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.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

生境片段化对千岛湖马尾松林内土壤种子库的影响

土壤种子库是森林群落更新的主要来源之一,对森林的演替和恢复等具有重要意义。生境片段化现象正日益严重地影响着森林群落,并可影响森林土壤种子库。研究了千岛湖地区的大陆及岛屿次生马尾松林内土壤种子库的组成及其影响因素(e.g.,岛屿面积,形状指数,隔离度和距岛屿边缘距离等)。根据大陆和岛屿的面积及边缘梯度,采用大数量小样方法,分别在土壤种子库最大化(初冬,2015年12月)和最小化(晚春,2016年4月)时期对马尾松林内土壤进行了机械取样。对土壤样品进行萌发实验,检测了两个时期的土壤种子库上层(0—2 cm)和下层(2—5 cm)种子组成,并通过广义线性混合效应模型等手段分析其影响因素。结果显示:(1)所有316个土壤样本中,萌发出幼苗1422株,隶属于29科、40属、41种。其中,木本植物幼苗占13种1024株,草本占28种398株。(2)Jaccard指数和相关性分析均显示初冬、晚春时期的土壤种子库组成具有很高的相似性;土壤种子库上、下层组成的相似性也很高。(3)广义线性混合效应模型分析显示,在大陆和岛屿上,土壤种子库下层种子含量低于上层;而大陆样地土壤种子库中的木本植物种子数较岛屿样地高。岛屿上,土壤种子库中的种子数随土层的加深而降低;随边缘梯度升高也下降,尤其是草本植物的种子。对于岛屿上的木本植物,不耐阴种的种子数量远大于耐阴种,尤其是土壤下层。表明千岛湖地区马尾松林内土壤种子库组成受到生境片段化的影响,进而可能作用于该类型森林群落的演替。     

Factors influencing vascular plant diversity on 22 islands off the coast of eastern North America

Aim The influence of physiographic and historical factors on species richness of native and non‐native vascular plants on 22 coastal islands was examined. Location Islands off the coast of north‐eastern USA and south‐eastern Canada between 41° and 45° N latitude were studied. Island size ranges from 3 to 26,668 ha. All islands were deglaciated between 15,000 and 11,000 yr bp ; all but the four New Brunswick islands were attached to the mainland until rising sea level isolated them between 14,000 and 3800 yr bp . Methods Island species richness was determined from floras compiled or revised since 1969. Simple and multiple regression and rank correlation analysis were employed to assess the relative influence of independent variables on species richness. Potential predictors included island area, latitude, elevation, distance from the mainland, distance from the nearest larger island, number of soil types, years since isolation, years since deglaciation, and human population density. Results Native vascular plant species richness for the 22 islands in this study is influenced most strongly by island area, latitude, and distance from the nearest larger island; richness increases with island area, but decreases with latitude and distance from the nearest larger island as hypothesized. That a similar model employing distance from the mainland does not meet the critical value of P confirms the importance of the stepping‐stone effect. Habitat diversity as measured by number of soil types is also an important predictor of native plant species richness, but at least half of its influence can be attributed to island area, with which it is correlated. Two historical factors, years since deglaciation and years since isolation, also appear to be highly correlated with native species richness, but their influence cannot be separated from that of latitude for the present sample size. Non‐native vascular plant species richness is influenced primarily by island area and present‐day human population density, although human population density may be a surrogate for the cumulative effect of several centuries of anthropogenic impacts related to agriculture, hunting, fishing, whaling, tourism, and residential development. Very high densities of ground‐nesting pelagic birds may account for the high percentage of non‐native species on several small northern islands. Main conclusions Many of the principles of island biogeography that have been applied to oceanic islands apply equally to the 22 islands in this study. Native vascular plant species richness for these islands is strongly influenced by physiographic factors. Influence of two historical factors, years since deglaciation and years since isolation, cannot be assessed with the present sample size. Non‐native vascular plant species richness is influenced by island area as well as by human population density; human population density may be a surrogate for other anthropogenic impacts.     

The effects of island area, isolation and source population size on the presence of the grayling butterfly Hipparchia semele (L.) (Lepidoptera: Satyrinae) on British and Irish offshore islands

Records of Hipparchia semele on British and Irish islands have been modelled against island area, isolation (sea and land distance) and the size of the nearest potential source populations. All three variables have been found to contribute significantly to the presence or absence of H. semele on the islands. Isolation is a more significant predictor than island area. This result differs from the multiple species case where area was found to be a more important influence than isolation. Records on islands are also shown to depend on the size of populations at the nearest sources; this underpins the relationships identified for the multiple species case, first, between the number of species on islands and at nearest sources and, second, between the incidence of species on islands and at nearest sources. There are clear indications that smaller islands may become increasingly marginalized for H.semele; with ongoing habitat loss, because isolation increases and source populations become sparser, the probability of H. semele recolonizing islands also decreases.     

