Tree species richness complementarity, disturbance and fragmentation in a Mexican tropical montane cloud forest

Tropical montane cloud forest exhibits great heterogeneity in speciescomposition and structure over short geographic distances. In central Veracruz,Mexico, the conservation priority of seven cloud forest fragments was assessedby considering differences in woody plant species richness and complementarityof species among sites, forest structure, tree mortality, and timber andfirewood extraction as indicators of anthropogenic disturbance. Densities oftrees 5 cm dbh (360–1700 trees/ha) weredifferent among the sites, but basal area (35.3–89.3m²/ha) did not differ among fragments. The number of dead trees rangedfrom 10–30 to 170–200 trees/ha. The fragments'species composition was different but complementary. The Morisita–Hornindex indicated low similarity between fragments. A non-parametric estimator ofspecies richness indicated that more sampling effort would be necessary tocomplete the inventory (15 additional trees and two understory shrub species).Unfortunately, most of the fragments are threatened with deforestation. The numberof cut trees was similar among sites (0–15 cut trees/0.1 ha).Sites with immediate need for conservation were close to settlements, with highnumbers of cut trees and no legal protection. The selected sites represent thevariety of situations that exist in the region. Given the high complementarityobserved between fragments, a regional conservation approach will be required topreserve the last repositories of part of the tremendous biodiversity of theonce continuous forest in this region.     

Effect of forest fragmentation on bird species richness in Papua New Guinea

Tropical forests worldwide are being fragmented at a rapid rate, causing a tremendous loss of biodiversity. Determining the impacts of forest disturbance and fragmentation on tropical biotas is therefore a central goal of conservation biology. We focused on bird communities in the interior (>100 m from forest edge) of forest fragments (300, 600, and 1200 ha) in the lowlands of Papua New Guinea and compared them with those in continuous forest. We surveyed bird communities using point counts, mist‐netting, and random walks, and measured habitat and microclimate characteristics at each site. We also surveyed leaf‐dwelling arthropods, butterflies, and ants, and obtained diet samples from birds to examine food availability and food preferences. We recorded significantly fewer bird species per point in the 300‐ha forest fragment than in other study sites. Overall, we recorded 80, 84, and 88 species, respectively, in forest fragments, and 102 in continuous forest. Frugivores (especially large frugivores) and insectivores had lower species richness in forest fragments than continuous forest. Our results did not support the food scarcity hypothesis, that is, the decline of insectivorous birds in forest fragments is caused by an impoverished invertebrate prey base. We also found no significant differences among forest fragments and continuous forest in microclimates of forest interiors. Rather, we found that microhabitats preferred by sensitive birds (i.e., 30% of species with the strongest preferences for continuous forest) were less common in forest fragments (19%–31% of points) than in continuous forest (86% of points). Our results suggest that changes in microhabitats may make forest fragments unsuitable for sensitive species. However, limited dispersal capabilities could also make some species of birds less likely to disperse and occupy fragments. In addition, impoverished food resources, size of the forest fragment, or hunting pressure could contribute to the absence of large frugivorous birds in forest fragments. The forest fragments in our study, preserved as village‐based protected areas, were not large enough to sustain the bird communities found in continuous forest. However, because these fragments still contained numerous bird species, preservation of such areas can be an important component of management strategies to conserve rainforests and birds in Papua New Guinea.     

Possible effects of habitat fragmentation and climate change on the range of forest plant species

Global circulation models predict an increase in mean annual temperature between 2.1 and 4.6 °C by 2080 in the northern temperate zone. The associated changes in the ratio of extinctions and colonizations at the boundaries of species ranges are expected to result in northward range shifts for a lot of species. However, net species colonization at northern boundary ranges, necessary for a northward shift and for range conservation, may be hampered because of habitat fragmentation. We report the results of two forest plant colonization studies in two fragmented landscapes in central Belgium. Almost all forest plant species (85%) had an extremely low success of colonizing spatially segregated new suitable forest habitats after c . 40 years. In a landscape with higher forest connectivity, colonization success was higher but still insufficient to ensure large-scale colonization. Under the hypothesis of net extinction at southern range boundaries, forest plant species dispersal limitation will prevent net colonization at northern range boundaries required for range conservation.     

