Bryophytes as indicators of ancient woodlands in Schleswig-Holstein (Northern Germany)

Ancient woodlands, with their long ecological continuity, frequently harbor a high number of typical, rare and threatened species, and are therefore of particular importance for nature conservation. To pinpoint these habitats, a common application is the use of plants as “ancient woodland indicators”. The occurrence of these particular species allows for evaluating the continuity of woodland cover in time. While lists of ancient woodland vascular plants have been derived for many regions, the identification and use of bryophytes as ancient woodland indicators has been widely neglected. This is a bit surprising because certain woodland bryophytes are very sensitive to varying environmental conditions or changes in land management. It therefore appeared promising to compile an ecologically grounded list of ancient woodland indicator bryophytes for practical use.In this study, we present a set of ancient woodland indicator bryophytes based on the analysis of datasets from the North German federal state of Schleswig-Holstein. To compile this list, we systematically evaluated the bryophyte distribution data from floristic surveys in relation to ancient woodland cover data from state-wide inventories. In this way, we were able to determine ancient woodland bryophytes using consistent and repeatable statistical methods.The presented list of 31 ancient woodland indicator bryophytes is ecologically sound and corresponds well with data from the sparse literature. We could distinguish two groups of ancient woodland indicator bryophytes. The first group is linked to base-rich, semi-natural deciduous woodlands with high soil and air humidity. The second group comprises acidophilic bryophytes that occur not only in acidic beech and oak woods, but also in acidic mixed or coniferous forests on ancient woodland sites. Apart from the ancient woodland indicator bryophytes, we could identify one group of recent woodland bryophytes and four groups of bryophytes that are more or less indifferent with respect to woodland continuity.Finally, we provide recommendations for the application of ancient woodland indicator bryophytes in nature conservation practice. Management suggestions for the conservation of the typical bryophyte diversity of ancient semi-natural woodlands are also given.     

Relations between the floristic composition of isolated young woods and their proximity to ancient woodland

Abstract. Recent studies indicate that, in the present-day agricultural landscape, the floristic composition of young woodland communities can be fully developed if the woods are situated adjacent to ancient woodlands. Four 70-yr-old deciduous woods in the Carpathian foothills were examined in relation to three adjacent ancient oak-hornbeam and oak-pine woodlands, which are the nearest source of woodland species diaspores. On the basis of data from 208 plots, the frequencies of various species groups in the field layer of the woods were analysed. The dependence of vegetation differentiation within the recent woods on (a) distance to the border with the ancient woodlands and (b) light intensity was examined by Partial Detrended Canonical Correspondence Analysis (DCCA). A significant relation between distance to ancient woodland and species composition was found for recent woods on rich brown soils. The vegetatively propagating species, myrmecochores and small autochores attained higher cover values near ancient woodland; endozoochores and anemochores were most abundant further away. Within recent, more open woods on poor podzolic and leached brown soils, colonisation is strongly inhibited by dense growth of Carex brizoides; here, vegetation regeneration is much slower than in woods on rich soils much further away from the source of diaspores.     

Colonization of secondary woodlands by Anemone nemorosa

Migration of the herb species Anemone nemorosa from older woodlands into adjacent, recently established deciduous woodlands on former arable land was studied. The wood anemone had colonized part of all studied recent woods, varying in age between 30 and 75 years. Cover of A. nemorosa in the recent woods decreased with increasing distance from the older woodland at all sites but one, indicating dispersal limitation during colonization. The advancing edge of most populations was characterized by negative logarithmic decrease in cover with establishment of isolated pioneer individuals and later gradual infill between pioneers. Migration rates were calculated by three methods. The mean migration rate of A. nemorosa based on observed maximum cover in the recent woods was 0.20 m year¹, 0.40 m year^-1 based on half maximum cover and 0.85 m year¹ based on the individual found farthest from the former woodland border. The calculated migration rates were consistently higher than the rate of possible rhizome growth. Seed dispersal and establishment is thus very important for colonization of new woodlands. Migration rates increased with tree canopy cover–especially cover of broad-leaved species with quickly decomposing litter–and with soil pH. Migration rates decreased with increasing grass cover in the field layer. These differences in migration rates may be due to increased micro-site availability for establishment at high canopy cover, low grass cover and high soil pH. Our results show that the wood anemone generally colonizes recent woods from nearby source populations. However, colonization proceeds relatively slowly and is limited by both seed dispersal and availability of suitable micro-sites. Gradients in abundance of A. nemorosa within secondary woods may be detectable for long periods of time and indicate the recent origin of a woodland.     

