A GIS-based Model of the Distribution of the Rare Liverwort Aneura maxima at the Landscape Scale for an Improved Assessment of Its Conservation Status

Testing whether rare species are limited by narrow habitat conditions from correlations between species occurrence and measured environmental factors is usually hampered by the lack of statistical power associated to the low number of observations. Aneura maxima is an exceedingly rare liverwort in Europe whose recent discovery precluded the inclusion within the Red Data Book of European bryophytes. A series of new observations resulting from intensive bryophyte grid-mapping at the border between France and Belgium allowed the statistical investigation of the factors accounting for its distribution at the landscape scale. The species was systematically observed in deep ravines on damp, loamy soils under light tree cover within broadleaf woodlands. These conditions differ from those of other European localities, suggesting that A. maxima is not limited by a narrow ecological range. Attempting to predict the occurrence of A. maxima at the landscape scale from data on land use and soil conditions resulted in a model where the species was actually observed in only a half of the squares where its probability of occurrence was the highest. The species thus does not seem to have colonized all its potential habitats, which may be explained by the poor dispersal ability of this dioecious, rarely fertile species. Given the relative frequency of A. maxima in the Semois river basin, but taking into account its seemingly low dispersal ability and its preference for swampy habitats that are threatened by draining and spruce plantations, a status as 'conservation dependent' to 'near threatened' within the category 'lower risk' is proposed.     

Patterns of bryophyte diversity and rarity at a regional scale

Information on bryophyte diversity and rarity were combined withinformation on soil conditions and land use for Walloon Brabant (centralBelgium, 1091 km²) in order to investigate whichlandscape features sustain the most rare and diverse species assemblages.Presence–absence of 325 bryophyte species was recorded in 87 grid-squaresof 4 × 4 km. Species diversity was significantly correlatedwith forest cover (r = 0.71, P <0.001), sandy soils (r = 0.61, P <0.001), loamy soils, (r = –0.68,P < 0.001), and agricultural fields(r = –0.49, P < 0.001). Themost diverse grid-squares possessed up to 182 species and were characterized byat least 10% forest cover and the presence of unique micro-habitats.Grid-squares with forest cover reaching at least 10% but lacking uniquemicro-habitats contained between 90 and 130 species. Below 10% forest cover,diversity ranged between 55 and 110 species per grid-square. However, even theleast diverse cultivated areas included a significant amount of the regionallyrare species. A number of the latter are characteristic in other areas forspecific primary habitats lacking in Walloon Brabant but display an unexpectedability to disperse throughout hostile areas and colonize secondary habitats.The tendency of such species to occur in man-made habitats decreased our abilityto predict species richness and rarity from landscape features and soilconditions.     

影响广东黑石顶树附生苔藓分布的环境因子

通过对广东省黑石顶自然保护区内5个2500 m2样地内树附生苔藓的调查及有关环境因子的测定,研究了树附生苔藓的分布格局及其与环境因子的关系.树附生苔藓在不同高度的分布存在一定梯度,20 cm高处树附生苔藓的种类数与盖度均大于60 cm及更高处,且其群落优势种的数量组成与后者存在较大差异.不同树种附生苔藓盖度和种数差异较大.基于附生苔藓植物的盖度进行DCA排序及聚类分析将树种分成四组,马尾松(Pinus massoniana)因具有两种特有的网藓(巴西网藓Syrrhopodon prolifer和鞘刺网藓S.armatus)单独一组,福建青冈(Quercus chungii)亦与其余各种的差异均较大,形成一组,其余阔叶树种根据其所处的森林类型分成两组,针阔叶混交林内的阔叶树种和次生阔叶林内的阔叶树种各形成一组.对环境因子及树皮含水量和pH的分析显示,垂直梯度上空气湿度的差异可能是造成附生苔藓在不同高度分布差异的主要影响因子之一,不同树种附生苔藓的差异在一定程度上受树皮pH的影响,而与树皮含水量无关.同一树种上树附生苔藓的分布又在一定程度上受森林类型的影响.     

