Vegetation transition following drainage in a high‐latitude hyper‐oceanic ecosystem

Questions: How does draining affect the composition of vegetation? Are certain functional groups favoured? Can soil parameters explain these differences? Location: Central Faroe Islands, treeless islands in the northern boreal vegetation zone. Since 1987, an area of 21 km² at 100–200 m a.s.l. was drained in order to provide water for hydro‐electric production. Method: Vegetation and soil of a drained area and a control, undrained neighbouring area of approximately the same size were sampled in 2007. Six sites were sampled in each area. The vegetation was classified with cluster analysis. Results: Four plant communities were defined in the area: Calluna vulgaris–Empetrum nigrum–Vaccinium myrtillus heath, Scirpus cespitosus–Eriophorum angustifolium blanket mire, Carex bigelowii–Racomitrium lanuginosum moss‐heath, Narthecium ossifragum–Carex panacea mire. Heath was more extensively distributed within, and was the dominant community of the drained area, whereas moss‐heath was more extensive in the undrained area. Blanket mire and mire had approximately the same distribution in both areas. For the blanket mire, species composition indicated drier conditions in the drained than in the undrained area. The drained area had higher frequencies of woody species and lichens, grasses had finer roots and available soil phosphate was considerably higher, whereas the undrained area had higher frequencies of grasses and sedges. Conclusion: The dominant plant communities were different in the two areas, which indicated that the blanket mire was drying in the drained area. Higher concentration of soil phosphate in the drained area also indicated increased decomposition of organic soils owing to desiccation.     

A phytosociological study of a mid-altitude heath in County Durham, Northern England

A phytosociological survey of Hedleyhope Fell, a mid-Durham heath, was carried out. Much of the Fell is rank Nardus grassland, with damper Juncus and Eriophorum patches. Disturbed areas to the west are covered by poor Agrostis/Trifolium/Festuca grasslands. The eastern end of the Fell forms a locally valuable area of heathland. 115 relevés were taken and 131 species of plant recorded. Classification by two-way indicator species analysis suggested the delimitation of 19 noda. The principal ordination axes, produced by detrended correspondence analysis, emphasised the importance of soil moisture gradients, organic/mineral soil gradients and levels of disturbance in determining the vegetation. The Fell, at 300 m, falls between the lowland heaths of southern England and the upland mires of the north. Indeed, its vegetation contains some assemblages and species characteristic of both habitats. The heaths of central, mid-altitude County Durham are found where the coal measures lie close to the surface and so this habitat type has suffered extensively from opencast mining activity in the recent past. Proposed opencast sites, if they go ahead, will destroy the Fell.     

Patterned fens in Scotland: evidence from vegetation and water chemistry

Mires with attributes similar to the patterned fens of boreal continental regions have recently been reported from northern Scotland. This paper reports on the vegetation and water chemistry of 18 of these sites. Most of the 18 mires displayed remarkable uniformity in their vegetation, with Carex lasiocarpa being a main character species. Major variation of plant communities is associated with microtopographical niches, similar to those found on ombrotrophic mire systems and secondarily with pH and conductivity of surface waters. Nine vegetation noda are described and compared with existing phytosociological associations and National Vegetation Classification (NVC) communities. The dominant Sphagnum - Carex communities are intermediate between the Oxycocco-Sphagnetea and the Caricion lasiocarpae and are thus difficult to assign to existing associations or to NVC communities. Although there is a large variation in pH of surface waters (3.9–6.6), mean measurements for terrestrial and aquatic communities are 5.12 and 5.51 respectively. pH levels are comparable with poor fen systems elsewhere in Britain and slightly higher than patterned fens in continental regions of Europe and North America. The vegeta-tional and chemical attributes of these sites therefore confirm the existence of true patterned fen systems in the British Isles.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Off-season uptake of nitrogen in temperate heath vegetation

