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.     

Biodiversity and ecology of the invertebrates of Atlantic Oakwoods

Summary

Woodland in general supports the richest invertebrate fauna of any habitat in Britain. The number of species depends on the structure of the wood and the age of the trees within it. The greatest biodiversity is associated with oaks although the difference between the communities found on the two oak species is not clearly understood, the Sessile Oak woods have been less well studied than their lowland equivalents, particularly so in Scotland. A number of minor habitats within the wood are important for many invertebrates and the presence of many of these may be due to past management. Every part of the oak tree, the leaves, flowers, acorns, bark and wood, supports invertebrates, herbivores, predators and parasites, and the fallen leaves and deadwood provides habitat for a complex of detritus feeders. Within the woodland the other trees, the fungi and the ground flora all have characteristic woodland invertebrates. Only a few studies, on beetles and spiders, have looked at which invertebrates are particularly associated with acidic oakwoods and there have been no specific studies of Atlantic oakwoods. It is known that there are some rare and local invertebrate species associated with western woods on ancient sites, including some recently discovered suggesting that the biodiversity of these sites and their management for invertebrates might be more important than currently thought.     

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 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.     

Restoring planted ancient woodland sites — Assessment,silviculture and monitoring

Summary

This paper deals with three aspects of the process of restoring planted ancient woodland sites(PAWS) to semi-natural conditions. Firstly, we describe a baseline assessment of botanical interest within a PAWS. This survey has been undertaken to determine the impact of clearfelling, particularly on lower plants and the subsequent colonisation of ground vegetation into areas currently dominated by needle litter. Secondly, we discuss some of the main considerations when undertaking restoration through alternative silvicultural systems to clearfell. Finally, we describe the main requirement for successful site monitoring for management purposes.

The last decade has seen considerable attention given to the benefits of restoring plantations on ancient woodland sites (PAWS) to semi-natural conditions. The survival of species and communities associated with ancient woodland through the process of conversion will be a critical measure of success for restoration practice. It is generally assumed that a gradual approach will improve the chances of such success.

Glencripesdale National Nature Reserve (NNR) is a heterogeneous area of semi-natural open ground, ancient woodland and Sitka spruce plantation. The plantation blocks contain occasional discrete elements of semi-natural vegetation, including some veteran trees. Because of difficult access and poor stability, silvicultural options are limited and clear felling is the only practical option. We describe a simple method of monitoring changes to cryptogamic communities and ground layer vegetation prior to and following clearfelling of the plantation matrix. Baseline data are presented.

In more stable and accessible stands, there are a number of alternative silvicultural approaches to consider when gradually restoring a PAWS. This paper addresses the question of how to secure ancient woodland remnants. We outline some of the initial silvicultural considerations such as stability, thinning/felling pattern and light requirements of native tree species. An approach to site monitoring is presented to allow managers to assess whether conditions are improving or declining and whether they are delivering objectives.     

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.     

The potential contribution of conifer plantations to the UK Biodiversity Action Plan

Summary

The detrimental effects of conifer plantations on open ground habitats have been well catalogued and discussed, but the potential contribution of planted forests to the conservation of woodland biodiversity has not been quantified to the same extent. This quantification is needed urgently to help forest managers fulfil commitments to biodiversity enhancement as outlined in the UK Biodiversity Action Plan, the UK Forestry Standard and the UK Woodland Assurance Scheme (UKWAS). Results are presented from a five-year programme of research aimed at obtaining baseline information on biodiversity in planted forests and evaluating the contribution of planted forests to the conservation of native flora and fauna. Fifty-two plots were surveyed in total, covering a range of different tree crops (Scots pine Pinus sylvestris L., Sitka spruce Picea sitchensis (Bong.) Carr., Norway spruce Picea abies L. and Corsican pine Pinus nigra var maritima (Aitón) Melville) and stand ages (pre-thicket, mid-rotation, mature and over-mature) in three contrasting bioclimatic zones (upland, foothills and lowlands) throughout Britain. Additional plots were established in semi-natural woodland to allow comparisons between the biodiversity of plantations and native stands. Over 2000 species were recorded in total, including 45 Red Data Book species. Planted stands had similar or richer fungal and invertebrate communities to those of the native stands but poorer lichen and vascular plant communities. The latter were strongly affected by shading, dense, mid-rotation Sitka spruce stands having the lowest species counts. In contrast, these stands had a high diversity of mycorrhizal fungi, including a number of rare and threatened species normally associated with native pine wood. Bryophyte species-richness was related more to climate than woodland type, with the wetter upland spruce and native oak stands having the most diverse communities. Compared to the younger planted stands, over-mature planted stands had a higher proportion of species characteristic of semi-natural woodland stands. This related to greater structural diversity and higher deadwood volumes in the over-mature stands. It is concluded that conifer plantations make a positive contribution to biodiversity conservation in the UK and hence to the UK Biodiversity Action Plan. No single stand or crop type provides ‘optimal’ conditions for biodiversity, but the habitat value of plantations could be enhanced by increasing the area managed under alternative systems to clear-felling, such as ‘continuous cover’ and/or non-intervention natural reserves.     

