Plant species diversity in Mediterranean old-growth forests: A case study from central Italy

Abstract

To investigate the differences in understorey composition and diversity between old-growth and managed forests, we analyzed an old-growth and a managed beech stand in the same area displaying similar abiotic features. We considered variations in understorey species composition and richness. The sampled understorey species were characterized in terms of functional traits, Ellenberg's indicator values and taxonomic distinctness; next, we calculated four different pairwise plot-to-plot dissimilarity matrices based on species composition, functional traits, Ellenberg's indices and taxonomic distances. We applied a permutational multivariate extension of ANOVA to test whether the forest stands significantly differ in the considered features. Indicator values of all plant species in managed and old-growth stands were evaluated.

The old-growth forest had a higher species richness; permutational analysis of variance showed significant differences between the two stands in plant species composition, functional traits, Ellenberg indices and taxonomic distances. Indicator species analysis highlighted 14 indicator species for the unmanaged stand, while only 3 indicators were found for the managed one.

The results suggest that forest management determines ecological differences that strongly affect plant species composition.

The knowledge of natural stands dynamics could allow development of new approaches and practices in forest management focusing on biodiversity conservation.     

Structural variables drive the distribution of the sensitive lichen Lobaria pulmonaria in Mediterranean old-growth forests

We tested the hypothesis that structural variables related to old-growth features affect the distribution of Lobaria pulmonaria in a Mediterranean National Park of Italy. A total of 36 plots, with old-growth characteristics and representing overall three forest types (beech- oak- and mixed- forests) were studied. The lichen was absent in about half of the sites, suggesting that the selection of old-growth forests based solely on structural features is not sufficient to predict the presence of this species, which therefore proves to be rather sensitive and selective. Its abundance was related to high tree circumference and basal area, and to availability of deadwood, confirming the relevant effect of substrate and stand continuity in the conservation of this species and emphasizing its role of indicator species of undisturbed forest ecosystems with long ecological continuity.     

Loss of biodiversity and shifts in aboveground biomass drivers in tropical rainforests with different disturbance histories

Tropical forests account for more than half of the global carbon forest stock and much of the biological diversity on Earth. However, disturbances such as deforestation and forest degradation threaten the maintenance of these ecosystem services. This study aimed to understand how different disturbance histories affect the forest stand biomass, as well as species and functional diversity, and to what extent these differences can change the relationships between biomass and their drivers. We used data from forests with clear-cut and selectively logged disturbance histories, and from old-growth forests, situated in the Brazilian Atlantic forest. Forests with logging disturbances showed significant losses in their aboveground biomass compared to those of old-growth forests (50% loss in selectively logged forests and 80% loss in clear-cut forests). Interestingly, only clear-cut secondary forests showed differences in species and functional diversity, and were dominated by species with acquisitive trait values, commonly found early in succession. Shifts in stand biomass drivers were observed in selectively logged forests. The mass-ratio hypothesis (mainly through the functional trait of maximum height) was the most important biomass driver in clear-cut secondary and old-growth forests, whereas the importance of the niche complementarity hypothesis (through functional richness and dispersion) was higher in selectively logged forests. Our study highlights that disturbance histories can affect forest aboveground biomass and its drivers. Moreover, our results reinforce the need for conservation of intact forests but highlight the importance of including degraded forests in conservation mechanisms based in carbon stocks, as these forests retain high values of species and functional diversities that are crucial to biomass and consequently carbon stock acquisition.     

Impact of a strong typhoon on the structure and dynamics of an old-growth beech (<Emphasis Type="Italic">Fagus crenata</Emphasis>) forest,southwestern Japan

Typhoon no. 19 of 1991 (T9119) caused multiple treefalls and created large openings in an old-growth beech (Fagus crenata) forest at Mt. Daisen, in the Daisen Forest Reserve, southwestern Japan. The area of the largest opening was about 1.7 ha (300 m by 70 m). To predict the dynamics of the beech stand after the disturbance of T9119, we investigated the damage to the stand and the density and growth rate of trees with DBH=5–10 cm in a 1-ha plot covering a large part of the largest opening and the adjacent closed canopy. The beech did not regenerate immediately. The regeneration and growth rate of trees with DBH=5–10 cm were related to the frequency of the typhoon attack for at least the past century. In beech forests, small gap formation is the prevailing mode of disturbance. Our results indicate that typhoons affect the structure and dynamics of this beech stand. We suggest that both small gap formation and large-scale disturbance are important for the maintenance of beech forest in some areas.     