Plant diversity patterns in the Aljibe Mountains (S. Spain): a comprehensive account

The Aljibe Mountains are located in the southern tip of the Iberian Peninsula and have a remarkable biogeographical interest. The complete plant species list (trees, climbers, shrubs, perennial and annual herbs, ferns, lichens, bryophytes and macroscopic algae) was recorded in four 0.1 ha plots from each of the most representative community types (Quercus suber woodland, Q. canariensis forest, open heathland and Q. coccifera shrubland). Up to 119 plant species were found in total in the Q. suber woodland plot. The diversity of woody plants was analysed from 44 samples of cover (100 m line), and the herbaceous layer was explored in 200 quadrats (of 0.5 × 0.5 m). Three biodiversity components (species richness, endemism, and taxonomic singularity) were evaluated in both shrub and herbaceous layers. Open heathlands showed the highest richness of endemic species, both woody and herbaceous. The highest number of woody species was found in the evergreen Q. suber woodland, and of herbaceous species in the semi-deciduous Q. canariensis woodland. Taxonomic singularity was higher in Q. canariensis woodlands and Q. coccifera shrublands for woody species, but there were no significant differences in the herbaceous layer. Local species diversity of heathlands in this region resembles that of South African heathlands (fynbos), despite the obvious geographic and floristic distance, and contrasts with the low diversity of biogeographically closer, European temperate heathlands. The Aljibe Mountains show high diversity values for different life forms (from trees to mosses) and spatial scales (from community to region), and are rich in endemic species. Thus, this area should be recognised as a relevant unit within the Mediterranean plant diversity hot spots.     

Quantifying island isolation – insights from global patterns of insular plant species richness

Isolation is a driving factor of species richness and other island community attributes. Most empirical studies have investigated the effect of isolation measured as distance to the nearest continent. Here we expanded this perspective by comparing the explanatory power of seventeen isolation metrics in sixty‐eight variations for vascular plant species richness on 453 islands worldwide. Our objectives were to identify ecologically meaningful metrics and to quantify their relative importance for species richness in a globally representative data set. We considered the distances to the nearest mainland and to other islands, stepping stone distances, the area of surrounding landmasses, prevailing wind and ocean currents and climatic similarity between source and target areas. These factors are closely linked to colonization and maintenance of plant species richness on islands. We tested the metrics in spatial multi‐predictor models accounting for area, climate, topography and island geology. Besides area, isolation was the second most important factor determining species richness on the studied islands. A model including the proportion of surrounding land area as the isolation metric had the highest predictive power, explaining 86.1% of the variation. Distances to large islands, stepping stone distances and distances to climatically similar landmasses performed slightly better than distance to the nearest mainland. The effect of isolation was weaker for large islands suggesting that speciation counteracts the negative effect of isolation on immigration on large islands. Continental islands were less affected by isolation than oceanic islands. Our results suggest that a variety of immigration mechanisms influence plant species richness on islands and we show that this can be detected at macro‐scales. Although the distance to the nearest mainland is an adequate and easy‐to‐calculate measure of isolation, accounting for stepping stones, large islands as source landmasses, climatic similarity and the area of surrounding landmasses increases the explanatory power of isolation for species richness.     

Effects of woodlot isolation on the dispersion of plants with fleshy fruits

By relating species presence, number and density to the perimeter and isolation of small (0.1–2.75 ha) homogeneous woodlots (n = 43), indications were obtained that some fleshy-fruited species are negatively affected by woodlot isolation. The number of fruit-bearing herbaceous perennial species increased with woodlot area, probably because of the increasing heterogeneity of the herb layer. The density of individuals of the investigated species were negatively correlated to woodlot area. There were also some indications thatthe presence/absence of species was related to the length of the forest edge (e.g. Hederá helix). Because woodlot area and shape index were strongly correlated, the effects of increased forest area and forest edge were hard to separate. The probability of occurrence of Lonicera periclymenum and the density oí Ilex aquifolium decreased with an increasing degree of isolation of present forest islands. The chance of occurrence of L. periclymenum decreased also with the degree of isolation in the past, measured from topographical maps of about 60 yr ago. The total area of broad-leaved forest within a radius of 500 m, as well as the distance to the nearest connecting landscape element, seem to be good indicators of woodlot isolation.     

