalist species was not affected by patch area reduction or by the increase in patch isolation, either during the breeding or during the non-breeding season. When the analyses were carried out for individual species, we found four forest species and one habitat generalist species whose responses (the relationship between population abundance and patch area or with isolation) were different between breeding and non-breeding seasons. The negative effects of forest fragmentation were found mainly during the breeding season. Our results suggest that reduction of patch area may lead to a reduction of more than 65% of the population abundance of forest bird species, during both the breeding and the non-breeding season. Therefore, there is an urgent need to conserve large forest patches within the region as irreplaceable elements for the conservation of populations of several species.     

栖息地破碎化与鸟类生存

栖息地破碎化给野生动物带来的不良后果是全球生态学家和保护生物学家共同关心的问题.自从提出栖息地破碎化是导致生物多样性丧失的关键因素之一的论点后,近20年来,栖息地破碎化研究一直是生态学和保护生物学最活跃的前沿研究领域之一.栖息地破碎化是一动态过程,可在多尺度上发生并蕴涵着复杂的空间模式变化.栖息地破碎化对鸟类的生态学效应主要体现在面积效应、隔离效应和边缘效应等.这些效应影响着鸟类的分布、基因交流、种群动态、扩散行为、种间关系和生活史特征等,最终影响着鸟类的生存.介绍和总结了栖息地破碎化过程、研究的理论依据及栖息地破碎化对鸟类生存产生的诸多影响.     

黑龙江省完达山地区马鹿生境破碎化及其影响因子

应用景观生态学原理和地理信息系统技术,分析黑龙江省完达山地区马鹿生境相关因子重要性、对景观连接度进行模糊相对赋值,建立了景观连接度评价模型及景观斑块指数,研究了黑龙江省完达山地区关于马鹿生境的景观连接度水平、生境的适宜性以及景观的空间结构。结果表明:(1)在155.6km2的面积中,适宜地区的总面积仅为14.81km2,占研究地区的9.52%;次适宜地区的总面积为9.57km2,占研究地区的6.15%;一般适宜地区的总面积为130.05km2,占研究地区的83.58%;不适宜地区的总面积为1.17km2,占研究地区的0.75%;(2)研究地区马鹿各类适宜地区呈多个斑块且相互隔离,在空间分布上处于破碎状态,而且不适宜地区斑块(人为活动景观)的面积比例虽小,在生态系统中形态上的破碎化程度较小,但对马鹿的生境的生态功能的丧失起到重要作用。     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

Landscape modification and habitat fragmentation: a synthesis

Landscape modification and habitat fragmentation are key drivers of global species loss. Their effects may be understood by focusing on: (1) individual species and the processes threatening them, and (2) human-perceived landscape patterns and their correlation with species and assemblages. Individual species may decline as a result of interacting exogenous and endogenous threats, including habitat loss, habitat degradation, habitat isolation, changes in the biology, behaviour, and interactions of species, as well as additional, stochastic threats. Human-perceived landscape patterns that are frequently correlated with species assemblages include the amount and structure of native vegetation, the prevalence of anthropogenic edges, the degree of landscape connectivity, and the structure and heterogeneity of modified areas. Extinction cascades are particularly likely to occur in landscapes with low native vegetation cover, low landscape connectivity, degraded native vegetation and intensive land use in modified areas, especially if keystone species or entire functional groups of species are lost. This review (1) demonstrates that species-oriented and pattern-oriented approaches to understanding the ecology of modified landscapes are highly complementary, (2) clarifies the links between a wide range of interconnected themes, and (3) provides clear and consistent terminology. Tangible research and management priorities are outlined that are likely to benefit the conservation of native species in modified landscapes around the world.     