Traits reveal how habitat‐quality gradients structure small mammal communities in a fragmented tropical landscape

Understanding the relationships among community structure, vegetation structure and availability of food resources are a key to unravelling the ecological processes that structure biological communities. In this study, we tested (i) whether the composition of small mammal communities changed across gradients in habitat quality in tropical forest fragments, and (ii) whether any observed change could be explained by the functional traits of species. We sampled 24 trapping grids in fragments of semi‐deciduous forest, in each of two 6‐month periods. We considered each trapping grid as a sampling unit, for which we collected three datasets: an environmental matrix (vegetation structure and food resource availability), the abundance of small mammal species (community structure) and a matrix of functional traits (ecological and morphological traits which express tolerance to habitat disturbance and trophic guild). We used an RLQ approach to evaluate the association between traits and environmental gradients. Forest‐specialist and scansorial–arboreal species were associated with more complex habitat that had greater litter and canopy cover and more fallen logs. In relation to trophic guilds, granivore (fruit seeds), insectivorous and omnivorous species were also associated with higher complexity habitat, while frugivores were associated with shrub cover and availability of fruits. We conclude that functional traits (habitat use, use of vertical strata and diet) provide valuable insights into the distribution of small mammals along gradients of habitat quality in tropical forest fragments. We highlight that communities studies in fragmented landscapes should investigate the different components of biodiversity not only in landscape‐scale but also in habitat scale. Abstract in Portuguese is available with online material.     

Trends in diversity and abundance of obligate epiphytic bryophytes in a highly managed landscape

Although forest stands represent 47% of the total land area in Europe, alterations to the forest habitat through logging and plantation of exotic trees has led to significant changes in forest biocenoses. Due to their peculiar biology and life history, epiphytic bryophytes, which include a number of species of high conservation value, are especially concerned. Ordinal logit regression was used to test whether trends in diversity and abundance of obligate epiphytic bryophytes are explained by forest cover and spruce plantation and determine specific optima and degree of reliance to these factors at the landscape scale. Spruce plantations had a negative impact on both species diversity and abundance. Although large forest patches were important for a set of species exclusively or more frequently occurring under the forest cover, the abundance of a number of species previously identified as woodland bryophytes decreased or was uncorrelated with increasing forest cover. Furthermore, the species pool adapted to edge-related abiotic conditions was important. The global epiphytic diversity did consequently not decrease with decreasing forest cover at the landscape scale. If large forest patches are important for the conservation of a set of species exclusively or more frequently occurring under the forest cover, the conservation of epiphytic bryophytes thus also involves the conservation of pioneer trees in open landscapes. A series of management measures, which may help maximize the species diversity and probability of occurrence of key-species of high conservation interest, are proposed.     

BryForTrait – A life‐history trait database of forest bryophytes

In ecological research, plant functional trait analyses are widely applied to understand to what extent the inter‐specific variation in trait attributes has an adaptive value or to predict ecosystem processes and changes. Compared to vascular plants, trait studies using bryophytes are scarce, which is likely due to missing trait information for bryophyte species. With the BryForTrait database, we want to reduce this deficit. Our database represents a compilation of autecological information and morphological and regenerative trait data on different stages of the life cycle of bryophytes occurring in forest ecosystems. The database contains information for 35 traits and 721 Central European bryophyte species; in total more than 23,000 trait values. The BryForTrait database will enable future trait studies, providing new insights into bryophyte‐dominated ecosystems.     

Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

Light and warming drive forest understorey community development in different environments

Plant community composition and functional traits respond to chronic drivers such as climate change and nitrogen (N) deposition. In contrast, pulse disturbances from ecosystem management can additionally change resources and conditions. Community responses to combined environmental changes may further depend on land‐use legacies. Disentangling the relative importance of these global change drivers is necessary to improve predictions of future plant communities. We performed a multifactor global change experiment to disentangle drivers of herbaceous plant community trajectories in a temperate deciduous forest. Communities of five species, assembled from a pool of 15 forest herb species with varying ecological strategies, were grown in 384 mesocosms on soils from ancient forest (forested at least since 1850) and postagricultural forest (forested since 1950) collected across Europe. Mesocosms were exposed to two‐level full‐factorial treatments of warming, light addition (representing changing forest management) and N enrichment. We measured plant height, specific leaf area (SLA) and species cover over the course of three growing seasons. Increasing light availability followed by warming reordered the species towards a taller herb community, with limited effects of N enrichment or the forest land‐use history. Two‐way interactions between treatments and incorporating intraspecific trait variation (ITV) did not yield additional inference on community height change. Contrastingly, community SLA differed when considering ITV along with species reordering, which highlights ITV’s importance for understanding leaf morphology responses to nutrient enrichment in dark conditions. Contrary to our expectations, we found limited evidence of land‐use legacies affecting community responses to environmental changes, perhaps because dispersal limitation was removed in the experimental design. These findings can improve predictions of community functional trait responses to global changes by acknowledging ITV, and subtle changes in light availability. Adaptive forest management to impending global change could benefit the restoration and conservation of understorey plant communities by reducing the light availability.     

An intraspecific application of the leaf‐height‐seed ecology strategy scheme to forest herbs along a latitudinal gradient

We applied the leaf‐height‐seed (LHS) ecology strategy scheme (a combination of three ecologically important traits: specific leaf area (SLA), seed mass and plant height) intraspecifically to two widespread European forest herbs along a latitudinal gradient. The aims of this study were to quantify LHS trait variation, disentangle the environmental factors affecting these traits and compare the within‐species LHS trait relationships with latitude to previously established cross‐species comparisons. We measured LHS traits in 41 Anemone nemorosa and 44 Milium effusum populations along a 1900–2300 km latitudinal gradient from N France to N Sweden. We then applied multilevel models to identify the effects of regional (temperature, latitude) and local (soil fertility and acidity, overstorey canopy cover) environmental factors on LHS traits. Both species displayed a significant 4% increase in plant height with every degree northward shift (almost a two‐fold plant height difference between the southernmost and northernmost populations). Neither seed mass nor SLA showed a significant latitudinal cline. Temperature had a large effect on the three LHS traits of Anemone. Latitude, canopy cover and soil nutrients were related to the SLA and plant height of Milium. None of the investigated variables appeared to be related to the seed mass of Milium. The variation in LHS traits indicates that the ecological strategy determined by the position of each population in this three‐factor triangle is not constant along the latitudinal gradient. The significant increase in plant height suggests greater competitive abilities for both species in the northernmost populations. We also found that the studied environmental factors affected the LHS traits of the two species on various scales: spring‐flowering Anemone was affected more by temperature, whereas early‐summer flowering Milium was affected more by local and other latitude‐related factors. Finally, previously reported cross‐species correlations between LHS traits and latitude were generally unsupported by our within‐species approach.     

Predictability of plant species composition from environmental conditions is constrained by dispersal limitation

Despite recent modelling approaches integrating the effects of niche-based processes and dispersal-based processes on local plant species composition, their relative importance is still not clear. We test whether the predictability of local species composition from environmental conditions is influenced by dispersal traits. We analyzed a large database with co-occurrence data, using ordination techniques (DCA and CCA) to identify the major environmental determinants of species composition. The percentage of explained variance in occurrence was quantified for individual species with CCA. Effects of life-history traits on the predictability of occurrence patterns were tested by means of regression analysis, using a generalized linear models approach. The results reveal close correlations between species composition and environmental conditions, implying that the predictability of the set of species that might occur in a given environmental setting ("habitat species pool") is high. The habitat species pool, however, reflects the potential species composition, and not the actual local situation. At the level of individual species, a large proportion (>90%) of the variation in occurrence remained unexplained. Predictability of species occurrence patterns was increased by a greater capacity for long-distance dispersal, greater adult longevity and the capacity to build a persistent seed bank. The results indicate that the predictability of species composition from environmental conditions is reduced by a few orders of magnitude by dispersal limitation and that poor dispersers are underrepresented.     