In this field study we show that temperate coastal heath vegetation has a significant off-season uptake potential for nitrogen, both in the form of ammonium and as glycine, throughout winter. We injected ¹⁵N-ammonium and ¹⁵N 2×(¹³C)-glycine into the soil twice during winter and once at spring. The winter temperatures were similar to those of an average winter in the northern temperate region of Europe, with only few days of soil temperatures below zero or above 5°C. The vegetation, consisting of the evergreen dwarf shrub Calluna vulgaris, the deciduous dwarf shrub Salix arenaria, and the graminoids Carex arenaria and Deschampsia flexuosa, showed high root uptake of both forms of nitrogen, both 1 day after labelling and after a month, in species specific temporal patterns. Plant uptake of ¹³C was not significant, providing no further evidence of intact uptake of glycine. Translocation of the labelled nitrogen to shoots was generally evident after 1 month and increased as spring approached, with different translocation strategies in the three plant functional types. Furthermore, only the graminoids showed shoot growth during winter. Increasing plant nitrogen concentration from fall to spring at temperate heaths may, hence, be due to nitrogen uptake. Our results suggest that the potential for nitrogen uptake in plants at winter is of the same order of magnitude as at summer. Hence, winter nitrogen uptake in ecosystems in the temperate/boreal region should be considered when making annual nitrogen budgets of heath ecosystems, and the view of plant nutrient uptake as low in this climatic region during winter should be revised.     

Vegetation change in Southeast Greenland? Tasiilaq revisited after 40 years

Questions: Are there changes in species composition of the oceanic, Low‐Arctic tundra vegetation after 40 years? Can possible changes be attributed to climate change? Location: Ammassalik Island near Tasiilaq, Southeast Greenland. Methods: Species composition and cover of 11 key vegetation types were recorded in 110 vegetation survey plots in 1968–1969 and in 11 permanent plots in 1981. Recording was repeated in 2007. Temporal changes in species composition and cover between the surveys were tested using permutation tests linked with constrained ordinations for vegetation types, and Mann–Whitney tests for individual species. Changes in vegetation were related to climate change. Results: Although climate became warmer over the studied period, most of the vegetation types showed minor changes. The changes were most conspicuous in mire and snowbed vegetation, such as the Carex rariflora mire and Hylocomium splendens snowbed. In the C. rariflora mire, species number and cover of vascular plants and cover of bryophytes increased, whereas in the H. splendens snowbed species numbers of vascular plants, bryophytes, and also lichens increased. Lichen richness increased in the Carex bigelowii snowbed and cover of bryophytes in the Salix herbacea snowbed. No such changes occurred in the Alchemilla glomerulans meadow, Alchemilla alpina snowbed and Phyllodoce coerulea heath. There was no change of species composition within the Salix glauca scrub, A. alpina snowbed, lichen grassland and the Empetrum nigrum and Phyllodoce coerulea heaths. Most changes resulted from increasing frequency or cover of some species; there were very few decreasing species. Most of the increasing species indicate drier substrate conditions. Conclusions: Only minor changes in species composition and cover were detected in the vegetation types studied. These changes were probably caused by milder winters and warmer summers during the years before the 2007 sampling. Climate warming may have reduced the duration of snow cover and soil moisture, particularly in snowbed and mire habitats, where species composition change was most pronounced. However, its magnitude was insufficient to cause a major change in species composition. Thus, on the level of plant community types, tundra vegetation near Tasiilaq was rather stable over the last 40 years.     

Biodiversity and vegetation dynamics in the coastal heathlands of western Norway

The heaths of Lygra, western Norway, were investigated, and 19 communities at the association level were distinguished. Distinct ecotypes of Ranunculus acris and Poa humilis and microspecies of Euphrasia were restricted to the grass heaths. No distinct ecotypes were found in the Calluna -dominated vegetation. It is suggested that the regular burning, which has been part of the management of the area, has activated the seed bank of the ericaceous heath, and thus maintained genetic diversity and slowed down ecotype differentiation. The grass heaths, with no combustible material, have been unaffected in this way. Permanent plots in the two dominant heath types, dry heath (Vaccinio-Callunetum) and wet heath (Ericetum tetralicis) were followed over six years and the data analysed using a modification of Jaccard's index, allowing for species abundances.     