The scientific value of the largest remaining old-growth red pine forests in North America

Old growth red pine forests (Pinus resinosa) cover less than 1% of their original range in North America and are essential for maintaining biodiversity at stand and landscape scales. Despite this, the largest remaining old-growth red pine forest in the world, the Wolf Lake Forest Reserve, is currently threatened by mining claims in Northern Ontario and has been receiving considerable media and public attention in recent months. We provide a timely review of how large old growth red pine forests maintain biodiversity at several taxonomic levels (with a focus on trees and plants) through heterogeneous partitioning of limiting resources such as light and nitrogen, formation of complex habitats through increased accumulation of coarse woody debris, and the maintenance of natural disturbance-driven succession. These processes shape the overstory community, allowing for the regeneration of pines, coexistence of early-mid successional shade intolerant species and cross-ecotonal establishment of late successional tree species in response to regional warming over the past three decades. Using Wolf Lake as a case study, we review legislation and policy complexities around this issue and provide scientific arguments for the preservation of this forest. We invoke recent insights into the ecological role of refugia, the development of criteria for assessing endangered ecosystems, and the challenges of conservation in the face of climate change and disturbance regimes. These forests are ecologically important and provide a scientifically irreplaceable system for assessing baseline ecosystem function, processes and services. As the largest remaining old-growth red pine forest in the world, Wolf Lake Forest Reserve deserves intensive study, monitoring and full protection from future development.     

Nuovi reperti di intersessualità nelle Cucurbitacee

Abstract

The aim of this paper is to investigate differences in plant species composition between managed and unmanaged forests, and to assess if these difference give rise to a higher plant diversity in the unmanaged forest. Furthermore our aim is to relate forest structure to differences in plant species composition, identifying the structural attributes more strongly related to the unmanaged forest vegetation. We compared an old-growth forest and a managed highforest in the Abruzzo Lazio and Molise National Park (Central Italy). Plant species composition and diversity, deadwood components and live structure have been analyzed. We used permutational multivariate analysis of variance to test the response of species composition to management factor; furthermore, we compared species richness and beta diversity. Redundancy analysis has been used to relate plant species abundances to structural variables; the importance of dead and living wood components has been compared through variation partitioning. Plant species composition proved to be significantly different in the two sites, and the old-growth stand showed a higher plant diversity. From a structural point of view, we found differences especially in the amount and quality of deadwood, and in the diameter class distribution. These variables are also the most important in determining the old-growth stand plant species composition according to redundancy analysis. Variation partitioning confirmed the greater importance of the deadwood variables. Our results suggest that including deadwood surveys in traditional forest inventories could help in finding forests with both structural and floristic old-growth properties to be considered in conservation programmes. The imitation of natural dynamics, through the creation of gaps avoiding deadwood removal, could be an effective strategy for restoring old-growth conditions, also in terms of plant diversity.     

云南省国家级和省级自然保护区森林生态服务功能价值评估

采取文献查阅、资料数据处理、统计分析等研究方法,选取合理、成熟、可操作的森林生态服务评估体系和方法（LY/T 1721-2008）,测算出云南省国家级、省级自然保护区森林生态系统在涵养水源、保育土壤、固碳释氧、积累营养物质、净化大气环境、生物多样性保护等6类生态系统服务功能的物质量和价值量,并从森林类型、质量、立地状况等多方面进行了分析。经评估,云南省自然保护区森林生态系统服务年总价值为2009.02亿元。其中,涵养水源价值为538.75亿元;保育土壤价值为493.79亿元;固碳释氧价值为122.09亿元;积累营养物质价值为16.11亿元;净化大气环境价值为83.21亿元;生物多样性保护价值为755.07亿元。保护区每年每公顷森林生态系统服务价值平均为12.31万元,相当于全国平均水平的2.9倍,云南省平均水平的2.4倍。     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

