海洋岛屿生物多样性保育研究进展

海洋岛屿生态系统因具有明显的海域地理隔离而区别于陆地生态系统,被誉为生物地理与进化生态学研究的"天然实验室".陆地或其它邻近岛屿的种源物种迁移到新的岛屿后,经历地理隔离、特征置换或适应辐射等一系列的岛屿进化过程,形成与种源物种具有显著遗传差异的岛屿特有种.岛屿在小面积范围内分化形成大量的特有种,是岛屿生物多样性最为重要的特点之一.但是,岛屿种群由于分布范围局限、生境脆弱且种群规模较小,岛屿种群较陆地种群具有更高的灭绝风险.本文通过对海洋岛屿物种的起源与演化、遗传结构以及岛屿物种的濒危与保护3个热点问题的讨论,阐述岛屿生物多样性的形成机制、濒危肇因以及岛屿生物多样性保育的重要性.     

人类活动效应对物种多样性影响的动力模拟——以洪湖湿地生境毁坏对水鸟物种多样性的影响为例

提出了随时间变化的人类活动效应对物种多样性影响的多物种竞争非自治动力模式,并以洪湖为例模拟了湿地水鸟物种多样性对人类活动效应（生境持续毁坏）的响应过程。模拟发现：对于强．强物种,生境的持续破坏使得湿地水鸟的物种多度大幅度减小,并发生优势种群的更替;对于弱-弱物种,将导致大批的弱物种种群迅速灭绝,而余下的弱物种种群将做准周期振荡;尽管停止对湿地生境的持续毁坏,仍会使一批弱物种种群继续走向灭绝,并且使得原来最强的几个种群最终灭绝。物种灭绝对生境毁坏的这种时间滞后性,即破碎的生境中存在着一些“活死者”,必须引起自然保护学家的关注,否则会低估了实际处于灭绝边缘的物种的数目,从而影响正确的物种保护决策的制订。     

以昆虫作为指示生物评估森林健康的生物学与生态学基础

昆虫是森林生物多样性的重要组成部分,在森林生态系统主体群落中,昆虫与植物密切关联.森林害虫胁迫及昆虫多样性变动,对于评估森林生态系统健康状况具有重要价值.昆虫多样性可用于对森林生态系统健康的快速评估,如基于昆虫指示生物评价森林生态系统的毒害水平、物种丰富度、多样性水平、靶标昆虫的地位和特有种水平等.针对影响森林环境健康的重要干扰因子,本文阐述了应用昆虫作为指示性生物监测与评估森林健康状况的生物与生态学依据,讨论了昆虫种群,特别是珍稀物种种群在生境破碎干扰下的种群波动,以及大气污染、干旱和二氧化碳浓度升高等对昆虫种群密度与分布的影响等.同时,分析了昆虫作为指示性生物监测与评估森林环境健康所存在的问题.     

四川蜂桶寨国家自然保护区地表甲虫物种多样性

在四川雅安蜂桶寨国家自然保护区（102°48′～103°18′ E,30°42′～30°54′ N）及其周边地区,就森林片段化和生态恢复程度的不同,选择4个代表地点,即位于保护区核心地带、以阔叶混交林为主的蜂桶寨天然林区（海拔1680～2080 m）,受到经济开发干扰但植被类型丰富的锅巴岩天然林区（海拔2 280～3 340 m）、以人工针叶林进行生态恢复的蚂蝗沟人工林区（海拔2430～2525 m）,以及森林高度片段化的双石镇农耕区（海拔870～1 165 m）,共设26块样地,以巴氏罐诱法为主研究地表甲虫的群落组成和多样性变化。本研究共采集甲虫标本 2.338 号,隐翅虫数量最多,占39.6%;步甲次之,占29.3%,拟步甲、象甲和叶甲的数量也各在5% 以上,它们共同构成该地区地表甲虫的优势类群。锅巴岩物种的个体数量和丰富度（S）以及多样性指数（H′）较高;蚂蝗沟均匀度指数（J）较高,丰富度较低;蜂桶寨天然林区个体数量较少;双石镇农耕区的多样性和均匀度指数较低。锅巴岩、蜂桶寨和蚂蝗沟间物种分布都有一定程度的相似性,但后两者相似性程度更高,双石镇物种分布与其他3个地点差异较大,这反映了不同地点间的生境异质性和森林植被片段化程度的差异。整个鞘翅目、隐翅虫科和步甲科的个体数量分布在针叶林内较多,在阔叶林内较少;拟步甲科的数量分布在高山灌丛内较多;叶甲科的数量分布在针叶林较多;象甲科除了竹林外,在其他植被内的数量均较多。在总体趋势上,随着海拔的升高,在蜂桶寨和锅巴岩两个邻近的天然林地点,整个鞘翅目以及优势甲虫类群的种类和个体数量也逐渐增多。比较蜂桶寨林区内的个体数量、丰富度和多样性指数,北坡均大于南坡。以上结果表明,物种多样性与海拔、坡向以及生境类型密切相关,森林片段化和生态恢复对物种多样性有显著的影响。因此,在改善森林片段化进行生态恢复时,采取合理措施,增加生境异质性,有助于提高该地区地表甲虫物种多样性。     