Influence of Microclimate on the Occurrence of Cyanobacteria in the Phyllosphere in a Premontane Rain Forest of Costa Rica

Abstract: The occurrence of cyanobacteria in the phyllosphere of several understory plants was investigated during a one and a half year study in a primary premontane rain forest in Costa Rica. Altogether, seven species of epiphyllous cyanobacteria were identified and are discussed here. The ecology of the two most frequent species in the understory, Scytonema javanicum and Sc. hofmannii , was studied in further detail. Within the Understory, the relative abundance of these species, as well as of epiphyllous bryophytes, was more influenced by air humidity than by light. The humidity, characterized as potential evaporation, also had influence on the pseudosuccession of the phyllosphere vegetation. While at moist sites bryophytes and both species of Scytonema appeared nearly simultaneously on 6–9-month-old leaves, at dryer sites Sc. javanicum , and especially Sc. hofmannii , appeared 6–9 months after bryophytes had established. On 13–15-month-old leaves bryophytes and Scytoriema were usually well established. On average, 20–30% of the leaf area of 2–5-year old leaves of Spathacanthus hoffmannii (Acan-thaceae, an abundant understory shrub) were covered by bryophytes, 2–3% by Sc. javanicum , and 0.1–0.2% by Sc. hofmannhi . An influence of phorophyte species on the occurrence and abundance of these two species of Scytonema could not be detected. However, Sc. hofmannii was more frequent on leaves with higher bryophyte cover, while Sc. javanicum was independent from bryophytes. A comparison of leaves from the understory and from the canopy showed that the composition and abundance of cyanobacteria species changed with height above forest floor.     

Spatio‐temporal colonization patterns of forest plant species in a mixed deciduous forest

Abstract. An integrated analysis of the colonization patterns of forest plant species was carried out in a 34‐ha, mixed deciduous forest in northern Belgium. First, we sought to describe the relationships between land use history and environmental conditions. Land use history and soil type were related and negative correlations between pH and secondary forest age were found. The density of the shrub layer increases with secondary forest age. Litter quantity and cover of Urtica dioica were mainly indirectly influenced by land use history. Litter starts accumulating at low pH values and high shrub density and Urtica dioica grows vigorously on nutrient enriched soils where much light can reach the ground. Next, the importance of these human‐altered environmental conditions for the colonization of forest plant species was assessed relative to the importance of dispersal limitation. Therefore, the distribution of 16 forest species was mapped and species‐specific spatio‐temporal isolation measures were calculated. The analysis revealed that the colonization patterns of the slowly colonizing species (i.e. ‘ancient forest plant species’) are best explained by a combination of spatio‐temporal isolation, soil type, pH and the (non‐)cover of Urtica dioica. By contrast, spatio‐temporal isolation was never a limiting factor for good colonizing forest species. Our results suggest that colonization of ‘ancient forest plant species’ is hampered by a combination of dispersal‐ and recruitment limitation and that the relative importance of both factors is species‐specific.     

Diversity and distribution pattern of epiphyllous liverworts and its ecological determinants

Epiphyllous liverworts form a special group of bryophytes that primarily grow on leaves of understory vascular plants, occurring in constantly moist and warm evergreen forest in tropical and subtropical regions. Epiphyllous liverworts may influence ecosystem processes including carbon, nitrogen and water cycles. Furthermore, they are very sensitive to climate change and forest fragmentation, and can be used as bioindicators for changes in ecological conditions and escalating loss of biodiversity. In this paper, we reviewed studies on morphological traits, species diversity, geographical distribution and environmental conditions (including characteristics of hosts and habitats) of epiphyllous liverworts, discussed the possible reasons for the mechanisms underlying the diversity pattern of epiphyllous liverworts. According to these studies, we proposed that further ecological studies on epiphyllous liverworts should be more focused on their formation, mechanisms of matter exchange and energy flux, ecological functions in forest ecosystem, the response to climate changes and their broader-scale ecology.     