The management of long term set-aside for nature conservation

Summary

Long term set-aside offers an opportunity to create new grasslands which may be visually appealing and valuable for wildlife. These grasslands, whilst not equalling the nature conservation value of old semi-natural grasslands, may have higher value for wildlife than the arable crops and grass leys which they replace. In this paper we compare the nature conservation value of vegetation established by either natural regeneration or by sowing a seed-mixture based on MG5 Cynosurus cristatus — Centaurea nigra grassland on set-aside at two sites in Scotland. The initial results suggest that sown vegetation is generally more botanically diverse than natural regeneration. Sheep grazing resulted in reduced botanical diversity in the sown plots and early cutting, with cuttings removed and aftermath grazing by cattle, resulted in the highest diversity estimates. However, some effects of management treatment were site specific.     

Pollen analysis and planted ancient woodland restoration strategies: a case study from the Wentwood, southeast Wales, UK

This paper considers the contribution of pollen analysis to conservation strategies aimed at restoring planted ancient woodland. Pollen and charcoal data are presented from organic deposits located adjacent to the Wentwood, a large planted ancient woodland in southeast Wales. Knowledge of the ecosystems preceding conifer planting can assist in restoring ancient woodlands by placing fragmented surviving ancient woodland habitats in a broader ecological, historical and cultural context. These habitats derive largely from secondary woodland that regenerated in the 3rd–5th centuries a.d. following large-scale clearance of Quercus-Corylus woodland during the Romano-British period. Woodland regeneration favoured Fraxinus and Betula. Wood pasture and common land dominated the Wentwood during the medieval period until the enclosures of the 17th century. Surviving ancient woodland habitats contain an important Fagus component that probably reflects an earlier phase of planting preceding conifer planting in the 1880s. It is recommended that restoration measures should not aim to recreate static landscapes or woodland that existed under natural conditions. Very few habitats within the Wentwood can be considered wholly natural because of the long history of human impact. In these circumstances, restoration should focus on restoring those elements of the cultural landscape that are of most benefit to a range of flora and fauna, whilst taking into account factors that present significant issues for future conservation management, such as the adverse effects from projected climate change.     

Trees,Woodland and Woodmanship in Early Mediaeval Ireland

Summary

The evidence for woodmanship in mediaeval Ireland is discussed. Palynological evidence dispels the continuing myth that there were vast mediaeval woods. Nonetheless, place name and documentary sources indicate discrete areas of woodland. At these places underwood was probably managed, livestock sheltered and fattened, berries and nuts gathered and timber felled. A range of historical texts and archaeological excavations support these claims. Woodland was an important aspect of the agricultural landscape and economy.     

Population changes of some bird species in Scottish and Welsh Atlantic Oakwoods between the 1980s and 2003/4 using data from the repeat Woodland Bird Survey

Summary

Atlantic Oakwoods support a unique breeding avifaunal community, characterised by redstarts, wood warblers, pied flycatchers and tree pipits. We have used a sample of data from the national Repeat Woodland Bird Survey to examine population changes in these four species within Atlantic Oakwoods over the last two decades. We used data from 29 woods in Argyll in western Scotland surveyed in 1985, and from 27 woods in Gwynedd in northwest Wales surveyed in 1983 and compare them with data from repeat surveys carried out in 2003 and 2004. In this paper, we report the population changes for these species between these two periods and examine whether changes differed between Scottish and Welsh sites.     