景观遗传学原理及其在生境片断化遗传效应研究中的应用

景观遗传学是近年来在景观生态学和种群遗传学之间形成的一个交叉领域,强调景观的组成、空间构型和环境梯度对基因流、种群遗传空间结构和局域种群适应的影响。景观遗传学尚未成为一门独立的学科,其理论基础主要来自分子遗传学、种群生物学和景观生态学。现代分子遗传标记技术、遥感和GIS支持下的景观分析和空间统计方法的综合运用是景观遗传学主要研究途径。系统介绍了景观遗传学的基础概念,关键科学问题,基本理论框架,及其与相邻研究领域的关系,综述了景观遗传学最为关注的现实课题——景观碎裂化的种群遗传效应的研究进展,主要涉及生境片断化的遗传效应、不同属性的物种响应、以及生境片断化过程的选择作用等方面。通过采取一种跨尺度的视角,景观遗传学研究显著深化了关于景观碎裂化对生物多样性影响的理解。     

荒漠破碎化生境中长爪沙鼠集合种群野外验证研究

近年来,人类活动和自然干扰,导致内蒙古阿拉善荒漠区生境的破碎化,出现了长爪沙鼠在不同斑块间的不连续分布,每一斑块内可能存在一个局域种群,而集合种群建立的前提条件,是局域种群斑块状分布在离散的栖息地环境中。2002~2012年每年的4~10月,在阿拉善荒漠区禁牧、轮牧、过牧和开垦4种人为不同利用方式形成的生境斑块中,采用标志重捕法对长爪沙鼠（Meriones unguiculatus）种群进行定点监测。通过分析长爪沙鼠种群动态,计算各局域种群的灭绝风险,利用Spearman秩相关系数检验种群动态的空间同步性,同时以种群周转率对长爪沙鼠扩散能力进行评估,以检验阿拉善荒漠区长爪沙鼠种群空间结构是否具有经典集合种群的功能。结果表明：（1） 不同生境斑块可被长爪沙鼠局域种群占据,11年间捕获长爪沙鼠2~7次不等;（2） 长爪沙鼠所有局域种群均具有灭绝风险,在轮牧区和禁牧区灭绝率高达1.000 0,开垦区灭绝率最低,也达到0.333 4,而本研究期间最大局域种群（2008年过牧区,26只/hm²）,在2010年发生了局域灭绝;（3） 不同生境斑块间没有明显的空间隔离而阻碍局域种群的重新建立,长爪沙鼠扩散能力较强,绝大部分月份的种群周转率在50.0%以上,特别是周转率达到100.0%的月份较多;（4） 不同生境斑块间仅轮牧区和禁牧区中长爪沙鼠种群密度显著正相关（P<0.05）,而其他生境斑块间相关性均不显著（P >0.05）,长爪沙鼠局域种群整体显示出明显的非同步空间动态。阿拉善荒漠区长爪沙鼠种群满足作为经典集合种群物种区域续存的4个条件,具有作为研究小哺乳动物集合种群的潜在价值。     

栖息地片断化对动物种群间基因流的影响及其测定方法

栖息地片断化指在人为活动和自然干扰下,大面积连续分布的自然栖息地被其它非适宜栖息地分隔成许多面积较小的斑块(岛屿)的过程。栖息地片断化是导致生物多样性丧失和物种绝灭的主要因素,也是威胁自然界生物生存的重要因素之一。栖息地片断化将影响基因在种群间和种群内的运动(基因流),开展栖息地片断化研究对保持物种间遗传多样性具有重要意义。本文介绍了片断化程度和基因流的测定方法。回顾了国内外的研究现状,并探讨了片断化对种群间基因流的影响和对濒危物种保护的意义。     