Positive Effects of Non-Native Grasses on the Growth of a Native Annual in a Southern California Ecosystem

Fire disturbance is considered a major factor in the promotion of non-native plant species. Non-native grasses are adapted to fire and can alter environmental conditions and reduce resource availability in native coastal sage scrub and chaparral communities of southern California. In these communities persistence of non-native grasses following fire can inhibit establishment and growth of woody species. This may allow certain native herbaceous species to colonize and persist beneath gaps in the canopy. A field manipulative experiment with control, litter, and bare ground treatments was used to examine the impact of non-native grasses on growth and establishment of a native herbaceous species, Cryptantha muricata. C. muricata seedling survival, growth, and reproduction were greatest in the control treatment where non-native grasses were present. C. muricata plants growing in the presence of non-native grasses produced more than twice the number of flowers and more than twice the reproductive biomass of plants growing in the treatments where non-native grasses were removed. Total biomass and number of fruits were also greater in the plants growing in the presence of non-native grasses. Total biomass and reproductive biomass was also greater in late germinants than early germinants growing in the presence of non-native grasses. This study suggests a potential positive effect of non-native grasses on the performance of a particular native annual in a southern California ecosystem.     

Effects of habitat isolation on pollinator communities and seed set

Destruction and fragmentation of natural habitats is the major reason for the decreasing biodiversity in the agricultural landscape. Loss of populations may negatively affect biotic interactions and ecosystem stability. Here we tested the hypothesis that habitat fragmentation affects bee populations and thereby disrupts plant-pollinator interactions. We experimentally established small ”habitat islands” of two self-incompatible, annual crucifers on eight calcareous grasslands and in the intensively managed agricultural landscape at increasing distances (up to 1000 m) from these species-rich grasslands to measure effects of isolation on both pollinator guilds and seed set, independently from patch size and density, resource availability and genetic erosion of plant populations. Each habitat island consisted of four pots each with one plant of mustard (Sinapis arvensis) and radish (Raphanus sativus). Increasing isolation of the small habitat islands resulted in both decreased abundance and species richness of flower-visiting bees (Hymenoptera: Apoidea). Mean body size of flower-visiting wild bees was larger on isolated than on nonisolated habitat islands emphasizing the positive correlation of body size and foraging distance. Abundance of flower-visiting honeybees depended on the distance from the nearest apiary. Abundance of other flower visitors such as hover flies did not change with increasing isolation. Number of seeds per fruit and per plant decreased significantly with increasing distance from the nearest grassland for both mustard and radish. Mean seed set per plant was halved at a distance of approximately 1000 m for mustard and at 250 m for radish. In accordance with expectations, seed set per plant was positively correlated with the number of flower-visiting bees. We found no evidence for resource limitation in the case of mustard and only marginal effects for radish. We conclude that habitat connectivity is essential to maintain not only abundant and diverse bee communities, but also plant-pollinator interactions in economically important crops and endangered wild plants. Received: 7 May 1999 / Accepted: 19 July 1999     

Origin and evolution of endemic plants of the Bonin (Ogasawara) Islands

The Bonin Islands are typical oceanic islands, located at the western part of the North Pacific Ocean and approximately 1,000 km south of mainland Japan. This archipelago consists of about 20 small islands. Although floristic diversity is low due to the small area and limited environmental diversity, the Bonin Islands harbor unique endemic flora as in other well-known oceanic islands. This paper presents a brief summary of the results obtained from recent studies on the endemic flora of the Bonin Islands. The results are reviewed in relation to the four stages of the evolution of endemic flora in the oceanic islands; migration, establishment, enlargement and diversification. The ancestors of the flora originated mostly from tropical and subtropical Southeast Asia or mainland Japan by rare events of long distance dispersal. The proportion of bird-dispersed species is relatively high as for other oceanic islands. Genetic data sets obtained from allozyme variation in some endemic species suggest that migration occurred several million years ago and genetic diversity is correlated with current population size. At the time of establishment, self-compatible plants are expected to have an advantage. However, the percentage of dioecious plants is relatively high. This is partly due to evolutionary changes from hermaphroditic ancestors to dioecy which occurred in two genera in the Bonin Islands. In addition, there are some examples of evolutionary changes from herbaceous ancestors to woody endemics. Adaptive radiation is found in some genera, although the number of congeneric endemic species is less than five. Studies of allozyme variation inPittosporum, Symplocos andCrepidiastrum showed that genetic identity is generally very high between congeneric species in spite of their distinct morphologies. This result suggests that divergence of these species occurred rather recently and distinct morphological differences are based on a limited number of genetic changes.     

