La zone némorale xérotherme sud-européenne

Abstract

The South-European nemoral region. - The objective of this paper is to present a checklist and a biogeographical arrangement of the main vegetation types on the whole submediterranean zone (and supramediterranean belt in the mountains) of the European southern nemoral region. This revision has been made possible by the recent publication of a number of synthetic vegetation maps. It is suggested that this cartographic information may facilitate the design of an ecological study of this zone, particularly of its parameters.     

Dispersal ecology of the endangered woodland lichen Lobaria pulmonaria in managed hemiboreal forest landscape

Changes in the forest management practices have strongly influenced the distribution of species inhabiting old-growth forests. The epiphytic woodland lichen Lobaria pulmonaria is frequently used as a model species to study the factors affecting the population biology of lichens. We sampled 252 L. pulmonaria individuals from 12 populations representing three woodland types differing in their ecological continuity and management intensity in Estonia. We used eight mycobiont-specific microsatellite loci to quantify genetic diversity among the populations. We calculated the Sørensen distance to estimate genetic dissimilarity among individuals within populations. We revealed that L. pulmonaria populations have significantly higher genetic diversity in old-growth forests than in managed forests and wooded meadows. We detected a significant woodland-type-specific pattern of genetic dissimilarity among neighbouring L. pulmonaria individuals, which suggests that in wooded meadows and managed forests dominating is vegetative reproduction. The vegetative dispersal distance between the host trees of L. pulmonaria was found to be only 15–30 m. Genetic dissimilarity among individuals was also dependent on tree species and trunk diameter. Lobaria pulmonaria populations in managed forests included less juveniles compared to old-growth forests and wooded meadows, indicating that forest management influences life stage structure within populations. We conclude that as intensive stand management reduces the genetic diversity of threatened species in woodland habitats, particular attention should be paid to the preservation of remnant populations in old-growth habitats. Within managed habitats, conservation management should target on maintenance of the stand’s structural diversity and availability of potential host trees.     

Mind the gap: how open were European primeval forests?

There are two competing hypotheses about the structure and dynamics of primeval forests in lowland Europe: the high-forest and the wood-pasture hypotheses, both of which influence current European forest conservation policies. In a recent study using pollen-analytical data from across lowland Europe, Mitchell provides support for rejecting the wood-pasture hypothesis. His study is important for future forest management planning and for showing how hypotheses about large herbivores as biotic factors can be tested using palaeoecological data.     

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.     

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.     

Role of domestication in shaping <Emphasis Type="Italic">Castanea sativa</Emphasis> genetic variation in Europe

The genetic structure of sweet chestnut (Castanea sativa Mill.) across Europe was assessed using 73 inter-simple sequence repeat markers to screen 1,768 individuals from 68 stands distributed across 29 sites in five European countries (Italy, France, Spain, Greece, and UK). At each site, trees were sampled from three distinct management types (domestication levels): naturalized stands, managed coppice, and grafted fruit orchards. In more than a third of the orchards, nonlocal genetic material (grafted clones) were evident, showing (as predicted) large differences from the other two domestication levels for most of the within-population genetic diversity parameters estimated. Randomly generated linkage disequilibrium analysis revealed weak though significant differences in two-locus allelic correlations between naturalized stands and coppice, suggesting that long-term management techniques may influence the genetic makeup of the populations. Multivariate analysis revealed the existence of five distinct gene pools across the study area; three were located in Greece, one on the northwestern coast of the Iberian peninsula and a large gene pool covering the rest of the Mediterranean basin. The implications of the results are discussed in relation to developing conservation strategies for chestnut genetic resources in Europe.     

The effect of coppice management on moth assemblages in an English woodland

Coppice woodlands in Britain may become the target of increased management due to the rise in demand for woodfuel. The biodiversity value of sweet chestnut (Castanea sativa) coppice and the effect of coppice management upon this has received limited study. Moths were the focus of this study in an actively coppiced sweet chestnut woodland in southern England. Coupes, with between one and 20 years of coppice regrowth, were systematically sampled for night flying micro- and macro-moths and coppice structure and ground vegetation were described. Using differences in moth assemblage, three stages of coppice development were distinguished: with one-four, five-nine and more than 10 years of coppice regrowth. Differences in moth assemblage related to habitat conditions within each coppice stage. The young coppice stage moth assemblage was characterised by species typically associated with open habitats; moths of the middle coppice stage assemblage fed on trees and were species typically associated with open woodland and scrub habitats; moths of the mature coppice stage assemblage were species typically associated with closed canopy woodland and contained specialist species whose larval food consists of material such as lichen and decaying leaves. All three coppice stages supported species of listed conservation status; the mature coppice stage contained a distinctive range of scarce and threatened species. The study showed that active coppicing promotes a change in moth assemblage but consequently will temporarily eliminate many species of mature stage coppice. Management which provides a range of coppice age classes within a woodland, appears key in promoting moth diversity.     