The Habitat Amount Hypothesis implies negative effects of habitat fragmentation on species richness

The habitat amount hypothesis (HAH) predicts that species richness in a habitat site increases with the amount of habitat in the ‘local landscape’ defined by an appropriate distance around the site, with no distinct effects of the size of the habitat patch in which the site is located. It has been stated that a consequence of the HAH, if supported, would be that it is unnecessary to consider habitat configuration to predict or manage biodiversity patterns, and that conservation strategies should focus on habitat amount regardless of fragmentation. Here, I assume that the HAH holds and apply the HAH predictions to all habitat sites over entire landscapes that have the same amount of habitat but differ in habitat configuration. By doing so, I show that the HAH actually implies clearly negative effects of habitat fragmentation, and of other spatial configuration changes, on species richness in all or many of the habitat sites in the landscape, and that these habitat configuration effects are distinct from those of habitat amount in the landscape. I further show that, contrary to current interpretations, the HAH is compatible with a steeper slope of the species–area relationship for fragmented than for continuous habitat, and with higher species richness for a single large patch than for several small patches with the same total area (SLOSS). This suggests the need to revise the ways in which the HAH has been interpreted and can be actually tested. The misinterpretation of the HAH has arisen from confounding and overlooking the differences in the spatial scales involved: the individual habitat site at which the HAH gives predictions, the local landscape around an individual site and the landscapes or regions (with multiple habitat sites and different local landscapes) that need to be analysed and managed. The HAH has been erroneously viewed as negating or diminishing the relevance of fragmentation effects, while it actually supports the importance of habitat configuration for biodiversity. I conclude that, even in the cases where the HAH holds, habitat fragmentation and configuration are important for understanding and managing species distributions in the landscape.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

Effects of forest fragmentation on European birds: implications of regional differences in species richness

Aim In this paper, we adopted a large‐scale approach to evaluate the effect of regional richness of forest birds on the number of bird species retained by forest fragments in several localities across Europe. Location We studied bird assemblages in fourteen forest archipelagos embedded in agricultural matrices from southern Norway to central Spain. Tree composition varied from oak and beech forests of the northern localities to oak and pine xerophitic woodlands of the southern ones. The number of fragments in each forest archipelago ranged from eighteen to 211. Methods We used the Gleason equation (s = a + z log A; where s and A are, respectively, the species richness and size of forest fragments and z the rate of species loss) to estimate the species richness for 1‐ and 15‐ha fragments in each archipelago. The regional richness of forest birds was estimated by modelling the geographical distribution of species richness in the European atlas of breeding birds. Results The latitudinal distribution of regional richness displayed a convex form, with the highest values being in central Europe. Along this gradient, the number of species retained by fragments and the rate of species loss was positively related to regional richness. In addition, the percentage of the regional pool of species sampled by fragments decreased in the southern localities. Main conclusions Relationships between regional richness of forest birds and richness in fragments seem to explain why fragments in central Europe shelter more species than their southern counterparts. The decreased ability of southern forest fragments to sample the regional richness of forest birds, could be explained as an effect of the low abundance of many species in the Mediterranean, which could depress their ability to prevent extinction in fragments by a rescue effect. Alternatively, high beta diversity in the Mediterranean could produce undersampling by fragments of the regional pool of species. These regional differences in the response of bird assemblages to forest fragmentation are used to discuss the usefulness of large‐scale, biogeographical approaches in the design of conservation guidelines.     