福建集体林权制度改革对生态公益林水源涵养能力变化的影响——基于723个村的实证研究

水源涵养等生态功能是生态公益林的首要功能,维护生态安全是集体林权制度改革（简称"林改"）的重要目标。为了厘清福建林改对生态公益林水源涵养能力变化的影响,以促进生态公益林质量的精准提升。运用综合蓄水能力法测算分析了福建省5县（区）723个村林改前后生态公益林水源涵养能力变化,并运用多元线性回归模型探讨了林改对生态公益林水源涵养能力变化的关键影响因素。结果表明：（1）林改后生态公益林水源涵养量普遍增加,但单位面积水源涵养能力多数下降,少数提升。（2）产权归属通过经营管护行为对单位面积水源涵养能力产生显著影响,其中,国有权属存在正向影响,集体权属与私有权属存在负向影响。（3）面积、人工林占比、防护林占比、蓄积量变动和降雨量等因素,主要通过资源结构、经营条件和外部环境对单位面积水源涵养能力产生显著影响。据此提出,在提高生态公益林数量的同时要注重提升生态公益林质量,加大政府赎买,重点挖掘幼龄人工生态林生态潜力,做好天然林和防护林保护等政策建议。     

Stand selection by birds in Scots pinewoods in Scotland: the need for more old-growth pinewood

A study at Abernethy Forest, Scotland, examined stand selection by some of the birds of conservation concern that inhabit pinewoods. Abernethy Forest was ideally suited for the study because all four main stand types of forest development (stand initiation, stem exclusion, understorey reinitiation and old-growth) were represented. Capercaillies Tetrao urogallus (both males and females) and crossbills Loxia spp. preferred old-growth pinewood in winter, and Redstarts Phoenicurus phoenicurus and Tree Pipits Anthus trivialis preferred old-growth in summer. Great Spotted Woodpecker Dendrocopos major selected snags in old-growth to excavate nest and roost sites. In winter, Crested Tits Parus cristatus had a weak selection for old-growth stands and Bullfinches Pyrrhula pyrrhula preferred stands at initiation. Possible reasons for these preferences are put forward. Although the area of conifer woodland in Scotland is 888 000 ha, it largely comprises North American species destined to be cut before reaching old-growth. Old-growth pinewood in Scotland is largely confined to the fragments of ancient native pinewood, amounting to only 16 000 ha. To conserve some pinewood birds of conservation concern, a mechanism to allow more Scots Pine Pinus sylvestris woods to reach and remain in an old-growth state needs to be pursued.     

Diversity of wood-decaying fungi under different disturbance regimes—a case study from spruce mountain forests

Rapid destruction of forest habitats has led to the establishment of protected areas in formerly managed forests with the aim of restoring biodiversity. Conservation in spruce-dominated reserves is often contradicted by salvage logging after insect outbreaks. Here we study the community characteristics of wood decaying fungi in a high montane Norway Spruce forest with three different management types: (1) a formerly managed area disturbed by a large-scale bark beetle outbreak, (2) an area with continuous salvage logging, and (3) an old-growth forest. Bark beetle activity in the disturbed area resulted in downed wood amounts comparable to those of the old-growth forest. However, species accumulation curves for the disturbed forest were more similar to those of the logged forest than to those of the old-growth forest. This arose because of differences in the diversity of wood decay classes; wood decay in the disturbed forest was more homogeneous. Logs in the disturbed forest originated almost exclusively from bark-beetle-infested trees, but the causes of tree mortality in the old-growth forest were manifold. Although most red-listed species were clearly confined to old-growth forest, Antrodiella citrinella was most abundant in the disturbed forest. Our analysis furthermore showed that the between stand scale is the most effective unit for diversity wood-decaying fungi. We therefore suggest a conservation strategy for preserving old-growth forests and establishing protected forest stands to enhance structural heterogeneity in spruce-dominated forests. For this, a careful screening of protected areas throughout Europe is necessary to provide managers with guidelines for conservation.     