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.     

Distribution and extent of unimproved lowland grassland National Vegetation Classification (NVC) types in Scotland

Summary

National Vegetation Classification (NVC) data from surveys of Scottish unimproved lowland grasslands were reviewed. The surveys dated from 1980 to 2000. The mapped area of each lowland grassland NVC type was measured or, where vegetation maps were not of sufficient quality, estimated. These measurements and estimates were summed to give the total recorded area of each lowland grassland NVC type. An estimate of the total area likely to occur throughout Scotland was made for the more thoroughly recorded lowland grassland NVC types.

The data from this review has been used to create Scottish Natural Heritage's Lowland Grassland Database, containing details of all lowland grassland sites recorded by the grassland surveys, their grid reference, conservation status, site area, surveyor, date of survey, NVC lowland grassland types present and their areas. Council Area, old local authority District and SNH Area are also included.

A total of 8700 ha of unimproved lowland grassland NVC types (excluding coastal grasslands) has been recorded in Scotland. This habitat is semi-natural and of high conservation value.

The total area of unimproved lowland grassland in Scotland is estimated to be in the order of 30,000 ha.     

Factors influencing patch occupancy and within‐patch habitat use in an apparently stable population of Willow Tits Poecile montanus kleinschmidti in Britain

Capsule Birds selected younger woodland patches with fewer conifers and higher soil moisture content; within these patches, they selected areas with higher vegetation cover at 2–4 m, and less bracken.

Aims For an apparently stable population, to investigate habitat differences between occupied and unoccupied woodland patches and habitat selection within occupied woodlands.

Methods We surveyed woodland patches for breeding Willow Tits in April 2006 in an area with an apparently stable population. Habitat was measured at points spread throughout each woodland patch and at points where Willow Tits were located. We compared habitat between occupied and unoccupied sites. In addition, within occupied sites, we examined habitat differences between Willow Tit locations and the points spread throughout the wood.

Results We surveyed 65 woodland patches, of which 29 were occupied. Willow Tits were more likely to be found in deciduous woodland that was younger and had higher levels of soil moisture. Probability of occupancy fell from 60% for woods aged 20–25 years to only around 15–30% for woods aged over 80 years. Within occupied woods, Willow Tits were more likely to be found in areas with more vegetation cover at the 2–4 m level and in areas with less bracken.

Conclusions Habitat management for Willow Tits should involve provision of young woodland patches with moist soils. Changes to the age structure and/or the soil moisture content of woods may be implicated in the dramatic decline of Willow Tits in Britain.     

The Modification of Ancient Woodland Ground Flora at Arable Edges

The last five decades have witnessed an intensification of agriculture with a marked increase in the use of agrochemicals. This study investigated whether arable edges affect ancient woods and the extent of any modifications to the ground flora. Plant species were recorded from a total of 84 transects into the centre of the woods, of which 59 were taken from the arable–woodland interface and 25 were taken from non-arable land or woodland rides. Ellenberg indicator and CSR values were used to infer fertility and features of the environment. Results suggested that, after adjusting for light and moisture conditions, elevated nitrogen levels might occur in ancient woodlands to a distance of at least 100 m from arable farmland. Furthermore, the transects from the arable fields had a substantially altered herb-layer with respect to species composition, notably a greater percentage cover of ruderal and nitrophilous species, and a lower cover of stress tolerator species. Smaller woods contained a greater cover of competitor and nitrophilous species and a lower cover of stress tolerators and ancient woodland indicator species. We conclude that farming may be unwittingly modifying the ground flora of ancient woodlands adjacent to arable land and that small and irregularly shaped woods are most at risk. It is recommended that farmers apply agrochemicals in such a way that drift is minimised, and that large, valuable ancient woods be protected by a substantial buffer zone.     