Factors influencing vegetation gradients across ancient‐recent woodland borderlines in southern Sweden

Abstract. We studied gradients in field layer vegetation across ecotone‐type borderlines between 12 ancient woodlands and adjacent secondary deciduous woodlands on former arable land. The aim of the study was to determine how distance from the borderline influences species distributions as compared with soil factors and degree of canopy closure. Correspondence Analysis showed that distance from the borderline is closely related to the first ordination axis at all study sites. Canonical Correspondence Analysis with variation partitioning revealed that distance from the borderline was the single most important factor in explaining vegetation variation. In general, the results suggest the following order of decreasing importance: Distance from the borderline < Soil reaction < Soil nitrogen < Soil moisture < Canopy cover. However, the sum of soil variables, as estimated by weighted averages of Ellenberg indicator values for moisture, reaction and nitrogen, accounted for as much as ca. 50–70% of the total variation explained by environmental variables. Important gradients in field layer vegetation are due to a decrease in typical woodland species and an increase in other species with increasing distance from the ancient woodland. The results suggest dispersal limitation of woodland species as an important determinant of secondary forest succession. However, the importance of distance to species distributions decreases with increasing stand age as most woodland species gradually colonize the recent woodlands. After 70 yr, ca. 50 % of the woodland species present at a site showed complete colonization within 50 m from the ancient woodland border.     

The macrolepidoptera of farm woodlands: determinants of diversity and community structure

The Farm Woodland Scheme, which provided incentives to convert agricultural land to timber production, contained an implicit assumption that farm woodlands produce important benefits for wildlife. The moth fauna of 18 farm woodlands in the Vale of York was surveyed between May and November 1991. The aims were twofold. The first was to determine if there were benefits for moth species diversity. The second was to ascertain whether concepts of island biogeography and the plant species richness of the woods were related to the moth species composition.Eleven families, 214 species and over 16 000 individuals of moths were recorded. Classification of the species presence/absence matrix indicated that small woods (less than 1ha) did not have characteristic woodland moth communities. Woods larger than 5ha were judged to be more valuable for the long-term conservation of woodland moth diversity. The best predictor of moth species richness was the herbaceous plant species richness within woodlands. Species richness of the family Geometridae was positively related to woodland area, as well as to woodland shape (compact shapes being preferable to elongated shapes). Characteristic woodland species are influenced by isolation (less isolated woods being richer in species). The implications of different powers of dispersal between moth families are discussed. Farm woodlands will be of more value for the conservation of the Macrolepidoptera if they are large, compact and incorporate remnants of existing woodland with extant herbaceous vegetation. These should be factors which are taken into consideration when providing incentives to establish and manage farm woodlands.     

The ground beetle fauna of ancient and recent woodlands in the lowlands of north-west Germany (Coleoptera, Carabidae)

In north-west Germany the ground beetle fauna was investigated using pitfall traps at 79 sampling sites in ancient woodlands and recent woodlands (with and without direct contact to old stands). Two woodland types were considered: The Quercion robori-petraea-woodlands (oak-beech-type) on mainly sandy soils and the Stellario-Carpinetum-woodlands (hornbeam-type) on mainly loamy soils. The number of recorded ground beetle species inhabiting exclusively or predominantly woodlands in the investigation area is significantly higher in ancient stands of both woodland types than in recent ones. No statistically substantiated relation between habitat size (both about 1800 and in 1990) and the number of characteristic woodland ground beetle species could be ascertained. Carabus glabratus and Abax parallelus show a distinct focus in ancient woodlands. Significantly more records of Carabus violaceus and Abax parallelepipedus are known from ancient woodlands than from recent ones. Twelve of the 16 ground beetle species, for which no difference in the colonisation of ancient and recent woodlands was ascertained, are macropterous. Half of the eight brachypterous woodland species is exclusively or predominantly found in ancient woodlands, suggesting that power of dispersal is an important factor which determines the species number in woodland fragments of different age.     