Heavy metal pollution negatively correlates with anuran species richness and distribution in south‐eastern Australia

Heavy metal pollution has likely played an important role in global biodiversity decline, but there remains a paucity of information concerning the effects of metals on amphibian diversity. This study assessed anuran species richness and distribution in relation to sediment metal content and water chemistry in wetlands located along the Merri Creek corridor in Victoria, south‐eastern Australia. Anurans were present in 60% (21/35) of study sites, with a total of six species detected: the eastern common froglet (Crinia signifera), the eastern sign‐bearing froglet (Crinia parinsignifera), the southern brown tree frog (Litoria ewingii), the growling grass frog (Litoria raniformis), the eastern banjo frog (Limnodynastes dumerilii) and the spotted marsh frog (Limnodynastes tasmaniensis). Mean species richness was 1.77 ± 0.32 per site, and species richness ranged from zero to six species per site. Across sites, species richness correlated negatively with sediment concentrations of six heavy metals: copper, nickel, lead, zinc, cadmium and mercury. Species richness also correlated negatively with wetland water electrical conductivity (a proxy for salinity) and concentrations of orthophosphate. Distributions of the three most commonly observed frog species (C. signifera, L. tasmaniensis and L. ewingii) were significantly negatively associated with the total level of metal contamination at individual sites. The study is the first to provide evidence for an association between metal contamination and anuran species richness and distribution in the southern hemisphere, adding to a small but growing body of evidence that heavy metal pollution has contributed to global amphibian decline.     

Local versus landscape-scale effects of anthropogenic land-use on forest species richness

The study investigated the effects of human-induced landscape patterns on species richness in forests. For 80 plots of fixed size, we measured human disturbance (categorized as urban/industrial and agricultural land areas), at ‘local’ and ‘landscape’ scale (500 m and 2500 m radius from each plot, respectively), the distance from the forest edge, and the size and shape of the woody patch. By using GLM, we analyzed the effects of disturbance and patch-based measures on both total species richness and the richness of a group of specialist species (i.e. the ‘ancient forest species’), representing more specific forest features. Patterns of local species richness were sensitive to the structure and composition of the surrounding landscape. Among the landscape components taken into account, urban/industrial land areas turned out as the most threatening factor for both total species richness and the richness of the ancient forest species. However, the best models evidenced a different intensity of the response to the same disturbance category as well as a different pool of significant variables for the two groups of species. The use of groups of species, such as the ancient forest species pool, that are functionally related and have similar ecological requirements, may represent an effective solution for monitoring forest dynamics under the effects of external factors. The approach of relating local assessment of species richness, and in particular of the ancient forest species pool, to land-use patterns may play an important role for the science-policy interface by supporting and strengthening conservation and regional planning decision making.     

海南五指山热带山地雨林植物物种多样性研究

王指山热带山地雨林的物种种类十分丰富,在１ｈｍ＾２样地中共出现５４个科,１００个属的乔木树种１７７种,１３３７个个体。区系地理分析表明属的分布区类型以热带分布型成分占优,为８９．０％,充分体现了五指山山地雨林的热带性质。多样性指数分析结果为：样地１Ｍａｒｇａｌｅｆ指数１７．８２２,Ｓｈａｎｎｏｎ－Ｗｉｅｎｅｒ指数５．６２１,均匀度０．８２３,Ｓｉｍｐｓｏｎ指数０．０５０,均匀度６．７７５,表明五指     

宏生态尺度上景观破碎化对物种丰富度的影响

生物多样性的地理格局及其形成机制是宏生态学与生物地理学的研究热点。大量研究表明,景观尺度上的生境破碎化对物种多样性的分布格局具有重要作用,但目前尚不清楚这种作用是否足以在宏生态尺度上对生物多样性地理格局产生显著影响。利用中国大陆鸟类和哺乳动物的物种分布数据,在100 km×100 km网格的基础上生成了这两个类群生物的物种丰富度地理格局,进一步利用普通最小二乘法模型和空间自回归模型研究了物种丰富度与气候、生境异质性、景观破碎化的相关关系。结果表明,景观破碎化因子与鸟类和哺乳动物的物种丰富度都具有显著的关联关系,其方差贡献率可达约30%—50%(非空间模型)和60%—80%(空间模型),略低于或接近于气候和生境异质性因子。方差分解结果显示,景观破碎化因子与气候和生境异质性因子的方差贡献率的重叠部分达20%—40%。相对鸟类而言,景观破碎化对哺乳动物物种丰富度的地理格局具有更高的解释率。     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Topographic controls on the distribution of tree islands in the high Andes of south‐western Ecuador