Long-Term Impacts of Forest Ditching on Non-Aquatic Biodiversity: Conservation Perspectives for a Novel Ecosystem

Artificial drainage (ditching) is widely used to increase timber yield in northern forests. When the drainage systems are maintained, their environmental impacts are likely to accumulate over time and along accompanying management, notably after logging when new forest develops on decayed peat. Our study provides the first comprehensive documentation of long-term ditching impacts on terrestrial and arboreal biodiversity by comparing natural alder swamps and second-generation drained forests that have evolved from such swamps in Estonia. We explored species composition of four potentially drainage-sensitive taxonomic groups (vascular plants, bryophytes, lichens, and snails), abundance of species of conservation concern, and their relationships with stand structure in two-ha plots representing four management types (ranging from old growth to clearcut). We found that drainage affected plot-scale species richness only weakly but it profoundly changed assemblage composition. Bryophytes and lichens were the taxonomic groups that were most sensitive both to drainage and timber-harvesting; in closed stands they responded to changed microhabitat structure, notably impoverished tree diversity and dead-wood supply. As a result, natural old-growth plots were the most species-rich and hosted several specific species of conservation concern. Because the most influential structural changes are slow, drainage impacts may be long hidden. The results also indicated that even very old drained stands do not provide quality habitats for old-growth species of drier forest types. However, drained forests hosted many threatened species that were less site type specific, including early-successional vascular plants and snails on clearcuts and retention cuts, and bryophytes and lichens of successional and old forests. We conclude that three types of specific science-based management tools are needed to mitigate ditching effects on forest biodiversity: (i) silvicultural techniques to maintain stand structural complexity; (ii) context-dependent spatial analysis and planning of drained landscapes; and (iii) lists of focal species to monitor and guide ditching practices.     

Ground Layer Plant Species Turnover and Beta Diversity in Southern-European Old-Growth Forests

Different assembly processes may simultaneously affect local-scale variation of species composition in temperate old-growth forests. Ground layer species diversity reflects chance colonization and persistence of low-dispersal species, as well as fine-scale environmental heterogeneity. The latter depends on both purely abiotic factors, such as soil properties and topography, and factors primarily determined by overstorey structure, such as light availability. Understanding the degree to which plant diversity in old-growth forests is associated with structural heterogeneity and/or to dispersal limitation will help assessing the effectiveness of silvicultural practices that recreate old-growth patterns and structures for the conservation or restoration of plant diversity. We used a nested sampling design to assess fine-scale species turnover, i.e. the proportion of species composition that changes among sampling units, across 11 beech-dominated old-growth forests in Southern Europe. For each stand, we also measured a wide range of environmental and structural variables that might explain ground layer species turnover. Our aim was to quantify the relative importance of dispersal limitation in comparison to that of stand structural heterogeneity while controlling for other sources of environmental heterogeneity. For this purpose, we used multiple regression on distance matrices at the within-stand extent, and mixed effect models at the extent of the whole dataset. Species turnover was best predicted by structural and environmental heterogeneity, especially by differences in light availability and in topsoil nutrient concentration and texture. Spatial distances were significant only in four out of eleven stands with a relatively low explanatory power. This suggests that structural heterogeneity is a more important driver of local-scale ground layer species turnover than dispersal limitation in southern European old-growth beech forests.     