Woody plant encroachment reduces species richness of herb‐rich woodlands in southern Australia

Abstract Woody plants have been increasing in many woodland and savanna ecosystems owing to land use changes in recent decades. We examined the effects of encroachment by the indigenous shrub Leptospermum scoparium (Myrtaceae) on herb‐rich Eucalyptus camaldulensis woodlands in southern Australia. Species richness and compositional patterns were examined under the canopy of L. scoparium and in surrounding open areas to determine the species most susceptible to structural changes. Richness was significantly lower in areas of moderate to high L. scoparium cover (>15%), suggesting that a threshold shrub cover caused major change in this ecosystem. Shrubs were associated with a significant reduction in above‐ground biomass of the ground‐layer flora and a significant shift in community composition. The few species that were positively associated with high L. scoparium cover were also common in the woodland flora; no new species were recorded under the shrub canopy. Important environmental changes associated with L. scoparium cover were decreased light availability and increased litter cover, which were likely a consequence of encroachment. Leptospermum scoparium cover was also associated with greater surface soil moisture, which may be a consequence of increased shading under the shrub canopy or indicate favourable soil conditions for L. scoparium establishment. Reductions in species richness and abundance of the germinable seed bank were found in soil samples taken from under L. scoparium. With ongoing recruitment of L. scoparium and consequent increases in shrub cover, ground‐layer diversity in these species‐rich woodlands should continue to decline over time.     

Woodland restoration on agricultural land: long‐term impacts on soil quality

Woodland restoration is underway globally to counter the negative soil quality and ecological impacts of agricultural expansion and woodland fragmentation, and restore or enhance biodiversity, ecosystem functions and services. However, we lack information about the long‐term effects of woodland restoration on agricultural soils, particularly at temporal scales meaningful to woodland and soil development. This study utilized soil and earthworm sampling across a chronosequence of sites transitioning from “agricultural land” to “secondary woodland” (50–110 years) and “ancient woodland” (>400 years), with the goal of quantifying the effects of woodland restoration on agricultural land, on key soil quality parameters (soil bulk density, pH, carbon and nitrogen stocks, and earthworm abundance, biomass, species richness and diversity). Broad‐leaved woodland restoration led to significantly greater soil organic carbon (SOC) stocks compared to arable land, and young (50–60 years) secondary woodland increased earthworm species and functional diversity compared to both arable and pasture agricultural land. SOC stocks in secondary broad‐leaved woodlands (50–110 years) were comparable to those found in long‐term ancient woodlands (>400 years). Our findings show that broad‐leaved woodland restoration of agricultural land can lead to meaningful soil ecological improvement and gains in SOC within 50–110 years, and provide intel on how restoration activities may be best targeted to maximize soil quality and functions.     

“Ghosts of the past”: flightless saproxylic weevils (Coleoptera: Curculionidae) are relict species in ancient woodlands

Historic maps show that the Central European landscape was influenced by exploitive human land-use during the middle ages and in the following centuries. A mixture of ancient woodlands, which survived the period of woodland destruction, and recent woodlands, which were established after 1800, cover about 10% of the study area in NW Germany today. Weevils (Coleoptera: Curculionidae) of the subfamily Cryptorhynchinae with the genera Acalles, Kyklioacalles, Ruteria and of the subfamily Molytinae, tribe Acicnemidini with the genus Trachodes are all flightless and possibly influenced by landscape history. The aims of this investigation are (1) to examine the spatial distribution of flightless saproxylic weevils in ancient and recent woodlands in NW Germany and (2) to test the frequency of possible relict species in relation to historical and current woodland size. Based on a field study in 29 deciduous woodlands and species records in collections and literature, six flightless saproxylic weevils were found to be associated with ancient woodlands in NW Germany. None of these were recorded in any of the 14 recent woodlands studied. The frequency of these relict species is correlated with historical, but not with current, woodland size. Distribution maps for Lower Saxony and data on the phenology of the relict species are presented. These weevils are relict species of ancient woodland, because they were unable to colonise isolated woods that were established after 1800. All of them are dependent on dead or dying wood for larval development. The results show that ancient broadleaved woodlands with long-lasting habitat continuity are of high conservation value for invertebrate species such as saproxylic weevils.     