Plant communities in broad-leaved deciduous forests in Hordaland county, Western Norway

The vegetation of broad-leaved deciduous forests in Hordaland, Western Norway is described with regards to species composition and structure. The investigation is based on phytosociological analyses of native forest stands dominated by Corylus avellana, Fraxinus excelsior, Ulmus glabra , or Tilia cordata. Two-way-indicator analysis (TWINSPAN) and correspondence analysis (CA) are used to distinguish different vegetation types and assess possible gradients in the vegetation data. The vegetation types are discussed in relation to equivalent forest communities and to syntaxonomical units. Two forest types are described at the first hierarchical TWINSPAN level: 1) A hygrophilous and slightly thermophilous Fraxinus excelsior-Cirriphyllum piliferum forest characterised by an open tree canopy, a dense field layer of tall herbs and ferns, and a high cover of bryophytes. 2) A thermophilous and less hygrophilous Corylus avellana-Brachypodium sylvaticum forest characterised by a dense tree layer, a more open field layer with larger elements of small herbs, and a somewhat lower bryophyte cover. The CA analysis clearly separates the samples from the first TWINSPAN division along the first ordination axis. Five forest types have been described at the second hierarchical level, mainly associated with differences in mesotrophic and eutrophic species, but there is clearly a gradient structure in the species composition between these plant communities. In relation to syntaxonomy, the first TWINSPAN division supports the separation of the west Norwegian broad-leaved deciduous forests into a hygrophilous plant community, the Eurhynchio-Fraxinetum (Blom 1982) Øvstedal 1985 and a drier and more thermophilous community, the Primulo-Ulmetum (Blom 1982) Øvstedal 1985.     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Phytosociological amplitudes of some Dutch coastal species and their ecological interpretation

A 4-axis Reciprocal Averaging ordination of phytosociological tables of Dutch plant communities revealed environmentally interpretable gradients: (1) overall moisture, (2) salinity, (3) soil fertility, and (4) structural complexity of vegetation. The quantitative occurrence of all species along these gradients has been computed. Some examples of the distribution of species from coastal communities are given, to show how these data may help to summarize and visualize existing knowledge and how they can form a basis for further study.Nomenclature of species follows Heukels-van Ooststroom (1975), that of syntaxonomical units Westhoff & den Held (1969).     

Plant communities in the terrestrial-aquatic transition zone in the paramo of Chingaza, Colombia

Plant communities in the terrestrial-aquatic transition zone in the paramo of Chingaza, Colombia. High Andean paramo ecosystems are an important water resource for many towns, and major cities in this region. The aquatic and wetland vegetation of different paramo lakes, pond, swamps and bogs was studied according to the classical phytosociological approach, which is based on homogenous stands, but excludes any border phenomena or transitional zone. The present research aimed at determining the aquatic and wetland vegetation along different moisture gradients. A total of 89 species in 30 transects were reported, of which Crassula venezuelensis, Carex honplandii, Callitriche nubigena, Eleocharis macrostachya, Ranunculus flagelliformis, R. nubigenus, Eleocharis stenocarpa, Galium ascendens y Alopecurus aequalis were present in more than one third of the transects. Numerical classification and indicator species analysis resulted in the definition of the next 18 communities: 1) Calamagrostis effusa, 2) Sphagnum cuspidatum, 3) Cyperus rufus, 4) Eleocharis stenocarpa, 5) Carex acutata, 6) Poa annua,7) Valeriana sp., 8) Ranunculus flagelliformis, 9) Carex bonplandii, 10) Festuca andicola. 11) Muhlenbergia fustigiata, 12) Elatine paramoana, 13) Iso?tes palmeri, 14) Crassula venezuelensis, 15) Lilaeopsis macloviana, 16) Callitriche nubigena, 17) Potamogeton paramoanus and 18) Potamogeton illinoensis. The ordination of communities reveals the presence of three different aquatic-terrestrial gradients which are related to the life form structure of species that characterized the various communities. We concluded that patchiness and heterogeneity of the vegetation is mainly the result of alterations caused by human activities (burning, cattle raise and material extraction for road and dam construction).     