集合种群的似Allee效应

从局域种群出发,建立了一个既包括局域种群动态,又包含集合种群侵占率的整合模型,并在这两个层次上进行了计算机模拟,结果表明：（1）同局域种群的Allee效应相类似,集合种群的斑块（适宜生境）侵占比例也存在一个临界值,即使有足够的适宜生境,当斑块的侵占比例低于这个临界值时,集合种群优将趋于灭绝。（2）这个临界值与局域种各的Allee效应密切相关,这将给自然保护,尤其稀有生物的保护以很大的启示。     

基于目标种保护的生态廊道构建——以崇明岛为例

景观破碎化和适宜生境面积减少是生物多样性降低与物种灭绝的重要因素之一。生态廊道的构建为陆行野生动物扩大活动范围提供了空间途径,从而增加了物种基因交流的机会, 提高了种群的生存能力。上海市崇明岛是一个地理上孤立的冲积沙岛,随着岛上森林斑块的减少与破碎化,依赖林地生境生存的陆行动物个体与种群数量大大减少。本文以崇明岛为例,选择上海市一级保护动物刺猬作为目标种,在分析其生活习性和基质斑块空间分布特征的基础上,利用最小耗费模型和GIS手段,实现了生境斑块之间的最佳连接,建立了以目标动物保护为目的的生态廊道规划方案。     

坝上地区农田和恢复生境地表甲虫多样性

研究采用陷阱法对河北坝上崇礼县农田、禁牧草地及再造林地3种常见生境中地表甲虫进行了取样调查,分别从鞘翅目分科、步甲分种两个分类水平比较分析了不同生境中的生物多样性状况以评估生境恢复状况及其对生物多样性的影响,并探索在以生物多样性为指标评估生境恢复状况过程中,研究结论是否受分类水平的影响。结果显示,草地和林地具有较为相似鞘翅目和步甲的结构组成,但均显著不同于农田。农田较再造林地和禁牧草地具有更多的鞘翅目科数及步甲物种数,且均显著高于林地;农田中步甲群落的Fisher's α多样性指数也显著地高于林地,但鞘翅目分科水平上的Fisher's α多样性指数在各生境中没有差异。结果说明:(1)再造林和草地禁牧的生境修复活动在当前状态下对生物多样性恢复的作用尚不明显,仅是表现出促进了景观尺度上较高的Beta和Gamma多样性,而农田则在维持当地较高的生物多样性中扮演重要角色;(2)在鞘翅目分科和步甲科分种研究数据所反映的不同生境条件下的多样性和组成状况基本相同,因此在需要快速评估生境恢复状况且物种水平鉴定存在困难的时候,可以采用鞘翅目分科的数据替代指示生境恢复的状况并反映生境恢复对生物多样性影响的趋势,从而减少分类的难度并提高工作效率。但是物种水平的多样性状况更能显著反映生境间的差异。     

Biodiversity of man-made open habitats in an underused country: a class of multispecies abundance models for count data

Since the 1960s, Japan has become highly dependent on foreign countries for natural resources, and the amount of managed lands (e.g. coppice, grassland, and agricultural field) has declined. Due to infrequent natural and human disturbance, early-successional species are now declining in Japan. Here we surveyed bees, birds, and plants in four human-disturbed open habitats (pasture, meadow, young planted forest, and abandoned clear-cut) and two forest habitats (mature planted forest and natural old-growth). We extended a recently developed multispecies abundance model to accommodate count data, and used the resulting models to estimate species-, functional group-, and community-level state variables (abundance and species richness) at each site, and compared them among the six habitats. Estimated individual-level detection probability was quite low for bee species (mean across species = 0.003; 0.16 for birds). Thirty-two (95% credible interval: 13–64) and one (0–4) bee and bird species, respectively, were suggested to be undetected by the field survey. Although habitats in which community-level abundance and species richness was highest differed among taxa, species richness and abundance of early-successional species were similar in the four disturbed open habitats across taxa except for plants in the pasture habitat which was a good habitat only for several exotic species. Our results suggest that human disturbance, especially the revival of plantation forestry, may contribute to the restoration of early-successional species in Japan.     