Long-term effects of forest fragmentation on Amazonian ant communities

Aim To analyse the effects of forest fragmentation on ant communities in an Amazonian landscape that has been fragmented for over a century. Location The region surrounding the village of Alter do Chão in the Brazilian Amazonian state of Pará (2°30′ S, 54°57′ W). Methods Collection of ants and measurements of tree density were performed along transects established in eight sites in continuous forest and in 24 forest fragments surrounded by savanna vegetation. Data on size, perimeter, and degree of isolation (distance to continuous forest and distance to nearest area of forest > 5 ha) of each fragment were obtained from a georeferenced Landsat image of the study area. Results There were significant differences in species richness and composition between fragments and continuous forest, and these differences were not related to intersite variation in vegetation structure (tree density). Fragments supported fewer ant species per plot, and these species tended to represent a nested subset of those found in continuous forests. Fragments had significantly fewer rare species and fewer ant genera. However, fragments and continuous forest had similar numbers of species that also occur in the savanna matrix (i.e. that are not forest specialists). Multiple linear regression analyses indicated that species richness and composition in the fragments are significantly affected by fragment area, but not by fragment shape and degree of isolation. More species were found in larger fragments. Main conclusions Forest fragmentation influences the organization of ant communities in Amazonian savanna/forest landscapes. Forest fragments harboured, on average, 85% of the species found in continuous forest. That these fragments, despite their long history of isolation, support a relatively large complement of the species found in continuous forest is surprising, especially given that in some recently fragmented landscapes the proportion of species surviving in the fragments is lower. Differences in inter‐fragment distance and type of matrix between Alter do Chão and these other landscapes may be involved. The fact that fragments at Alter do Chão are surrounded by a natural (rather than an anthropogenic) habitat, and that most of them are less than 300 m from another forest area, may have helped to ameliorate the adverse effects of forest fragmentation.     

How fire interacts with habitat loss and fragmentation

Biodiversity faces many threats and these can interact to produce outcomes that may not be predicted by considering their effects in isolation. Habitat loss and fragmentation (hereafter ‘fragmentation’) and altered fire regimes are important threats to biodiversity, but their interactions have not been systematically evaluated across the globe. In this comprehensive synthesis, including 162 papers which provided 274 cases, we offer a framework for understanding how fire interacts with fragmentation. Fire and fragmentation interact in three main ways: (i) fire influences fragmentation (59% of 274 cases), where fire either destroys and fragments habitat or creates and connects habitat; (ii) fragmentation influences fire (25% of cases) where, after habitat is reduced in area and fragmented, fire in the landscape is subsequently altered because people suppress or ignite fires, or there is increased edge flammability or increased obstruction to fire spread; and (iii) where the two do not influence each other, but fire interacts with fragmentation to affect responses like species richness, abundance and extinction risk (16% of cases). Where fire and fragmentation do influence each other, feedback loops are possible that can lead to ecosystem conversion (e.g. forest to grassland). This is a well-documented threat in the tropics but with potential also to be important elsewhere. Fire interacts with fragmentation through scale-specific mechanisms: fire creates edges and drives edge effects; fire alters patch quality; and fire alters landscape-scale connectivity. We found only 12 cases in which studies reported the four essential strata for testing a full interaction, which were fragmented and unfragmented landscapes that both span contrasting fire histories, such as recently burnt and long unburnt vegetation. Simulation and empirical studies show that fire and fragmentation can interact synergistically, multiplicatively, antagonistically or additively. These cases highlight a key reason why understanding interactions is so important: when fire and fragmentation act together they can cause local extinctions, even when their separate effects are neutral. Whether fire–fragmentation interactions benefit or disadvantage species is often determined by the species' preferred successional stage. Adding fire to landscapes generally benefits early-successional plant and animal species, whereas it is detrimental to late-successional species. However, when fire interacts with fragmentation, the direction of effect of fire on a species could be reversed from the effect expected by successional preferences. Adding fire to fragmented landscapes can be detrimental for species that would normally co-exist with fire, because species may no longer be able to disperse to their preferred successional stage. Further, animals may be attracted to particular successional stages leading to unexpected responses to fragmentation, such as higher abundance in more isolated unburnt patches. Growing human populations and increasing resource consumption suggest that fragmentation trends will worsen over coming years. Combined with increasing alteration of fire regimes due to climate change and human-caused ignitions, interactions of fire with fragmentation are likely to become more common. Our new framework paves the way for developing a better understanding of how fire interacts with fragmentation, and for conserving biodiversity in the face of these emerging challenges.     

