Ground beetle diversity in ancient woodland remnants in north-western Germany (Coleoptera, Carabidae)

We investigated ground beetle communities (Coleoptera, Carabidae) in ancient woodland remnants in north-western Lower Saxony, Germany. A total of 90 pitfall traps was exposed in a stratified design in 10 stands of mature oak–beech and oak–hornbeam forests in the year 2003. Overall, 47 species (10,676 individuals) were recorded. Among these were the two relict species Carabus glabratus and Abax parallelus, and 14 further eurytopic forest species. Eleven species exhibited a high frequency and were found in all of the ten stands. Multiple linear regressions showed several significant relationships at two scales for species richness of different groups of carabids and for several of the measured environmental factors. Forest area, litter depth, amount of dead wood, distance to forest edge, and soil moisture were found to be key factors determining species richness. Furthermore, recent disturbance by logging reduced the number of forest species. According to direct gradient analyses soil moisture and litter depth have greatest influence on species communities of both, forest species and widespread species. Habitat suitability models for the two recorded relict species, A. paralellus and C. glabratus, were developed using logistic regression. The presence of A. parallelus in the mature ancient woodland remnants depends mainly on higher values of soil moisture, whereas for C. glabratus none of the measured environmental variables appeared to be key factors. Implications for the conservation of carabid assemblages in mature ancient woodlands include the advice to spread out logging over long periods of time and over various woodlands in order to keep the stand disturbance at a long-term low level. Variation in logging practices may help to conserve diverse structures. Afforestation with non-native tree species should be avoided in the managed ancient woodlands. Finally, especially the preservation of a high soil moisture seems to be important to conserve typical carabid communities.     

Plant species as indicators of ancient woodland in northwestern Germany

Abstract. 315 isolated semi-natural and natural stands in NW Germany were investigated floristically, 285 stands of ancient woodlands and 30 recent ones. In the study area nearly all semi-natural and natural ancient woodlands are found on mesotrophic or eutrophic sites and can be assigned to the Fagetalia. 54 selected herbaceous and five woody Fagetalia-species have been tested in their association to ancient woodlands. 21 of the 59 selected plant species show a highly significant association to ancient woodlands and seven species show a weak significance. 31 plant species are not significantly correlated with ancient woodland sites, but 25 of them have a low frequency. The result shows that investigations of the historical ecology can help to understand floristic composition of present-day woodlands. Restriction of many woodland species to ancient woodlands, especially rare species, emphasizes the importance of woodlands with a long continuous history for the preservation of endangered species. Irrespective of ecological conditions, the restriction of plant species to ancient woodlands seems to be mainly caused by their low ability to colonize recent woodlands, especially isolated stands. Differences in the association to ancient woodlands between European countries are assumed to be a function of both time and degree of isolation of woodlands, of the dispersal mode of the plant species and of the availability of suitable habitats.     

Historical ecology meets conservation and evolutionary genetics: a secondary contact zone between Carabus violaceus (Coleoptera, Carabidae) populations inhabiting ancient and recent woodlands in north-western Germany

Only very few cases have documented that an increase in connectivity after a period of fragmentation in ecological time has had an effect on the distribution, genetic structure and morphology of stenotopic species. In this study we present an example of clinal variability in a woodland ground beetle as a result of changes in the connectivity of a landscape during the last two centuries. The study area hosts both the nominate form Carabus violaceus s. str. and the subspecies Carabus violaceus purpurascens, which is ranked as a distinct species by some authors. We studied 12 Carabus violaceus populations from a 30 km transect of ancient and recent forests in north-western Germany. We analyzed three polymorphic enzyme loci, classified the elytron sculpture and measured the shape of the aedeagus tip of the specimens. Carabus violaceus showed secondary gradients both in allozyme markers and morphometric characters in our study area. A genetic differentiation of 16% between the populations is high but lies within the range of intraspecific variability in habitat specialists of the genus Carabus. Populations had no significant deficit of heterozygotes. We found many hybrid populations in terms of morphological properties. This study highlights the conservation value of ancient woodland and the consequences of landscape connectivity and defragmentation on the genetic setting of a ground beetle. Moreover, it shows that differences in the external shape of male genitalia do not prevent gene flow within the genus Carabus. Thus, the establishment of species status should not exclusively be based on this property.     