Consequences of forest fragmentation for polyporous fungi at two spatial scales

Greatly reduced area of old-growth forests and the very low amount of dead wood in managed forests in northern Europe have caused a marked decline in the populations of saproxylic species. It is less clear at which spatial and temporal scales these adverse changes are taking place, and more information is needed to reliably predict which species are especially sensitive to loss and fragmentation of habitat. Here we compare species richness, incidence of occurrence in forest fragments, and abundance of polyporous fungal species and species groups between two regions in Finland with contrasting histories of forestry and a marked difference in the amount and spatial configuration of old-growth forests. We also analyse the consequences of increasing loss of connectivity on the presence and abundance of polypores in a study region with a documented short-term history of old-growth fragmentation. Our results show that the species number, incidence of occurrence, and abundance of especially the rare, threatened, and near-threatened species are much lower in the old-growth fragments in Häme in southern Finland in comparison with Kuhmo in eastern Finland, most probably because of the longer history of intensive forestry in Häme. Among the rare species, the species that show the greatest difference between the two regions (at the scale of 500 km) also tended to respond most strongly to the more recent forest fragmentation within the study region in Kuhmo (at the scale of 50 km). Polypores associated with spruce seem to be more strongly affected by forestry than species associated with pine, possibly reflecting the differences in the natural dynamics of spruce-dominated and pine-dominated forests.     

The effects of forestry on carabid beetles (Coleoptera: Carabidae) in boreal forests

As compared to natural forests, managed boreal forests are younger, more homogeneous in terms of tree age and species composition, and consist of smaller fragments. Here we examine the effects of such characteristics caused by forestry on carabid beetles (Coleoptera, Carabidae) in the boreal region. The main results are the following. (1) Fragmentation of forests and the size of a fragment appear not to be crucial for the survival of the majority of forest carabids, as they tend to be distributed over various successional stages, but species requiring old-growth habitats suffer. (2) For carabids there appear to be no or very few edge specialist species, and forest-open land edges appear to be effective barriers for species associated with forest or open habitat. However, generalist species easily cross the edge, and edges of forest fragments may be invaded by species from the surrounding open habitat. (3) Habitat change following clear-cutting dramatically changes the composition of carabid assemblages: species restricted to mature forests disappear and open-habitat species invade, while habitat generalists survive at least in the short term. Carabid diversity can probably best be maintained if forest management mimics natural processes, maintains natural structures and includes the natural composition of vegetation and other structural elements (such as dead wood) within the stands, provided that these forest features can be maintained and recreated through forest management practices. At a larger scale, the whole spectrum of forest types and ages (especially old-growth forests), and different successional processes (especially fire) should be maintained. These require the development and use of innovative logging methods, and the planning, implementation, and assessment of landscape-scale ecological management strategies.     

The effect of deforestation on the genetic diversity and structure in <Emphasis Type="Italic">Acer saccharum</Emphasis> (Marsh): Evidence for the loss and restructuring of genetic variation in a natural system

The level and distribution of genetic diversity can be influenced by species life history traits and demographic factors, including perturbations that might produce population bottlenecks. Deforestation and forest fragmentation are common sources of population disturbance in contemporary populations of forest ecosystems. Although the genetic effects of forest fragmentation and deforestation have been examined by assessing levels of genetic variation in forest fragments that remain after logging, few considerations have been made of the populations that re-colonize once-cleared areas. Here we examine the effects of human-mediated population bottlenecks on the level and distribution of genetic diversity in natural populations of the long-lived forest tree species, Acer saccharum (sugar maple). We compared genetic variation and structure for populations of sugar maple found within old-growth forested area and in area that has re-colonized since logging. In this study the percent polymorphic loci and allelic richness estimates were reduced in the logged populations compared to old-growth populations. Jackknifed estimates of population genetic differentiation showed significantly higher differentiation among logged populations, with this result being consistently seen when individuals within populations were grouped according to diameter at breast height. The result of decreased genetic variation and higher levels of genetic structure among logged populations suggests that even one extensive bout of logging can alter the level and distribution of genetic variation in this forest tree species.     