Ecological change in British broadleaved woodland since 1947

Over the last 60 years Britain's broadleaved woodlands have undergone a complex pattern of ecological change. The total extent has expanded from c . 676 000 ha in 1947 to c . 904 000 ha in 2002, but there has also been significant turnover, with losses of ancient woodland and a gain from new planting. Structural change has occurred due, in part, to change in management. In 1947 21% of the broadleaved resource was classed as coppice, 28% as scrub and only 51% of the area as high forest, compared with 97% high forest in 2002. This has been accompanied by changes in ground flora and the regeneration pattern of tree species, which will impact upon the character of the woodlands. Woods have also become more ecologically isolated because of the decline in semi-natural vegetation in the surrounding countryside, although the long-term impacts of this are poorly understood. Other factors driving ecological change are increased pollution, change in grazing pressures, climate change, alien species and game management. Overall woodland specialist species and those of open habitats tend to be doing less well than woodland generalists. Progress has been made in reversing some adverse impacts such as acid deposition, and action is being taken to reduce the impact of others such as over-grazing by deer. However, some drivers, notably climate change, will be more difficult to address. In the long term integrated management of woods as landscape components whilst meeting economic and societal needs will be required.     

The effects of urban or rural landscape context and distance from the edge on native woodland plant communities

The increasing rate of urban sprawl continues to fragment European landscapes threatening the persistence of native woodland plant communities. The dynamics of woodland edges depend on the characteristics of woodland patches and also on landscape context. Our aim was to assess the extent of edge influences on the understorey vegetation of small native woodlands in rural and urban landscapes. The study was carried out in two cities of north-western France. Ten comparable woodlands, each of about 1.5 ha, were surveyed; five were situated adjacent to crops and five adjacent to built-up land. Vascular plant species were recorded in 420 3 × 3 m plots placed at seven different distances from the edge (from 0 to about 45 m from the edge). Soil pH, light levels, level of disturbance and tree and shrub cover were also recorded. Plant species were first classified as non-indigenous or indigenous and then three groups of indigenous species were distinguished according to their affinity for forest habitat (forest specialists, forest generalists and non-forest species). We inferred certain ecological characteristics of understorey vegetation by using Ellenberg values. An inter-class correspondence analysis was carried out to detect patterns of variation in plant community composition. Linear mixed models were used to test the effects of adjacent land use, distance from the edge and their interactions on the species richness of the different groups and on the ecological characteristics of vegetation. Total species richness, richness of forest generalists and of non-forest species decreased from edge to interior in both urban and rural woodlands. The number of non-indigenous species depended mainly on urban–rural landscape context. Urban woodland edges were not as rich in forest specialists as rural edges. More surprisingly, the number of forest specialists was higher in rural edges than in rural interiors. Community composition was mainly affected by urban–rural context and to a lesser degree by the edge effect: the community composition of urban edges resembled that of urban interiors whereas in rural woodlands vegetation near edges (up to 10 m) strongly differed from interiors with a pool of species specific to edges. Urban woodland vegetation was more nitrophilous than rural vegetation in both edges and interiors. A major difference between urban and rural vegetation was the distribution of basiphilous species according to distance from the edge. Generally edge vegetation was more basiphilous than interior vegetation however the presence of basiphilous species fell off quickly with distance from the edge in rural woodlands (in the first 10–15 m) and more slowly (from 25 m onwards) in urban woodlands. This pattern was linked to variation in measured soil pH. As regards the conservation of flora in small native woodlands, it appeared that invasion of exotic and non-forest species was currently limited in both urban and rural landscape contexts but might pose problems in the future, especially in urban woodlands. Forest species were not negatively affected by the edge effect and indeed edges seemed to provide important habitats for this group. Hence conservationists should pay particular attention to the protection of edges in urban woodlands.     