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.     

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.     

Explaining the saproxylic beetle diversity of a protected Mediterranean area

Saproxylic beetle diversity is high at the Cabañeros National Park (central Spain), where woodland habitats exhibit remarkable heterogeneity. Our aim was to explain the diversity of saproxylic beetles, focusing on species turnover among mature woodland types. We surveyed five woodland types that represented the heterogeneity of the park’s woodland habitats. Beetles were collected using window traps over a period of 20 months. The Jaccard Similarity Index was used as indirect value of beta diversity among woodlands and to test the relation between species turnover and geographical distance. We also identified the contribution of species turnover to landscape diversity by using a partitioning model. Moreover, the presence of mixed woodlands (more than one tree species) allowed us to attempt to valorise the effect of tree species (coupled with their historical management) on species turnover among woodlands. Finally, we looked for different saproxylic beetle preferences for habitat and tree species using an indicator value method. We found that saproxylic beetle species composition varied significantly among the studied woodlands. The variation in species turnover was independent from the distance among woodlands, which suggested that beetle dispersal abilities could not explain this high turnover. Tree species within woodlands were a key factor that increased diversity turnover in woodlands and, consequently, the diversity of the park. Moreover, we found saproxylic beetle species that had different habitat and tree species preferences. We conclude that woodland heterogeneity (highly affected by woodland composition) seems to be the driving force for saproxylic beetle diversity in this protected area.     

Simulating the impacts of southern pine beetle and fire on the dynamics of xerophytic pine landscapes in the southern Appalachians

Question: Can fire be used to maintain Yellow pine (Pinus subgenus Diploxylon) stands disturbed by periodic outbreaks of southern pine beetle? Location: Southern Appalachian Mountains, USA. Methods: We used LANDIS to model vegetation disturbance and succession on four grids representative of xeric landscapes in the southern Appalachians. Forest dynamics of each landscape were simulated under three disturbance scenarios: southern pine beetle, fire, and southern pine beetle and fire, as well as a no disturbance scenario. We compared trends in the abundance of pine and hardwood functional types as well as individual species. Results: Yellow pine abundance and open woodland conditions were best maintained by a combination of fire and southern pine beetle disturbance on both low elevation sites as well as mid‐elevation ridges & peaks. On mid‐elevation SE‐W facing slopes, pine woodlands were best maintained by fire alone. Conclusions: Our simulations suggest that fire can help maintain open pine woodlands in stands affected by southern pine beetle outbreaks.     

“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.     

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.     

Identification of one X‐linked and five autosomal microsatellite loci in Carabus violaceus (Coleoptera,Carabidae) and their applicability to related taxa

We characterize six microsatellite loci from the ground beetle Carabus violaceus. The polymorphism of the loci was assessed in 46 individuals from one population. The number of alleles ranges between three and eight, and the observed heterozygosities between 0.46 and 0.76. All males are homozygous for locus CVI08106CMPG, whereas most females are heterozygous. This suggests that this locus is situated on the X chromosome. Tests in eight different species of ground beetles showed that four loci could be amplified in C. coriaceus and one each in C. glabratus, C. nemoralis, C. problematicus and Abax parallelepipedus.     

Orchard and riparian habitats enhance ground dwelling beetle diversity in Mediterranean agro-forestry systems

The relative contribution of mixed orchard and riparian vegetation patches to local and regional diversity of Mediterranean landscapes dominated by cork-oak woodlands was tested in 2006–2007 using ground, rove and darkling beetles (Coleoptera: Carabidae, Staphylinidae, Tenebrionidae). Mixed orchard and riparian gallery habitats recorded higher values of abundance and species richness for overall beetle species, although most darkling beetle species were associated with the sclerophyllous cork-oak woodlands. Ground and rove beetle community structure changed from the orchard and riparian habitats to samples placed 100 m away into the surrounding cork-oak woodland, i.e., non-cork-oak patches enhanced beta diversity within the landscape mosaic. Analysis of ground beetle traits concerning moisture preferences revealed a higher proportion of hygrophilous species in mixed orchard and riparian gallery habitats while xerophilous species were dominant within the cork-oak woodlands. The results of this study suggested that land-use management promoting the maintenance of habitat heterogeneity enhances biodiversity conservation of important hygrophilous and xerophilous species, and subsequently the sustainable use of Mediterranean agro-forest mosaics.     