Aim To evaluate the hypothesis that geomorphometric parameters of upper montane Andean environments have an important influence on the regional fire ecology and consequently play a role in the spatial distribution of ‘remnant’ tree islands dominated by Polylepis. Location A glacial landscape located between 3600 and 4400 m elevation in Cajas National Park, south‐western Ecuador. Methods The eigenvalue ratio method was used to evaluate the regional geomorphometric parameters of a 30‐m digital elevation model for Cajas National Park. The landscape character was evaluated by quantifying the topographic roughness, organization, and gradient. This information was used to determine the spatial correlations between terrain characteristics and the distribution of tree islands in the region. Results We demonstrate a strong spatial correlation between areas of high topographic roughness and gradient, and the locations of the major tree islands. We find that there is a distinctive relationship between the topographic roughness and organization in the vicinity of the tree islands (e.g. increased upslope roughness and decreased topographic grain strength) that substantiates the notion that the tree islands are located in relatively inaccessible topography. Main conclusions In the northern and central Andes, the location of Polylepis‐dominated ‘forest islands’ has been shown to be a function of climate, terrain characteristics, and anthropogenic disturbances. Although the relative importance of various ecological factors has been debated, it remains clear that fires have exerted a strong influence on these ecosystems. Other authors have noted that tree islands are more likely to occur at the base of cliffs, above moist areas, and in other areas where fires do not burn frequently. Our results corroborate these observations, and demonstrate that the occurrence of Polylepis patches is strongly correlated with specific combinations of terrain features. Although we do not discount the importance of other factors in determining the spatial position and areal extent of these forests, we demonstrate strong support for fire‐related hypotheses.     

Long‐term effects of exclusion of grazing stock on degraded herbaceous plant communities in a riparian Eucalyptus camaldulensis forest in south‐eastern Australia

Abstract Stock grazing has degraded many riparian ecosystems around the world. However, the potential for ecosystem recovery following the removal of grazing stock is poorly known. We developed a conceptual model to predict the responses of native and exotic herbaceous plants to grazing exclusion, based on site productivity and the degree of initial vegetation degradation. The effects of excluding grazing stock on richness, cover and composition of herbaceous plants were examined over 12 years in the degraded understorey of a riparian forest in Gulpa Island State Forest in south‐eastern Australia. We predicted that grazing exclusion would lead to limited changes in vegetation cover, richness and composition, owing to presumed low site productivity and the high degree of understorey degradation. Results showed that the cover, richness and composition of native and exotic species varied significantly among years. Over all plots, regions and years, total cover was slightly but significantly lower in grazed than in ungrazed plots (43.4% vs. 50.8%). While the cover of native plants increased over time in both treatments, the rate of increase was slightly greater in ungrazed plots. Grazing exclusion had no effect on the richness of native or exotic species, but had a significant but minor impact on plant composition, with different common species (mostly exotics) being promoted or diminished in ungrazed plots. The composition of grazed and ungrazed areas did not become more different over time. Overall, the effects that could be attributed to grazing exclusion were relatively minor and transient. Results are consistent with predictions based on site productivity and initial degradation, and should not be extrapolated to other more productive, or less degraded, riparian systems.     