Forest fragmentation and selective logging have inconsistent effects on multiple animal-mediated ecosystem processes in a tropical forest

Forest fragmentation and selective logging are two main drivers of global environmental change and modify biodiversity and environmental conditions in many tropical forests. The consequences of these changes for the functioning of tropical forest ecosystems have rarely been explored in a comprehensive approach. In a Kenyan rainforest, we studied six animal-mediated ecosystem processes and recorded species richness and community composition of all animal taxa involved in these processes. We used linear models and a formal meta-analysis to test whether forest fragmentation and selective logging affected ecosystem processes and biodiversity and used structural equation models to disentangle direct from biodiversity-related indirect effects of human disturbance on multiple ecosystem processes. Fragmentation increased decomposition and reduced antbird predation, while selective logging consistently increased pollination, seed dispersal and army-ant raiding. Fragmentation modified species richness or community composition of five taxa, whereas selective logging did not affect any component of biodiversity. Changes in the abundance of functionally important species were related to lower predation by antbirds and higher decomposition rates in small forest fragments. The positive effects of selective logging on bee pollination, bird seed dispersal and army-ant raiding were direct, i.e. not related to changes in biodiversity, and were probably due to behavioural changes of these highly mobile animal taxa. We conclude that animal-mediated ecosystem processes respond in distinct ways to different types of human disturbance in Kakamega Forest. Our findings suggest that forest fragmentation affects ecosystem processes indirectly by changes in biodiversity, whereas selective logging influences processes directly by modifying local environmental conditions and resource distributions. The positive to neutral effects of selective logging on ecosystem processes show that the functionality of tropical forests can be maintained in moderately disturbed forest fragments. Conservation concepts for tropical forests should thus include not only remaining pristine forests but also functionally viable forest remnants.     

The conservation value of South East Asia's highly degraded forests: evidence from leaf-litter ants

South East Asia is widely regarded as a centre of threatened biodiversity owing to extensive logging and forest conversion to agriculture. In particular, forests degraded by repeated rounds of intensive logging are viewed as having little conservation value and are afforded meagre protection from conversion to oil palm. Here, we determine the biological value of such heavily degraded forests by comparing leaf-litter ant communities in unlogged (natural) and twice-logged forests in Sabah, Borneo. We accounted for impacts of logging on habitat heterogeneity by comparing species richness and composition at four nested spatial scales, and examining how species richness was partitioned across the landscape in each habitat. We found that twice-logged forest had fewer species occurrences, lower species richness at small spatial scales and altered species composition compared with natural forests. However, over 80 per cent of species found in unlogged forest were detected within twice-logged forest. Moreover, greater species turnover among sites in twice-logged forest resulted in identical species richness between habitats at the largest spatial scale. While two intensive logging cycles have negative impacts on ant communities, these degraded forests clearly provide important habitat for numerous species and preventing their conversion to oil palm and other crops should be a conservation priority.     

Impact of logging on the richness and diversity of forest butterflies in a tropical dry forest in Thailand

Studies on the impact of logging on tropical forest butterflies have been almost exclusively conducted in moist forest habitats. This study considers the impacts of small-scale logging on butterfly communities at three sites of varying disturbance intensity in a tropical dry forest in western Thailand. Butterfly species richness was similar at all sites, but the abundance of butterflies and diversity of the butterfly community decreased with increased logging disturbance. The recorded decrease in diversity at the relatively large sampling scale used lends further support to the hypothesis that disturbance effects are scale dependent. Species abundance data for butterflies fitted a log-normal distribution at all sites, but also a log-series distribution at the two disturbed sites. These analyses suggest a more complex butterfly community at the undisturbed site, but also that log-series and log-normal distributions may not to be sufficiently sensitive to be useful indicators of community changes following logging. Community ordination separates both the butterfly species and transect samples into three distinct regions corresponding to the three study locations. Ordination axes are correlated with tree density, understorey cover and understorey plant richness. Species with the smallest geographic ranges tend to be the least abundant and occurred most frequently in the undisturbed site. The observed diverging responses to disturbance among butterfly families diminishes the value of butterfly communities as biodiversity indicators, and forest managers should perhaps focus on restricted range species or of groups of recognized sensitive species for this purpose.     