Recreational Use of the Countryside: No Evidence that High Nature Value Enhances a Key Ecosystem Service

In Western Europe, recreational amenity is presented as an important cultural ecosystem service that, along with other values, helps justify policies to conserve biodiversity. However, whether recreational use by the public is enhanced at protected areas designated for nature conservation is unknown. This is the first study to model outdoor recreation at a national scale, examining habitat preferences with statutory designation (Site of Special Scientific Interest) as an indicator of nature conservation importance. Models were based on a massive, three year national household survey providing spatially-referenced recreational visits to the natural environment. Site characteristics including land cover were compared between these observed visit sites (n = 31,502) and randomly chosen control sites (n = 63,000). Recreationists preferred areas of coast, freshwater, broadleaved woodland and higher densities of footpaths and avoided arable, coniferous woodland and lowland heath. Although conservation designation offers similar or greater public access than undesignated areas of the same habitat, statutory designation decreased the probability of visitation to coastal and freshwater sites and gave no effect for broadleaved woodland. Thus general recreational use by the public did not represent an important ecosystem service of protected high-nature-value areas, so that intrinsic and existence values remain as the primary justifications for conservation of high nature value areas. Management of ‘green infrastructure’ sites of lower conservation value that offer desirable habitats and enhanced provision of footpaths, could mitigate recreational impacts on nearby valuable conservation areas.     

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.     

Woodland area,species turnover and the conservation of bird assemblages in lowland England

A study over 4 years into the number of breeding bird species and species turnover (extinctions and colonisations) in relation to area was conducted in 35 woodlands, set in an intensively farmed landscape, in north-east Essex, UK. A total of 46 species was recorded. The number of species breeding increased with woodland area; the slope of the species–area relationship did not differ between years. Habitat diversity was the only other measured variable to influence species richness. Absolute species turnover was independent of woodland area but relative turnover declined with increase in woodland area. The numbers of territories of nine species were determined. For four summer visitors the number of woods occupied increased as the overall populations increased but, for the other species, changes in overall population size led to changes in numbers in occupied woods. Chaffinch Fringilla coelebs and Song Thrush Turdus philomelos were more associated with woodland edges, Nightingale Luscinia megarhynchos, Garden Warbler Sylvia borin, Chiffchaff Phylloscopus collybita and Willow Warbler P. trochilus with interiors. Several species showed an inverse relationship between population density and woodland area. Collections of small woods hold similar species richness to single large woods. While the acquisition of large woods for conservation purposes should be a priority, the extension of smaller woods to a size of about 10 ha would be highly beneficial to both the species richness and population stability of regional woodland bird assemblages.     

The effect of woodland area on avian community composition in a fragmented southern UK landscape and associated management recommendations

ABSTRACT

Capsule: Smaller woodlands not only support fewer species but also show different avian community composition due to loss of woodland interior and an increase in edge habitat.

Aims: To use observed community composition changes, rather than traditional total species richness-area relationships, to make area-specific management recommendations for optimizing woodland habitat for avian communities in fragmented landscapes.

Methods: 17 woodlands were selected in Oxfordshire, UK, with areas between 0.2 and 120 ha. Three dawn area searches were conducted in each woodland between 1st April and 28th May 2016 to record encounter rates for each species. The impact of internal habitat variation on woodland comparability was assessed using habitat surveys.

Results: Woodlands with area less than 3.6 ha showed a significant positive relationship between total avian species richness and woodland area. Woodlands with area over 3.6 ha were all consistent with a mean (± se) total richness of 25.4?±?0.6 species, however the number of woodland specialists continued to increase with woodland area. Woodland generalists dominated the total encounter rate across the area range, however the fractional contribution of woodland specialists showed a significant positive correlation with woodland area, while the fractional contribution of non-woodland species significantly decreased. Non-woodland species numbers peaked in mid-sized woodlands with enhanced habitat heterogeneity.

Conclusions: Community composition analysis enabled more targeted recommendations than total species richness analysis, specifically: large woodlands (over 25?ha) in southern UK should focus conservation efforts on providing the specific internal habitats required by woodland specialists; medium-sized woodlands (between approximately 4 and 25?ha) should focus on promoting internal habitat variety, which can benefit both woodland species and non-woodland species of conservation concern in the surrounding landscape; small woodlands (under 4?ha) should focus on providing nesting opportunities for non-woodland species and on improving connectivity to maximize habitat for woodland generalists and facilitate movement of woodland specialists.