Maladaptive habitat selection of a migratory passerine bird in a human-modified landscape

In human-altered environments, organisms may preferentially settle in poor-quality habitats where fitness returns are lower relative to available higher-quality habitats. Such ecological trapping is due to a mismatch between the cues used during habitat selection and the habitat quality. Maladaptive settlement decisions may occur when organisms are time-constrained and have to rapidly evaluate habitat quality based on incomplete knowledge of the resources and conditions that will be available later in the season. During a three-year study, we examined settlement decision-making in the long-distance migratory, open-habitat bird, the Red-backed shrike (Lanius collurio), as a response to recent land-use changes. In Northwest Europe, the shrikes typically breed in open areas under a management regime of extensive farming. In recent decades, Spruce forests have been increasingly managed with large-size cutblocks in even-aged plantations, thereby producing early-successional vegetation areas that are also colonised by the species. Farmland and open areas in forests create mosaics of two different types of habitats that are now occupied by the shrikes. We examined redundant measures of habitat preference (order of settlement after migration and distribution of dominant individuals) and several reproductive performance parameters in both habitat types to investigate whether habitat preference is in line with habitat quality. Territorial males exhibited a clear preference for the recently created open areas in forests with higher-quality males settling in this habitat type earlier. Reproductive performance was, however, higher in farmland, with higher nest success, offspring quantity, and quality compared to open areas in forests. The results showed strong among-year consistency and we can therefore exclude a transient situation. This study demonstrates a case of maladaptive habitat selection in a farmland bird expanding its breeding range to human-created open habitats in plantations. We discuss the reasons that could explain this decision-making and the possible consequences for the population dynamics and persistence.     

Artificial nest predation in hedgerows and scrub forest in a human-dominated landscape of central Mexico

Hedgerows as well as other narrow corridors could be valuable habitats for birds in regions of intensive agriculture, however, it is still not clear how successful breeding birds are in different types of hedgerows as compared to birds nesting in their natural habitats. We used artificial nests to examine whether hedgerows were sinks (ecological traps) for birds by comparing rates of predation in two types of hedgerows with different vegetation structure (simple and complex), and in a tract of scrub forest in an agricultural landscape of central Mexico. We determined also the types of predators responsible for egg predation. Ground and elevated nests were baited with one Japanese quail Coturnix japonica egg and one plasticine egg and placed alternately along transects. Significantly, greater predation rates were found in scrub forest and complex hedgerows than in simple hedgerows. Higher predation rates in complex habitats seemed to reflect the higher number of predator types found there. The most important predator types were carnivores followed by rodents, birds, and humans. Carnivores and rodents mainly predated ground nests, whereas birds and humans predated elevated nests. Simple hedgerows in this landscape appeared to offer relatively safe nest sites in terms of predation pressure when compared to more complex habitats (complex hedgerows and scrub forest).     

Changes of traditional agrarian landscapes and their conservation implications: a case study of butterflies in Romania

Global biodiversity is decreasing as a result of human activities. In many parts of the world, this decrease is due to the destruction of natural habitats. The European perspective is different. Here, traditional agricultural landscapes developed into species-rich habitats. However, the European biodiversity heritage is strongly endangered. One of the countries where this biodiversity is best preserved is Romania. We analyse the possible changes in Romania's land-use patterns and their possible benefits and hazards with respect to biodiversity. As model group, we used butterflies, whose habitat requirements are well understood. We determined the ecological importance of different land-use types for the conservation of butterflies, underlining the special importance of Romania's semi-natural grasslands for nature conservation. We found that increasing modern agriculture and abandonment of less productive sites both affect biodiversity negatively — the former immediately and the latter after a lag phase of several years. These perspectives are discussed in the light of the integration of Romania into the European Union.     

Invasion success of alien plants: do habitat affinities in the native distribution range matter?