Applying life-history strategies for freshwater macroinvertebrates to lentic waters

• 1 Effective environmental management requires a sound understanding of the causal mechanisms underlying the relationship of species with their environment. Mechanistic explanations linking species and environment are ultimately based on species traits. Many tools for ecological assessment and biomonitoring lack such explanations. Elsewhere, we have defined life‐history strategies, based on the interrelations between species traits and their functional implications.
• 2 This study tests the hypothesis that life‐history strategies represent different solutions to particular ecological problems, thus connecting species and their environment through species traits. Data on aquatic macroinvertebrates in a variety of waterbodies were analysed in terms of life‐history strategies. These waterbodies differed in environmental conditions and macroinvertebrate assemblages. Solutions to the ecological problems present in each type of water body were expected to be reflected in the abundance of (species exhibiting) different life‐history strategies.
• 3 Results show clear differences in strategy composition between the different water types, which could be related to the prevailing environmental conditions through mechanistic explanations. For example, species with a long period of juvenile development and a synchronized emergence of short‐lived adults were most dominant in large mesotrophic waterbodies with stable and predictable environmental conditions. In contrast, species that have a rapid development and spread successive reproduction attempts over a longer time period were most abundant in waterbodies with fluctuating and less predictable environmental conditions.
• 4 Differences in strategy composition provide insight into the prevailing environmental conditions related to temporal predictability, and habitat favourability, from the perspective of the species themselves. By reducing diverse species assemblages to a small number of strategies, representing easily interpretable relationships, this approach may be useful in environmental quality assessment programmes, including those required by the European Water Framework Directive. Based on mechanistic explanations, life‐history strategies may generate testable predictions and guide future research. Further research may focus on expanding life‐history strategies to include other species groups and ecosystems.

     

Phylogenetic and morphological relationships between nonvolant small mammals reveal assembly processes at different spatial scales

The relative roles of historical processes, environmental filtering, and ecological interactions in the organization of species assemblages vary depending on the spatial scale. We evaluated the phylogenetic and morphological relationships between species and individuals (i.e., inter‐ and intraspecific variability) of Neotropical nonvolant small mammals coexisting in grassland‐forest ecotones, in landscapes and in regions, that is, three different scales. We used a phylogenetic tree to infer evolutionary relationships, and morphological traits as indicators of performance and niche similarities between species and individuals. Subsequently, we applied phylogenetic and morphologic indexes of diversity and distance between species to evaluate small mammal assemblage structures on the three scales. The results indicated a repulsion pattern near forest edges, showing that phylogenetically similar species coexisted less often than expected by chance. The strategies for niche differentiation might explain the phylogenetic repulsion observed at the edge. Phylogenetic and morphological clustering in the grassland and at the forest interior indicated the coexistence of closely related and ecologically similar species and individuals. Coexistence patterns were similar whether species‐trait values or individual values were used. At the landscape and regional scales, assemblages showed a predominant pattern of phylogenetic and morphological clustering. Environmental filters influenced the coexistence patterns at three scales, showing the importance of phylogenetically conserved ecological tolerances in enabling taxa co‐occurrence. Evidence of phylogenetic repulsion in one region indicated that other processes beyond environmental filtering are important for community assembly at broad scales. Finally, ecological interactions and environmental filtering seemed important at the local scale, while environmental filtering and historical colonization seemed important for community assembly at broader scales.     