Woodland birds in patchy landscapes: the evidence base for strategic networks

Habitat creation and management within wooded networks is a potentially effective strategy to reduce ecological isolation and the deleterious effects of fragmentation. However, questions remain over the relative advantages of different approaches, e.g. buffering patches vs. increasing connectivity. Potential effects of woodland fragmentation include reduction in regional woodland cover, reduced patch size, edge effects with loss of core habitat, and increased isolation with disruption of dispersal and metapopulation dynamics. We adopt an evidence-based approach to review how each of these affects woodland birds with an emphasis on studies from the UK and use this to identify management priorities for mitigation. There is evidence for both patch area and composition effects: larger woodlands support more woodland bird species, and woods located within sparsely wooded landscapes are less valuable to specialist woodland species. Bird assemblages show a nested pattern with respect to area, and thus species found in small woods also occur in large woods but not vice versa. However, small woods may be preferred by a few edge species, while small woods also have greater variability in bird species composition. Consideration of the metapopulation dynamics of specialist species with poor dispersal shows that creating or buffering large woodlands is more efficient than a greater total area of small fragments. Connectivity appears most useful for widespread generalist species with almost continuous populations. Woodland structure and quality are of overwhelming importance: as well as mature woodland, young growth, scrub and edges are also key components. There is an urgent need to examine the relationship between nest predation and landscape structure within UK woodlands.     

Vegetation differentiation and secondary succession on a limestone hill in southern Poland

Abstract. The vegetation of the SkoHzanka Reserve, situated on a limestone hill near Krakow, was examined over a 30 year period, using the Braun-Blanquet method. With the help of a numerical classification eight types of grassland communities were distinguished, belonging to the classes Sedo-Scleranthetea, Nardo-Callunetea and Festuco-Brometea, and 11 types of woodland communities, both remnants of ancient woodlands from the classes Querco-Fagetea and Vaccinio-Piceetea and more recent woods. In the case of grasslands the results of detrended correspondence analysis (DCA) indicated that the largest variation was associated with the soil type. Woods on sandy soils are very different from those on calcareous soils, and the younger recent woods are very different from the ancient woodlands. An analysis of historical cartographic material and aerial photographs shows that in the last few decades the area of recent woods has increased considerably at the expense of grasslands. Changes in species diversity in the woods during secondary succession and their successional convergence were examined and discussed. It was found that young and mature woods on sandy soils are more similar to each other than young and mature woods on calcareous soils. These differences are mainly related to the fact that many woodland herb species growing in rich, broadleaved ancient woods are unable to colonize recent woods since they have a limited colonization capacity.     

The nature of three ancient woodland soils in southern England

Abstract. Despite a wealth of published research on the nature of woodland soils, little is known about the nature of soils on sites that have supported woodland for many hundreds of years, namely ancient woodland. The properties and variability of soils in three ancient woods; one in the New Forest, Hampshire and two in Berkshire, were compared with those under recent woods. The acidity of ancient and recent woodland soils was high and remarkably similar. Only where cultivation of soils had preceded woodland establishment was soil acidity lower. The quantity of carbon in the soils studied was inversely related to soil acidity and the ancient woods had accumulated larger quantities of carbon than their recent counterparts. The quantities of Ca²⁺, Mg²⁺ and K⁺ were larger in the ancient woods except where prior cultivation had taken place. Total and organic phosphate contents of the ancient woodland soils were also consistently larger. The nature and pattern of soil variability in ancient woodland soils was quite distinct from that found in recent woods. Overall, the variability of soil acidity, carbon content and organic phosphate was larger in the ancient woodland soils but the pattern of variability differed between the soil properties. No clear association existed between the pattern of soil acidity and individual trees. At the surface of some of the woodland soils, however, carbon distribution appeared to be associated with individual trees. At depth in the ancient woodland soils, the association with the existing vegetation cover was not so clear. It is probable that the ancient woodland soils retained relict features of previous vegetation cover. Organic phosphate distribution was very strongly associated with the present vegetation cover. The pattern of distribution of organic phosphate appeared to be stronger than that of soil acidity and carbon content.     