Invasive African big-headed ants, Pheidole megacephala, on coral cays of the southern Great Barrier Reef: distribution and impacts on other ants

Infestation of islands by exotic ants is widespread and increasing due to human activities throughout the world. Exotic ants, particularly the invasive African big-headed ant, Pheidole megacephala (Fabricius), are of great conservation concern for coral cays at the southern end of the Great Barrier Reef, Australia. Little is known, however, about the distribution and ecological impacts of invasive ants in this insular system. We surveyed the ants of 14 vegetated coral cays recording a total of 24 ant species, including at least nine exotics. Pheidole megacephala was by far the most abundant and widespread species, occurring on 11 of 14 islands, often in very large numbers. The inter-island distribution of P. megacephala was best explained by human activities, with frequently visited, and to a lesser degree disturbed islands, more likely to be infested. On large islands (≥?10?ha) P. megacephala exhibited distinct habitat preferences, occurring in significantly lower abundances within heavily-shaded Pisonia grandis forest in the centre of islands, compared to more open, fringing woodland or shrubland. On smaller islands (<10?ha) with less extensive Pisonia stands, P. megacephala penetrated throughout the forest where its abundance was similar to that in open woodland. Despite considerable differences in biotic (floristic composition) and abiotic factors (e.g. island size) as well as the spatial configuration among islands, the severity of infestation by P. megacephala best explained variation in species richness, abundance and assemblage composition of other ants. We suggest a number of strategies to manage P. megacephala infestations on these islands.     

Winter bird communities in forest plantations in western England and their response to vegetation,growth stage and grazing

This paper presents the results of a study of wintering bird communities across a wide range of coniferous, broadleaved and mixed forest stands in the Forest of Dean, western England. Bird communities of broadleaved and coniferous woodland differed with respect to their species composition. The mean number of individual birds recorded increased linearly with woodland age and was not influenced by woodland type, stand size or the presence of grazing. Woodland age and type and the presence or absence of grazing all significantly influenced bird species richness and the proportions of the bird community made up by granivores, insectivores and omnivores. Broadleaved stands held more species than coniferous stands. Ungrazed stands held significantly more species, particularly seed-eating species, than grazed stands and this effect was independent of woodland type. Ordination was used to relate variation in tree species composition and stand structure to bird community composition. A larger number of species was associated with broadleaved stands and stands with abundant undergrowth than was associated with coniferous stands or stands with little undergrowth. Woodland age had less effect on bird community composition than the extent of undergrowth and the conifer to broadleaf ratio. The results of this work have relevance to the enhancement of winter bird communities in commercial forests.     

Influence of fire and soil nutrients on native and non-native annuals at remnant vegetation edges in the Western Australian wheatbelt

Abstract. The effect of fire on annual plants was examined in two vegetation types at remnant vegetation edges in the Western Australian wheatbelt. Density and cover of non-native species were consistently greatest at the reserve edges, decreasing rapidly with increasing distance from reserve edge. Numbers of native species showed little effect of distance from reserve edge. Fire had no apparent effect on abundance of non-natives in Allocasuarina shrubland but abundance of native plants increased. Density of both non-native and native plants in Acacia acuminata-Eucalyptus loxophleba woodland decreased after fire. Fewer non-native species were found in the shrubland than in the woodland in both unburnt and burnt areas, this difference being smallest between burnt areas. Levels of soil phosphorus and nitrate were higher in burnt areas of both communities and ammonium also increased in the shrubland. Levels of soil phosphorus and nitrate were higher at the reserve edge in the unburnt shrubland, but not in the woodland. There was a strong correlation between soil phosphorus levels and abundance of non-native species in the unburnt shrubland, but not after fire or in the woodland. Removal of non-native plants in the burnt shrubland had a strong positive effect on total abundance of native plants, apparently due to increases in growth of smaller, suppressed native plants in response to decreased competition. Two native species showed increased seed production in plots where non-native plants had been removed. There was a general indication that, in the short term, fire does not necessarily increase invasion of these communities by non-native species and could, therefore be a useful management tool in remnant vegetation, providing other disturbances are minimised.     