Rural–urban gradients and the population genetic structure of woodland ground beetles

Genetic diversity and differentiation of two carabid beetle species were examined in woodlands along rural–urban gradients in two cities (Brussels, Belgium, and Birmingham, UK), based on allozymes, studied in more than 1000 beetles. Compared to Abax ater, Pterostichus madidus showed higher levels of genetic diversity but lower genetic differentiation, probably because of its ability to survive in non-forest habitats. Higher genetic diversity in both species was observed in Brussels as compared to Birmingham. However, genetic differentiation among sites was higher in Birmingham corresponding to the more extreme degree of fragmentation and isolation between the Birmingham woodlands. The isolation-by-distance model did not explain genetic differentiation among sites within the two regions. Gene diversity in P. madidus Birmingham populations was higher in smaller urban forests with a small perimeter. A similar absence of genetic erosion in smaller and more highly modified (urban) populations was also obtained for A. ater populations from Brussels, with a higher genetic diversity in sites closer to woodland edges. This unexpected result is hypothesised to be the result of an adaptive increase of genetic diversity in more heterogeneous landscapes, closer to woodland edges and in smaller and more perturbed forests.     

Changes in the ground flora in Wytham Woods,southern England from 1974 to 1991 – implications for nature conservation

Abstract. The ground flora (vascular plants < 2 m high, excluding trees and shrubs) was recorded in 1974 and 1991 from 163 permanent 10 m x 10 m plots arranged on a systematic grid across Wytham Woods, near Oxford (UK). The Woods cover about 320 ha, are predominantly deciduous, but of varying ages and management types. The total number of species found (173, 167 respectively), the mean richness per plot (16.7, 17.2) and the breakdown of the species list between different species types (ancient woodland indicators, other woodland species, non‐woodland species) showed no significant differences between 1974 and 1991, but mean ground cover declined from 80% to 64%. Ancient woodland indicators as a group showed less change between the years than species associated with the open glades and grassland patches in the wood. Some species increased in frequency across the woodland including Arum maculatum, Brachypodium sylvaticum, Deschampsia cespitosa and Poa trivialis; while others such as Ajugareptans, Angelica sylvestris and Circaea lutetiana declined. Mean cover of Rubus fruticosus per plot declined from 35% to 6% and of Mercurialis perennis from 32 to 24%. More species were lost from plantations than from semi‐natural stands. The results are based on only two times, so inferences on possible causes must be drawn with caution. However the results are consistent with the effects of (1) the changing conditions associated with stand growth, particularly in the plantations, and (2) an increase in browsing/grazing by deer. No evidence was found for an effect of changing soil nitrogen levels on the vegetation. Changes in the ground flora as well as the woody layers, and in managed as well as unmanaged stands should be monitored, if nature conservation objectives are to be met.     

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.     

The number and distribution of vascular plant species in island forest communities in the northern part of the West Carpathian foothills

Phytosociological records were made in 80 stands lying in 50 small woodland islands in the Wierzbanówka valley. The types of communities were distinguished on the basis of numerical classification of records. These were mainly fragments of theTilio-Carpinetum association. For all the stands and for two groups of stands connected with various soils and localized mainly in the eastern or western part of the valley the statistical relationships between ten independent variables were examined (area of record, area of woodland island, its shape, distance to the nearest large woodland, mean distance to the three nearest woodland islands and others) and 10 dependent variables (number of species of trees, shrubs, herbaceous plants, species characteristic of the classesQuerco-Fagetea, Vaccinio-Piceetea, Molinio-Arrhenatheretea, Rudero-Secalietea and others). It was found that the floristic composition of the communities investigated was connected primarily with the isolation of the woodland islands. With increasing isolation the total number of species decreased, as did the number of herbaceous plants and that of herbaceous plants of the classesQuerco-Fagetea andVaccinio-Piceetea, while the number of species of the classesMolinio-Arrhenatheretea andRudero-Secalietea increased. Conditions of the habitat such as the inclination of slopes, shading, and soil fertility were less important.     