Aim To assess how habitat affinities in the native distribution range influence the invasion success of 282 central European neophytes (alien plants introduced after ad 1500). Location Czech Republic. Methods Classification trees were used to determine which native habitats donate the most alien species, the correspondence between habitats occupied by species in their native and invaded distribution ranges, and invasion success of species originating from different habitats. Results The species most likely to naturalize in Central Europe are those associated with thermophile woodland fringes in their native range (81%), cultivated areas of gardens and parks (75%) and broad‐leaved deciduous woodlands (72%). The largest proportions of invasive species recruit from those that occur on riverine terraces and eroded slopes, or grow in both deciduous woodland and riverine scrub. When the relative role of habitats in the native range is assessed as a determinant of the probability that a species will become invasive in concert with other factors (the species’ residence time, life history, region of origin), the direct effect of habitat is negligible. However, the effect of native habitats on patterns of invasions observed in central Europe is manifested by large differences in the numbers of species they supply to the invaded region. More than 50 neophytes were recruited from each of the following habitats: dry grasslands, ruderal habitats, deciduous woodland, inland cliffs, rock pavements and outcrops, and tall‐herb fringes and meadows. Main conclusions Casual species recruit from a wider range of habitats in their native range than they occupy in the invaded range; naturalized but not invasive species inhabit a comparable spectrum of habitats in both ranges, and successful invaders occupy a wider range of habitats in the invaded than in the native range. This supports the idea that the invasive phase of the process is associated with changes in biological features that allow for extension of the spectrum of habitats invaded.     

基于“源地-廊道”生态安全格局构建逻辑范式的建设用地减量化研究

建设用地急剧扩张导致土地利用覆被发生剧烈变化,物种栖息地丧失、破碎化及分离,严重威胁到社会经济及生态可持续发展。以快速城市化地区苏锡常为研究区域、地区优势物种白鹭生境为源地、"源地-廊道"生态安全格局构建逻辑范式为重要性评价依据,识别并筛选出对维护、控制目标物种生态过程具有重要意义的、可增强地区生境整体性及连通性的网络构成要素及其缓冲区,对缓冲区内建设用地进行减量化,提出一套建设用地减量化系统性方法。结果表明：（1）研究区内重要生态"源地"及"对应廊道"内建设用地需减量化10170.44 hm²,其中城镇用地5408.38 hm²,农村居民点4107.96 hm²,交通用地654.10 hm²。（2）依据生态安全格局构建逻辑范式,增补保护不同类型"源地"及"对应廊道",可在提升区域生态安全水平的同时,识别出新的需要减量化的建设用地。该方法也可作为区域内存量及增量建设用地空间布局调整优化的依据,具有较好的普适性。     

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.     

Clone identification and clonal structure of the European aspen (Populus tremula)

The European aspen (Populus tremula) is thought to reproduce mostly asexually. Thus aspen forms clones, in which several ramets belong to one genetically defined genet. We compared the clonal structure of aspen in old-growth and managed forests in southern and northeastern Finland. Clones were identified using morphological characters and nine microsatellite loci originally developed for Populus tremuloides. There were more clones identified by microsatellites than morphotypes both in old-growth and managed forest. The average size of the clones was only 2.3 ramets and most clones (70%) consisted of just one ramet. The size of the clones showed no difference between managed and old-growth forests or between northeastern and southern Finland. The small size of the clones suggests that most of them are relatively young. Therefore, sexual reproduction may be more common than previously thought. There was an aggregated spatial genetic structure as measured by Moran's I (0-10 m) and by co-ancestry (rho(ij), 0-20 m). Low level of co-ancestry can be explained by relatively unrestricted gene flow, the important role of disturbance in reproduction, and/or local selection.     

Implications of afforestation for bird communities: the importance of preceding land-use type

Afforestation of open habitats is one of the principal land-use changes underway in Europe and elsewhere in the world at present, and it can have a considerable impact on local biodiversity. The sustainable expansion of global forest plantations requires an understanding of the factors that determine the ecological impacts of afforestation. This study set out to determine the importance of preceding land-use type in determining the outcomes of afforestation for bird communities. Paired comparisons of 5-year-old exotic conifer plantations and matching non-forested sites were studied in areas of low (peatland), intermediate (wet grassland) and high (improved grassland) management intensity. Afforestation resulted in an overall increase in total bird density in all three habitat types. The effects of forest planting on bird conservation were found to be positively related to prior management intensity at the site. The density of bird species of conservation concern increased in response to the planting of intensively managed grassland sites, but decreased in response to afforestation of peatlands and of grasslands under intermediate management intensity. This study shows that plantation forests can, in some contexts, offer opportunities for bird conservation, and the findings highlight the trade-offs that are an integral part of land-use change. Therefore, where afforestation planning includes consideration of its impact on bird communities, planting should take place predominantly on sites of low biodiversity value, such as agriculturally improved grasslands. Furthermore, the preservation of sites of high conservation value within areas of afforestation would confer advantages on bird communities.     