Vegetation-environment relationships in a heavy metal-dry grassland complex

Heavy-metal content is assumed to be the most important edaphic factor that determines vegetation composition on contaminated soil. We compared the effects of heavy metals on species composition and species richness in the heavy metal-dry grassland complex of the Bottendorf Hills (Central Germany) with those of other environmental factors. Based on 206 relevés, we distinguished nine communities of the classesKoelerio-Corynephoretea andFestuco-Brometea. Four communities in which the metallophytesArmeria maritima subsp.halleri andMinuartia verna subsp.hercynica occurred with high frequency were classified as heavy metal subassociations of four different dry grassland associations because of the dominance of dry grassland species. We measured the soil content of copper, zinc and lead, and the carbonate content, C/N ratio, pH and conductivity of the soil, soil depth and incident radiation per site. The first axis resulting from a DCA was positively correlated with the cover and height of the herb layer, the soil depth and soil carbonate content, and negatively with the soil content of copper, the proportion of rocks, the soil C/N ratio and incident radiation per site. The number of vascular plants, bryophyte and lichen species per plot increased with pH up to 7.5 and then decreased slightly. Species richness increased with carbonate content and conductivity of the soil and decreased with the soil C/N ratio. Heavy metal content of the soil and species richness were not correlated. The occurrence of the metallophytes was strongly related to the copper content of the soil. In conclusion our study has shown that heavy metal content is not necessarily the main factor determining the total composition and richness of grasslands on soil containing heavy metals. Heavy metal grasslands are not necessarily floristically distinct from “normal” dry grasslands.     

Demographic response to shading and defoliation in two woodland orchids

Many woodland herbs are long-lived, clonal geophytes that have evolved life histories favoring survival over reproduction. We examined the life history responses of natural populations of two woodland orchid species,Cypripedium calceolus andCephalanthera longifolia to defoliation and heavy shading conducted early in the growing seasons of 2002 and 2003. We asked whether, in view of the importance of growth for the survival of geophytes, treated plants were more likely to exhibit reduced flowering than reduced vegetative growth in the seasons following treatment. We also asked whether plants would suffer reduced ramet performance. Both treatments led to significant declines in flower number per ramet, number of leaves per ramet, and mean ramet height relative to controls inCypripedium. However, inCephalanthera, only shaded plants exhibited significant declines in flower number per ramet, and only defoliated plants exhibited declines in mean ramet height. The number of ramets per plant did not decline relative to controls in either species. Thus, these orchids, especiallyCypripedium, appeared to allocate resources preferentially to vegetative growth functions over sexual reproduction. Per-plant variation in leaf and flower number per ramet, as well as in mean ramet height, consistently declined in response to treatment, significantly so in the case of mean ramet height, suggesting that ramets became more similar within genets. These results suggest both similarities and differences in the ways in whichCephalanthera andCypripedium mobilize resources in response to stress.     

Habitat preferences of aphidophagous coccinellids [Coleoptera]

From May to July 1982 and 1983, coccinellids were sampled in habitats that differed in aphid density, insolation, and host plant type. The number of adults ofCoccinella 7-punctata (CS),C. 5-punctata (CQ),Propylea 14-punctata (PQ),Adonia variegata (AV),Adalia bipunctata (AB),A. 10-punctata (AD), andCalvia 14-guttata (CA) were recorded. AB, AV and CS preferred high, while CQ tolerated low aphid density. CQ, CS, AB and AV preferred sunny, while AD and CA tolerated shaded conditions. CS and AV preferred herbaceous plants, AD and CA large plants, particularly trees, AB showed little specialization, CQ preferred trees early in the season and sparse herbaceous plants later. PQ appeared to be a generalist with a wide tolerance of all these factors, but with a tendency to move from trees to herbaceous plants as the season progressed. Habitats with a high abundance of a species were assumed to be those with on optimum combination of the preferred conditions. There were slight differences in the breadth of habitat preferences, and a little habitat overlap between species. The tendency to aggregate was greater in AB than in other species. Annual variations in species abundance influenced the number of habitats occupied, but not the abundance in the favoured places.     

Small mammals on Massachusetts islands: the use of probability functions in clarifying biogeographic relationships

Summary Thirty-three insular small mammal communities along the coast of Massachusetts (USA) were surveyed to investigate the biogeographic relationships of the insular communities and to examine the distribution patterns of individual species. Nine species of terrestrial small mammals were observed in the total insular fauna, whereas thirteen occurred on the mainland. The species-area relation yielded a z value of 0.06, which is the lowest value yet reported for insular mammal communities.Multiple logistic regression was used to calculate probability functions for each species in order to identify variables potentially important in determining a species' occurrence on islands and to estimate probabilities of occurrence on islands. Statistically significant and ecologically interpretable functions were obtained for all but one species. Occurrence on islands was positively related to increasing island size in four species and to decreasing island isolation in four species. The extremely low z value, negative correlations of species number with isolation variables, and the inclusion of an isolation variable in the logistic functions of four species indicated that immigration was an important determinant of small mammal occurrence on these islands. There was a positive relationship between population density and number of islands occupied.Logistic regression has several advantages over linear discriminant function analysis, and we suggest that it may be useful in other ecological studies and in the preservation of endangered species.     

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.