An examination of scale of assessment,logging and ENSO-induced fires on butterfly diversity in Borneo

The impact of disturbance on species diversity may be related to the spatial scales over which it occurs. Here I assess the impact of logging and ENSO (El Niño Southern Oscillation) -induced burning and forest isolation on the species richness (477 species out of more than 28,000 individuals) and community composition of butterflies and butterfly guilds using small (0.9 ha) plots nested within large (450 ha) landscapes. The landscapes were located in three habitat classes: (1) continuous, unburned forest; (2) unburned isolates surrounded by burned forest; and (3) burned forest. Plots with different logging histories were sampled within the two unburned habitat classes, allowing for independent assessment of the two disturbance factors (logging and burning). Disturbance within habitat classes (logging) had a very different impact on butterfly diversity than disturbance among habitat classes (due to ENSO-induced burning and isolation). Logging increased species richness, increased evenness, and lowered dominance. Among guilds based on larval food plants, the species richness of tree and herb specialists was higher in logged areas but their abundance was lower. Both generalist species richness and abundance was higher in logged areas. Among habitat classes, species richness was lower in burned forest and isolates than continuous forest but there was no overall difference in evenness or dominance. Among guilds, generalist species richness was significantly lower in burned forest and isolates than continuous forest. Generalist abundance was also very low in the isolates. There was no difference among disturbance classes in herb specialist species richness but abundance was significantly higher in the isolates and burned forest than in continuous forest. Tree specialist species richness was lower in burned forest than continuous forest but did not differ between continuous forest and isolates.The scale of assessment proved important in estimating the impact of disturbance on species richness. Within disturbance classes, the difference in species richness between primary and logged forest was more pronounced at the smaller spatial scale. Among disturbance classes, the difference in species richness between continuous forest and isolates or burned forest was more pronounced at the larger spatial scale. The lower levels of species richness in ENSO-affected areas and at the larger (landscape) spatial scale indicate that future severe ENSO events may prove one of the most serious threats to extant biodiversity.     

Differences in the Effects of Selective Logging on Amphibian Assemblages in Three West African Forest Types

Making generalizations about the impact of commercial selective logging on biodiversity has so far remained elusive. Species responses to logging depend on a number of factors, many of which have not been studied in detail. These factors may include the natural forest conditions (forest types) under which logging impacts are investigated; but this question has so far remained unexamined. In a large‐scale replicate study we aimed at clarifying the relationship between logging and forest types on leaf litter frogs. We contrast three distinct and naturally occurring forest types, including wet evergreen, moist evergreen and semi‐deciduous forests. Selectively logged sites were compared with primary forest sites for each forest type. We found that the response of frog communities to logging varies in different forest types. In the wet evergreen forest, richness was higher in logged forest than primary forest, while diversity measures were not different between logged and primary forest habitats. In the moist evergreen, richness and diversity were higher in selectively logged areas compared with primary forest habitats. In the semi‐deciduous, logged forests were characterized by drastic loss of forest specialists, reduced richness, and diversity. These results indicate that the net effect of logging varies with respect to forest type. Forest types that are characterized by adverse climatic conditions (i.e., low rainfall and protracted dry seasons) are more likely to produce negative effects on leaf litter anuran communities. For comparisons of the impact of logging on species to be effective, future research must endeavor to include details of forest type.     

Road networks predict human influence on Amazonian bird communities

Road building can lead to significant deleterious impacts on biodiversity, varying from direct road-kill mortality and direct habitat loss associated with road construction, to more subtle indirect impacts from edge effects and fragmentation. However, little work has been done to evaluate the specific effects of road networks and biodiversity loss beyond the more generalized effects of habitat loss. Here, we compared forest bird species richness and composition in the municipalities of Santarém and Belterra in Pará state, eastern Brazilian Amazon, with a road network metric called ‘roadless volume (RV)’ at the scale of small hydrological catchments (averaging 3721 ha). We found a significant positive relationship between RV and both forest bird richness and the average number of unique species (species represented by a single record) recorded at each site. Forest bird community composition was also significantly affected by RV. Moreover, there was no significant correlation between RV and forest cover, suggesting that road networks may impact biodiversity independently of changes in forest cover. However, variance partitioning analysis indicated that RV has partially independent and therefore additive effects, suggesting that RV and forest cover are best used in a complementary manner to investigate changes in biodiversity. Road impacts on avian species richness and composition independent of habitat loss may result from road-dependent habitat disturbance and fragmentation effects that are not captured by total percentage habitat cover, such as selective logging, fire, hunting, traffic disturbance, edge effects and road-induced fragmentation.