Stand structure, species diversity and regeneration of an endemic palm forest on the Seychelles

The ‘Vallée de Mai’ on the island of Praslin in the Seychelles group is a World Heritage Site because of its unique forest vegetation dominated by the palm Lodoicea maldivica. However, despite its great conservation importance, the stand structure, species diversity and regeneration of this forest have not previously been studied. Moreover, the conservation value of the reserve is currently threatened by invading alien plants and limited regeneration of some native species. The species composition of the forest was studied in six permanent plots (1985 and 1998) and along seven trail transects (1997). To explain potential variation in tree regeneration, the light climate within all plots was investigated using hemispherical photographs. The results reveal little change in forest structure, species diversity and regeneration in the permanent plots over the 13‐year period, but some regeneration of most woody species, presence of some alien species and a relatively high species turnover. We found no significant influence of local light climate upon forest structure, species diversity or regeneration. Although the stand characteristics seem to be relatively stable, the permanent plots and transects should be monitored in order to detect future changes in stand structure and to optimize the protection of this unique forest reserve.     

Spatial Variation of Bird Species in Landscapes Dominated by Old-growth Forests in Northern Boreal Finland

An important issue in conservation planning is to study the distribution and abundance patterns of species in natural landscapes. Information of concordant and clumped species distribution pattertns can enhance attempts to focus conservation efforts in sites/areas preferred by various species. We compared bird species abundance, community composition and species distribution patterns in three large old-growth forest areas (size 40–120 km²) in northern boreal Finland on the basis of quantitative bird censuses. Total bird density and species composition, and mean density and variances of the most abundant species were highly similar between the three areas. Most of the bird species were distributed randomly within the areas and species preferring old-growth forests showed compensatory density variation without preference for any sub-area. Mean density of species preferring old-growth forests was significantly higher in the studied large old-growth forests than in smaller old-growth forests and was about threefold higher than regional density of species in a predominantly managed landscape matrix. Bird species preferring old-growth forests are probably not dependent on a certain specific part of a virgin boreal forest landscape but rather on the overall size of the high-quality and diverse old-growth forest area.     

Attitudes toward forest diversity and forest ecosystem services—a cross-cultural comparison between China and Switzerland

Aims Despite the current interest in services provided by ecosystems and the role of biodiversity, the relationship among human attitudes, biodiversity and ecosystem services has hardly been investigated. Moreover, few studies have examined attitudes toward nature in cross-cultural comparisons. This study investigates the attitudes of Chinese and Swiss people, both environmental experts and laypersons, toward forest biodiversity and ecosystem services.Methods Overall, 640 people in China and Switzerland were interviewed with the help of a standardized questionnaire. In each country, the study population was equally divided into an urban (80 city dwellers and 80 environmental science students) and a rural (80 forest visitors and 80 farmers) study group. The 15-minute interviews took place in the cities of Beijing and Zurich and in the rural forested areas of Dujiangyan, Sichuan Province and Lake Sempach, canton Lucerne. Attitudes toward forest biodiversity were investigated with the help of color photographs that depicted both monocultures and species-rich forests typical for China and Switzerland. Attitudes toward ecosystem services were investigated with the help of 13 statements on provisioning, regulating, cultural and supporting services of forests.Important findings On average, Chinese participants showed no strong preferences for biodiversity, whereas the Swiss clearly preferred species-rich forests over monocultures. However, Chinese environmental science students did prefer species-rich forests and attributed to them a higher conservation value because of their higher biodiversity. Although there were no strong preferences for Chinese versus Swiss forests, all participants correctly answered that Chinese forests are more species rich in terms of plants and animals and thus found them less boring and more interesting, but also less managed, than Swiss forests. All participants highly valued the ecosystem services provided by forests; especially the regulating and supporting ones. Environmental science students and farmers placed more importance on the provisioning services, whereas city dwellers and forest visitors emphasized more on the regulating services. The disjuncture between the high ecological quality of species-rich forests and their low attractiveness to Chinese study participants points to a potential conflict between conservation policies and the public's preferences. A better communication of ecosystem services provided by forest biodiversity to the public might change these preferences in favor of ecological quality, as already observed among Chinese environmental science students.     