Succession pattern at two mineralogically different spoil heaps in Central Slovakia

Plants colonize post-mining areas such as spoil heaps even without human interference, by spontaneous succession. Knowledge of this process, its direction and role in shaping the vegetation is desirable not only from an ecological point of view but also in the application of remediation techniques of such environmental burdens. The aim of our study was to analyse plant species composition and to describe a succession pattern at two mineralogically different spoil heaps near the city of Banská Bystrica (central Slovakia). Hierarchical cluster analysis was used in PC-ORD software to classify physiognomic vegetation types (PVT) at heaps and synoptic tables were generated in Juice software according to species percentage of frequency and their median cover. At the Hg-heap Ve?ká studňa we recorded a total of 335 taxa of vascular plants and 83 taxa of bryophytes within 21 identified PVT, which were grouped into 4 clusters (forest, wetland, grassland and ruderal cluster). A total of 111 taxa of vascular plants and 76 taxa of bryophytes were recorded at the Cu-heap Podlipa and here 9 PVT formed 3 clusters (initial, non-forest and forest cluster). At both heaps PVT, represented as individual successional stages, formed a distinct continuum along the moisture gradient (in dry and wet areas). We can consider both heaps as species-rich with different diagnostic and constant species and a development of various microhabitats. In addition, bryophytes greatly contribute to species composition and richness and their assamblages in initial stages of succession can be important towards the higher ones.     

Environmentally and behaviourally mediated co‐occurrence of functional traits in bird communities of tropical forest fragments

Two major theories of community assembly – based on the assumption of ‘limiting similarity’ or ‘habitat filtering’, respectively – predict contrasting patterns in the spatial arrangement of functional traits. Previous analyses have made progress in testing these predictions and identifying underlying processes, but have also pointed to theoretical as well as methodological shortcomings. Here we applied a recently developed methodology for spatially explicit analysis of phylogenetic meta‐community structure to study the pattern of co‐occurrence of functional traits in Afrotropical and Neotropical bird species inhabiting forest fragments. Focusing separately on locomotory, dietary, and dispersal traits, we tested whether environmental filtering causes spatial clustering, or competition leads to spatial segregation as predicted by limiting similarity theory. We detected significant segregation of species co‐occurrences in African fragments, but not in the Neotropical ones. Interspecific competition had a higher impact on trait co‐occurrence than filter effects, yet no single functional trait was able to explain the observed degree of spatial segregation among species. Despite high regional variability spanning from spatial segregation to aggregation, we found a consistent tendency for a clustered spatial patterning of functional traits among communities in fragmented landscapes, particularly in non‐territorial species. Overall, we show that behavioural effects, such as territoriality, and environmental effects, such as the area of forest remnants or properties of the landscape matrix in which they are embedded, can strongly affect the pattern of trait co‐occurrence. Our findings suggest that trait‐based analyses of community structure should include behavioural and environmental covariates, and we here provide an appropriate method for linking functional traits, species ecology and environmental conditions to clarify the drivers underlying spatial patterns of species co‐occurrence.     

叶附生苔植物物种多样性分布格局及生态成因

叶附生苔植物(以下简称叶附生苔)是一类附生在维管植物叶片表面的、一般只出现在热带雨林和常绿阔叶林中的苔类植物。它们具有重要的生态功能, 如影响碳、氮、水循环, 对气候变化和森林破碎化反应敏感, 可用作环境变化的指示植物。该文对叶附生苔的形态特性、物种多样性的研究历史和进展、地理分布格局, 以及对环境的要求(附主和生境的特性)等进行了综述, 探讨了叶附生苔多样性分布格局形成的可能原因, 即环境(空气湿度高、林冠层发育好、干扰少)和适应特征(无性繁殖、形体微小)的选择。根据全球叶附生苔研究进展状况及所存在的问题提出未来可开展的研究方向, 如叶附生苔的形成原因、与附主间的物质交换和能量流动机制、在森林生态系统中的生态功能、气候变化指示作用研究等, 大尺度上的研究也值得关注。     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.