Factors determining species richness of soil seed banks in lowland ancient woodlands

The demise of coppicing in UK ancient woodlands, combined with the planting of non-native, fast-growing conifers in the twentieth century, heightens the potential recharge value of ground flora seed banks. Soil cores from adjoining semi-natural and conifer-containing stands in four lowland ancient woods in central England were removed to establish seed bank species richness. During a fourteen-month germination trial soil from two depths yielded 6554 seedlings from 81 species, ten of which showed a strong affinity for ancient woodland conditions. Juncus effusus accounted for 80% of emergent seeds whilst 23 other species, including Lysimachia nummularia and Potentilla sterilis, were represented by only one individual. Species richness is described by a model that explains 40% of observed variance (P < 0.00001). The model has three significant variables: species richness increases as soil pH rises, and decreases with both depth and increasing time since the most recent planting/disturbance event. No difference was found in the density of seeds from species common to paired semi-natural and conifer-containing stands that were separated only by a woodland ride, suggesting prior management and environmental conditions have a greater influence on seed banks than current stand type. Sørensen similarity index values revealed poor congruence between above-ground vegetation and species in the seed bank. Taking pH measurements in conifer stands identified as younger in terms of planting/disturbance may help locate areas where greater numbers of species (including woodland specialists) are located. Caution is required, however, as these seed banks may also contain non-target, competitive species that may swamp the regeneration of woodland specialists.     

Lowland woodland structure and pattern and the distribution of arboreal,phytophagous arthropods

We investigated factors that limited the distribution of phytophagous species within a woodiand system in Midlothian, Scotland. A pattern analysis was conducted of phytophagous species on a total of 45 Fagus sylvatica within 15 woodlands. Species richness counted on collected leaves was tested against within-and between-wood variables. Variables used in a regression with arthropod data from Fagus were used to estimate the phytophage richness on Betula pendula and Quercus robur in the same woods. Convariance in the number of phytophages in sampled woods was found for Fagus over three years and for Fagus, Betula and Quercus in 1992. Association analysis was used to classify the woods into species rich or poor based on presence or absence matrices. The main factors that limit phytophages on Fagus (gaps along the woodland edge, depth and species richness of the field layer. density of leaf litter and the extent of contiguous woodland cover, when including hedgerows and lines of trees) affect phytophages of similar life history strategy on other tree species within the same woods. Eighty-six per cent of species were lost because certain life history stages were vulnerable to factors that prevail in woods of poor structure. The nature conservation value of woodlands could be assessed using the correlated vulnerability of particular phytophages across tree species under specific woodland conditions.     

Has Botanical Enhancement of Broad‐Leaved Plantations in Milton Keynes,United Kingdom,Resulted in More Woodland‐Like Insect Assemblages?

Two methods of invertebrate sampling were used to determine how the deliberate introduction of field layer vegetation to broad‐leaved plantations influenced the associated insect assemblages. Enclosed pitfall traps and tent‐like emergence traps were employed at (1) 25‐year‐old plantations where botanical enhancement had taken place a decade previously; (2) “nonenhanced” plantations of a similar age; and (3) local ancient woods, all in Milton Keynes, Buckinghamshire, United Kingdom. The aim was to discover whether enhancement had produced a community intermediate to nonenhanced plantations and naturally established woodland. Enhancement had not successfully created plantations botanically more woodland‐like and plantation types were not constant; however, plant species richness had increased. This was also true of the insect assemblages (Coleoptera, Diptera and Hemiptera), which showed more variation in composition, though woodland communities could still be distinguished. However, those insect species present in the naturally established woods did have a significantly greater frequency and abundance in the enhanced plantations compared to the nonenhanced plantations, especially so with “woodland edge” species. Mantel tests showed significant correlations between the plant species present and insect assemblages taken by both sampling types. Although insect species richness was not significantly higher in the enhanced plantations, this correlated strongly with plant species richness and a measure of vegetation volume. The overall findings suggest that the enhancement had created plantations only subtly more like the local woodlands, though the insect assemblages had responded. However, relative to the time scale of woodland development, it is still early days.     