Contrasting patterns in the invasions of European terrestrial and freshwater habitats by alien plants,insects and vertebrates

Aim To provide the first comparative overview on the current numbers of alien species that invade representative European terrestrial and freshwater habitats for a range of taxonomic groups. Location Europe. Methods Numbers of naturalized alien species of plants, insects, herptiles, birds and mammals occurring in 10 habitats defined according to the European Nature Information System (EUNIS) were obtained from 115 regional data sets. Only species introduced after ad 1500 were considered. Data were analysed by ANCOVA and regression trees to assess whether differences exist among taxonomic groups in terms of their habitat affinity, and whether the pattern of occurrence of alien species in European habitats interacts with macroecological factors such as insularity, latitude or area. Results The highest numbers of alien plant and insect species were found in human‐made, urban or cultivated habitats; if controlled for habitat area in the region, wetland and riparian habitats appeared to support relatively high numbers of alien plant species too. Invasions by vertebrates were more evenly distributed among habitats, with aquatic and riparian, woodland and cultivated land most invaded. Mires, bogs and fens, grassland, heathland and scrub were generally less invaded. Habitat and taxonomic group explained most variation in the proportions of alien species occurring in individual habitats related to the total number of alien species in a region, and the basic pattern determined by these factors was fine‐tuned by geographical variables, namely by the mainland–island contrast and latitude, and differed among taxonomic groups. Main conclusions There are two ecologically distinct groups of alien species (plants and insects versus vertebrates) with strikingly different habitat affinities. Invasions by these two contrasting groups are complementary in terms of habitat use, which makes an overall assessment of habitat invasions in Europe possible. Since numbers of naturalized species in habitats are correlated among taxa within these two groups, the data collected for one group of vertebrates, for example, could be used to estimate the habitat‐specific numbers of alien species for other vertebrate groups with reasonable precision, and the same holds true for insects and plants.     

Livestock management in Natura 2000: A case study in a Quercus pyrenaica neglected coppice forest

Due to the European Habitats Directive, Quercus pyrenaica habitat has been protected within the Natura 2000 network for biodiversity conservation. Many of these habitats have been widely managed as coppice forest with silvopastoral use throughout Spain. Currently, in many of these oak coppices fuelwood extraction is neglected and grazing is the main use. Management plans required in the Directive should consider all these circumstances and offer an integrated approach to ensure a favourable conservation status of these habitats. We propose a three phase methodology to assess the ability of the habitat to support a given livestock density annually and seasonally, and to evaluate some of the physical impacts of grazing animals on the physical structure and development of the habitat. We applied the methodology to a neglected coppice forest that supports livestock at different periods. Results show that there is a possible compatibility between annual livestock food requirements and pasture production. Current grazing periods seem to fit the seasonal pasture productivity. Negative impacts were assessed on oak shrubland stands due to intense browsing and on low polewood owing to the high density of broken trees. On more developed stands, positive livestock impact was due to the control of the regrowth of coppice shoots. Acorn absence is a result of the previous coppice management. This negative impact may endanger long-term viability of Q. pyrenaica in the study area. We propose a process of conversion into high forest through a sequence of thinning to start sexual regeneration. In this proposal, livestock have an important role to ensure biodiversity, control the regrowth of coppice shoots and thinning low polewood stands.     

Woodmanship

Summary

Woodmanship is the craft and technology of managing and adapting wild trees and natural woodland as a resource. It is widespread in the north temperate world (as yet there is curiously little evidence for the tropics) and goes back in time to the beginnings of settled civilisation. It mostly concerns either coppice-woods or wood-pasture. The latter integrates the management of trees and wild or domesticated livestock. Wood-pasture often takes the form of savanna, and may involve pollarding or shredding the trees to yield leaves on which to feed the animals. The study of woodmanship begins by knowing the behaviour of tree species and their different responses to felling, browsing, etc. It brings together evidence from a variety of sources: written documents, archaeological timber and wood, archaeological remains on the ground, pollen analysis, ancient living trees and surviving vegetation.