Changes in the decomposer community when converting spruce monocultures to mixed spruce/beech stands

The impact of the admixture of beech in spruce monocultures on structure and function of the decomposer community was studies in the Ore mountains (Saxony, Germany) on Dystric Cambisols between 2000 and 2002. The study sites represented four stages of forest conversion from a mature spruce stand to a mature spruce/beech stand. There, the functional profile of the nematodes, enchytraeids, lumbricids, and dipterans was analysed on the basis of ecological guilds, and their metabolic equivalences were calculated to characterize the decomposition potential of the invertebrate decomposer community. Because of the acidic parent soil the coenoses at all study sites were dominated by the enchytraeids with increasing importance of lumbricids and dipterans in progress of forest conversion. Gradual changes with rising coverage of beech culminated in intense differences of entire biomasses and metabolic equivalences between the mature stands, indicating a higher decomposition potential of the invertebrate decomposer community by the admixture of beeches in spruce forests. The quality of the beech litter is likely to be the important factors for these changes. To prove this assumption further investigations of the saprovore food chain are necessary, taking microbial parameters into account.     

Restoration of beech forest for saproxylic beetles—effects of habitat fragmentation and substrate density on species diversity and distribution

The influence of spatial location and density of beech snags on species diversity and distribution patterns of saproxylic beetles was studied in a 2,400 ha forest landscape in southern Sweden. Complete snag surveys were combined with a beetle survey using small window traps directly attached to the beech snags. The density of beech snags ≥30 cm dbh varied between one and seven snags per ha within the study area, corresponding to 1.1–5.1 m³/ha. A total of 2,610 specimens of 180 saproxylic beetles species were trapped, of which 19 species were red-listed. Within the study area, the number of red-listed and formerly red-listed species was highest around traps in old-growth stands, intermediate in managed stands contiguous with old-growth and lowest in managed stands isolated from old-growth by a two km-wide zone without beech forest. Logistic regressions revealed negative relationships between distance to old-growth forest and occurrence of eleven species, among them six red-listed or formerly red-listed species. The number of non red-listed species was not correlated with isolation from old-growth forest. The number of red-listed species also increased with snag density within 200–300 m around the traps. Our results suggest that red-listed species generally have a lower dispersal capacity than other saproxylic beetles. We conclude that retention of dead wood close to existing populations is more beneficial for red-listed species than an even distribution of snags across the forest landscape.     

Old rural parks support higher biodiversity than forest remnants

One of the main challenges in biodiversity conservation is to curb a further degradation and loss of high-quality habitats. In agricultural matrix landscapes, the detection of alternative habitats for habitat specialists may be a solution. Historic old parks or landscape gardens around manor houses and castles are cultural heritage of nobles, but their value in harbouring biodiversity is poorly acknowledged. Therefore we evaluated the potential of old rural parks to serve as a habitat for nemoral forest species. We recorded stand structure and the presence of forest biodiversity indicators in 74 closed-canopy stands of historic parks and compared them with 93 neighbouring mature forest remnants on ancient forest land. We estimated the importance of stand structure in relation to habitat type on biodiversity indicators. Finally we suggest single-value indicator-complexes for the cost-efficient assessment of the conservation value of forests and forest-like habitats. Park stands outclassed reference forests in several individual structural characteristics, and in combined indicators of habitat quality and biodiversity. Forests had higher estimates for the combined indicator of dead wood, but large-diameter dead wood types were more abundant in parks. Woodpeckers, several old-growth indicator epiphytes and forest herbs had successfully become established in planted forest-like park fragments. Old rural parks resemble high-conservation-value forests more than the best preserved contemporary forest remnants. After the century needed to overcome immigration delay, old parks do provide a refugium for temperate deciduous forest species. Consequently, biodiversity-targeted management should retain and enhance old-growth attributes in forests and on the peripheries of parks: e.g. preserving old trees to provide service for epiphytes, hollow trees and an understorey mosaic for birds and bats; dead wood elements for saproxylic insects and fungi; limited mowing frequency and increased cutting height for forest herbs. Forestry should enhance the recovery of mixed deciduous stands and avoid conifer plantations.