长白山森林景观破碎的遥感探测

天然林数量的减少以及质量的降低对中国的生态环境和经济发展产生不利影响。虽然林地的破碎对各种自然灾害起到关键性作用,林地是如何被破碎的尚不十分清楚。应用卫星遥感技术,以及地理信息系统和空间分析手段研究中国东北－原始的针阔混交林的破碎过程。结果表明,本来完整的原始森林,已经被一种“复叶”状小面积皆伐作业分割为零碎的斑真。采伐地的面积平均为15hm^2, 但随着采伐的继续、采伐地段连为一体,增大了采伐迹地的面积,从而保留的林地面积越来越小。采伐地点的选择没有考虑物种和环境的保护,林地破碎的特征是保留林地面积的缩小,中、小林地数量的增加,林地斑块形状的改变,以及林缘长度的增加。最后就保护东北森林植被完整性的角度提出林业政策与森林经营的改进意见。     

Gap-phase regeneration in a tropical montane forest: the effects of gap structure and bamboo species

To study the influence of gap structure and bamboo species on the regrowth of montane Atlantic forest, colonization by plants was characterized in 30 treefall gaps (30.3–500.5 m²). The study was conducted at Santa Virgínia (45°30 W, 23°17 S), a 4970-ha reserve of Atlantic montane forest in southeastern Brazil. Area covered by bamboos ranged from 0% to 100% of gap area. Average height of surrounding canopies ranged from 12 to 30 m. As gap are covered by bamboo and average height of surrounding canopies increased, both density and richness of pioneer woody species decreased. Density and richness of shade-tolerant species were negatively influenced by gap area. Low-light-demanding species of Miconia, Leandra and Rapanea accounted for the majority of both pioneer species and individuals sampled, whereas high-light demanding pioneers of Cecropia, Alchornea and Tibouchina were poorly represented. We suggest that in the Atlantic montane forest bamboo species compete for gaps, excluding other light-demanding pioneers. This results in an overall reduction of pioneer species richness in the Atlantic forest.     

Multiple successional pathways in human‐modified tropical landscapes: new insights from forest succession,forest fragmentation and landscape ecology research

Old‐growth tropical forests are being extensively deforested and fragmented worldwide. Yet forest recovery through succession has led to an expansion of secondary forests in human‐modified tropical landscapes (HMTLs). Secondary forests thus emerge as a potential repository for tropical biodiversity, and also as a source of essential ecosystem functions and services in HMTLs. Such critical roles are controversial, however, as they depend on successional, landscape and socio‐economic dynamics, which can vary widely within and across landscapes and regions. Understanding the main drivers of successional pathways of disturbed tropical forests is critically needed for improving management, conservation, and restoration strategies. Here, we combine emerging knowledge from tropical forest succession, forest fragmentation and landscape ecology research to identify the main driving forces shaping successional pathways at different spatial scales. We also explore causal connections between land‐use dynamics and the level of predictability of successional pathways, and examine potential implications of such connections to determine the importance of secondary forests for biodiversity conservation in HMTLs. We show that secondary succession (SS) in tropical landscapes is a multifactorial phenomenon affected by a myriad of forces operating at multiple spatio‐temporal scales. SS is relatively fast and more predictable in recently modified landscapes and where well‐preserved biodiversity‐rich native forests are still present in the landscape. Yet the increasing variation in landscape spatial configuration and matrix heterogeneity in landscapes with intermediate levels of disturbance increases the uncertainty of successional pathways. In landscapes that have suffered extensive and intensive human disturbances, however, succession can be slow or arrested, with impoverished assemblages and reduced potential to deliver ecosystem functions and services. We conclude that: (i) succession must be examined using more comprehensive explanatory models, providing information about the forces affecting not only the presence but also the persistence of species and ecological groups, particularly of those taxa expected to be extirpated from HMTLs; (ii) SS research should integrate new aspects from forest fragmentation and landscape ecology research to address accurately the potential of secondary forests to serve as biodiversity repositories; and (iii) secondary forest stands, as a dynamic component of HMTLs, must be incorporated as key elements of conservation planning; i.e. secondary forest stands must be actively managed (e.g. using assisted forest restoration) according to conservation goals at broad spatial scales.