苜蓿草地生境丧失与破碎化对昆虫物种丧失与群落重建的影响

生境破碎化包括生境丧失与破碎化两个相对独立的过程,为探讨这两个过程各自对生物多样性的影响,本文利用苜蓿草地实验模型系统(EMS)构建了36个小区研究不同生境丧失与破碎化对昆虫群落及不同类群的影响,包括18个破碎化小区与18个连续小区,破碎化小区全部采用1 m×1 m(H=1)破碎,连续小区苜蓿连片(H=0),生境丧失采...     

京杭运河邵伯高邮段栖息地破碎化对狗獾种群数量分布的影响

栖息地破碎是生物多样性下降的主要原因之一。栖息地破碎引起的面积效应、隔离效应和边缘效应能影响动物种群的绝灭阈值、分布、多度、种间关系以及生态系统过程, 最终影响动物种群的数量分布。2006年10-11月和2007年10-11月, 利用全球定位系统(GPS)、地理信息系统(GIS)和样方法定量分析京杭运河邵伯至高邮段狗獾栖息地破碎化程度, 确定不同斑块的面积、斑块距离、斑块隔离度以及栖息地质量对斑块中狗獾数量分布的影响。结果表明, 各个斑块内狗獾的洞口数、粪堆数与该斑块的面积显著的正相关(r=0.961, P=0.039; r=0.999, P=0.023), 但与斑块距离、斑块隔离度无显著的相关性(P>0.05)。栖息地的质量也会影响狗獾的数量分布, 多元线性逐步回归分析表明, 人类干扰和与栖息地的郁闭性显著的影响狗獾的数量分布。以上结果说明, 京杭运河邵伯高邮段栖息地的破碎化程度对狗獾的数量分布还没有造成显著的直接影响, 但会间接降低栖息地的质量, 进而影响狗獾的生存。     

Chaco forest fragmentation effects on leaf litter decomposition are not explained by changes in litter fauna

Forest fragmentation is a component of global change, with substantial impact on biodiversity and ecosystem functioning. Despite extensive evidence of forest fragmentation effects on above‐ground ecological processes, little is understood about its below‐ground effects. Abundance and richness of leaf litter fauna can be affected by forest fragmentation, and this can have cascading effects on the decomposition process. Here, we examine how fragmentation of a subtropical dry forest affects aspects of ecosystem structure and functioning, by unravel area and edge effects on leaf litter fauna and decomposition rates and testing whether changes in abundance or richness of litter fauna mediated fragment area and edge effects on litter decomposition. We incubated litterbags filled with a common substrate, at the edge and interior of 12 fragments of Chaco Serrano forest in Central Argentina, for 180 days. We found that invertebrate abundance was higher at the forest edge but independent of fragment area, whereas decomposition declined with fragment size independently of edge or interior location. According to our results, the effect of forest size on decomposition was not mediated by changes in abundance or richness of leaf litter fauna, suggesting independent changes in ecosystem structure and functioning.     

Slow response of plant species richness to habitat loss and fragmentation

We examined the response of vascular plant species richness to long-term habitat loss and fragmentation of Estonian calcareous grasslands (alvars). The current number of habitat specialist species in 35 alvars was not explained by their current areas and connectivities but it was explained by their areas and connectivities 70 years ago ( R ² = 0.27). We estimated the magnitude of extinction debt in local communities by assuming an equilibrium species richness in 14 alvars that had lost only a small amount of area and by applying this model to the remaining alvars, in which the average area has declined from 3.64 km² in the 1930s to 0.21 km² at present. The extinction debt estimated for individual alvars was around 40% of their current species number. Our conclusions are applicable to temperate grasslands in general, which have lost much area because of agricultural intensification and cessation of traditional management.