Effects of urban land cover on the local species pool in Britain

Regression models were used to quantify the relationship between the amount of urban land and composition of local species pools. There was no evidence that urban land cover increases the richness of plant species, based on a survey of 785 2-km squares of which 157 had > 10% urban land cover. However, the number of alien plants is significantly higher in urban areas. Complete urbanization approximately doubles the proportion of alien species, and the proportion of aliens is twice as high in southern Britain as in the north. The flora of urban tetrads consists of ubiquitous native species and introduced species characteristic of waste ground, but woodland species are poorly represented. At the tetrad scale. enhanced dispersal by man is not the main factor for maintaining the urban flora; availability of urban habitats and high levels of disturbance are more important. The planned housing expansion to greenfield sites in Britain will increase the proportion of alien species, yet the majority of native species should persist in urban areas if existing woodland is preserved.     

Landscape effects on birds in urban woodlands: an analysis of 34 Swedish cities

Aim To compare bird abundances in woodlands along gradients from the city centre to the peri‐urban area. To evaluate the importance of the proportion of woodland within the city and in the peri‐urban landscape to forest bird communities breeding in urban woodlands. To test whether fragmentation effects on birds were linked to the type of peri‐urban matrix. Location A total of 34 Swedish cities with > 10,000 inhabitants in south‐central Sweden. The study area covered 105,000 km², in which 84% of the Swedish population of 9.1 million lives. Methods Repeated point count surveys were conducted in 2004 in a total of 474 woodlands. General linear models were used to test for possible differences in abundance along urban to peri‐urban gradients, and to regress bird abundances in local urban woodlands on: (1) total woodland cover in the city, (2) total woodland cover in the peri‐urban landscape, (3) the interaction between woodland cover in the city and in the peri‐urban area, (4) region, and (5) human density. Results More than 12,000 individuals of 100 forest bird species were recorded. Of the 34 most common species detected, 13 bird species had higher abundances in urban than in peri‐urban woodlands, and seven species showed the opposite trend. The bird community of urban woodlands was characterized by species associated with deciduous forests and tree nesters, whereas the bird community of peri‐urban woodlands was characterized by species associated with coniferous woodland and ground nesters. Twelve species were significantly linearly associated with the proportion of urban woodland and/or the proportion of peri‐urban woodland, and a further eight species were associated with the interaction between these two factors. Local breeding bird abundances of four species were significantly positively associated with the proportion of urban woodland only in farmland‐dominated landscapes. Main conclusions Fragmentation effects on some urban birds are linked to the type of peri‐urban matrix. In farmland landscapes, peri‐urban woodlands may have been too scarce to act as a source of bird immigrants to fragmented urban woodlands. To maintain populations of specialized forest birds within cities in landscapes dominated by agriculture, it is of paramount importance to conserve any remaining urban woodlands.     

Species richness of small woodlands on the western Carpathian foothills

Sixty-six small woodland islands (0.008–2.16 ha) isolated in the agricultural landscape of the Wierzbanówka valley (western Carpathian foothills) were investigated. Analysis of historical cartographic material showed that 60 woodland islands are most probably the remnants of ancient woodlands. Among them both woodlands associated with more than one type of soil, and those with homogeneous soil conditions, were identified; the latter were divided into two groups differing in the period of isolation and anthropogenic pressure. These three groups of woodlands were analysed statistically using 12 independent variables (area, shape, distance to the nearest large woodland, mean distance to the five nearest woodland islands, cover of tree-and shrub layer, and others) and 20 dependent variables (number of all species, number of species of different growth forms, mode of dispersal, and phytosociological characters). The number of species was found to be related to habitat diversity, area, shape, isolation, and cover of tree and shrub layers. Compared with the more recently isolated and less disturbed woodlands, those isolated for longer periods and more anthropogenically disturbed were found to have fewer species, including fewer harbaceous species, woodland species, species of the Querco-Fagetea class, and also tree and shrub dyszoochores, as well as herbaceous endozoochores and myrmecochores. On the other hand, significantly more non-woodland species, species of the Molinio-Arrhenatheretea class, and tree anemochores were present here.