Micro-habitat drivers of saproxylic beetle assemblages in old woodlands of Mediterranean cork oak (Quercus suber)

1. Cork oak landscapes are fascinating ecosystems, historically managed for cork extraction. The persistence in this habitat of many hollow veteran trees provides suitable micro-habitats for saproxylic beetles.
2. We investigated the saproxylic beetle community of two isolated cork oak woodlands of central Italy with different degree of recovery after human transformation: (1) an open woodland and (2) a dense mixed woodland, both dominated by cork oak trees.
3. We found endemic, rare and threatened saproxylic beetles in both the areas, confirming the important conservation value of cork oak landscapes. In the open woodland we observed a higher number of species in all trophic categories, except for mycophagous specialists. Several microhabitat variables reflected the different stage of recovery of the two woodlands.
4. Our findings suggest the crucial role of diversified environments in protected areas: even a small difference in the degree of recovery (i.e., tree closeness) can affect the number of beetle species. Specifically, we found (1) more xerophilous species in the open woodland and (2) more mesophilous species in the dense mixed woodland.

     

Carabid beetle diversity and mean individual biomass in beech forests of various ages

Carabid beetle diversity and mean individual biomass (MIB) were analysed in three different successional stages of beech tree stands (60, 80 and 150 years old). Carabid beetles were captured using pitfall traps placed at nine sites (three per age class) in the Papuk Mountain of East Croatia during 2008. A cluster analysis identified three groupings that corresponded to the beech age classes. MIB values increased with stand age, ranging from 255 in 60-year-old stand to 537 in the oldest forests. The 80-year-old stand showed the highest species richness and diversity values. With respect to species composition, large species such as Carabus scheidleri and Carabus coriaceus were dominant only in the oldest forests. Furthermore, species that overwinter in the larval stage were more abundant in the oldest forests (45% of the total number of individuals from the 150-year-old stand) than in the younger ones (20% of individuals from 60-year-old, and 22% of individuals from 80-year-old stands). Our results showed that the analyses of species composition and life history traits are valuable for estimating the conservation values of older forests. Although the investigated sites form part of a continuous forested area and are only a couple of kilometres apart, MIB values detect significant differences associated with forest age and can be a useful tool in evaluating the degree to which a forest reflects a natural state.     

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.     

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.     

Flora, vegetation, and soil in broadleaved ancient and planted woodland, and scrub on RøsnÆs, Denmark

9 small ancient woodlands (>200 years), 9 planted woodland sites (25 — ca. 100 years), and 6 sites in grazed scrub (ca. 100 years) on the RøsnÆs peninsula, Denmark, showed characteristic differences: in ancient woodland, the tree and shrub layer was fairly rich, the field layer rather poor in species. The dominance of spring flowering geophytes, the abundance of Anemone nemorosa , and the occurrence of Corylus avellana and Polygonatum multiflorum were characteristic. pH of the soil was relatively low, organic matter content high, and light intensity at the forest floor in summer low. In planted woodland there was more light, and the field layer was rich in short-lived species, but poor in spring flowering geophytes. Many woodland species were rare in planted woodland, some did not at all occur there, and none were specific for this type of woodland. The scrub was marked by grazing and a strong relief, hence pH was high and organic matter content low. The field layer was rich both in shortlived species and in spring flowering geophytes. — It is suggested that ancient woodland species (i.e. species restricted to or preferably occurring in woodlands existing prior to the enclosure ca. 200 years ago) is a heterogenous group, consisting of a) species favoured by traditional woodland management; b) species restricted to woodlands where specific environmental (e.g. soil) conditions have had sufficient time to develop; c) species with limited ability to spread or establish; or d) species which in ancient woodland are represented by small and scattered populations.