藏北高原草地群落的数量分类与排序

采用TWINSPAN数量分类和DCA、CCA排序的方法,对藏北高原草地29个样点进行统计分析。结果显示:(1)TWINSPAN数量分类将藏北高寒草地群落划分成10种类型。(2)样点DCA排序第一轴基本反映了水分环境梯度,第二轴基本反映了热量梯度。(3)TWINSPAN分类所划分的各群落在DCA排序图上都有各自的分布范围和界限,说明DCA排序能较好的反应各优势群落与其环境资源之间的关系。(4)样点CCA排序表明,影响群落分布的首要环境因子是水分因子(年均降水量)和空间因子(经度),其次是热量因子(年均温度),CCA排序进一步阐明了群落分布决定于水分和温度等环境因子,并间接验证了TWINSPAN的分类结果。(5)物种CCA排序和TWINSPAN分类结果表明:植物群落中物种的分布格局与植物群落类型的分布格局存在一定的相似性。     

Zonation of plant cover and environmental factors in wetlands of the Sanjiang Plain,northeast China

The zonation of depressional and riparian wetlands in the Sanjiang Plain of northeastern China was studied to describe their vegetation composition and environmental variables. We sampled 108 plots in 6 depression and riparian wetlands. Samples were classified into 4 groups using two‐way indicator species analysis (TWINSPAN). Emergent marsh vegetation was characterized by Carex lasiocarpa and C. pseudocuraica, meadow marsh vegetation by tussock species such as Carex appendiculata and C. meyeriana, wet meadow vegetation by Calamangrostis angustifolia and Carex appendiculata, and shrub meadow vegetation by the shrubs Betula fruticosa, Alnus sibirica and Salix rosmarinifolia and the graminids Carex schmidtii and Calamagrostis angustifolia. CCA ordination showed that water table, organic matter and available N were the major factors explaining the vegetation zonation pattern. Compared with other Northern Hemisphere regions, bog and fen vegetation are completely absent due to climatic conditions unfavorable for peat formation. Out of four vegetation types, only the Carex lasiocarpa community and the C. pseudo‐curaica community have been found in other regions. However, at the species level many species are widespread and some species are vicariant or pseudovicariant to other regions in the world. Our study suggests that topography and hydrology may be the most important determinants of the vegetation pattern in this region.     

Composition and ecology of the Quercus coccifera L. communities along the eastern Adriatic coast (NE Mediterranean)

The aim of this study is to classify and describe all plant communities with Quercus coccifera covering the entire eastern Adriatic coast and islands from north Croatia to south Albania, and to relate their species composition, chorotypes and life forms to environmental factors using Pignatti ecological indicator values. From total 70 phytosociological relevés, we identified and described four floristically and ecologically distinctive vegetation communities (two new proposed subassociations, one association and stand each) using TWINSPAN and the Braun-Blanquet classification scheme. In Croatia and Montenegro, Q. coccifera is forming macchia within the Fraxino orni–Quercetum cocciferae pistacietosum lentisci. Quercus coccifera occurs only sparsely in south Croatia as a shrubland within Fraxino orni–Quercetum cocciferae nerietosum oleandri subassociation or macchia within the Erico arboreae–Arbutetum unedonis association. Despite the difference in biogeographic position and bioclimates, low shrubby Albanian Q. coccifera stands are more closely related to the Q. coccifera communities from the western Mediterranean. Eastern Adriatic communities appear exclusively within the Querceta ilicis vegetation zone and spread within the meso-Mediterranean belt. They nevertheless are an important part of the region's natural heritage and management plans must ensure that all forms of land are used in a sustainable way.     

Vegetation gradients in Atlantic Europe: the use of existing phytosociological data in preliminary investigations on the potential effects of climate change on British vegetation

• 1 This paper aims to demonstrate the use of available vegetation data from the phytosociological literature in preliminary analyses to generate hypotheses regarding vegetation and climate change.
  • 2 Data for over 3000 samples of calcareous grassland, mesotrophic grassland, heath and woodland vegetation were taken from the literature for a region in the west of Atlantic Europe and subjected to ordination by detrended correspondence analysis in order to identify the main gradients present.
    • 3 Climate data were obtained at a resolution of 0.5° from an existing database. The relationship between vegetation composition and climate was investigated by the correlation of the mean scores for the first two ordination axes for each 0.5° cell with the climate and location variables.
      • 4 The ordinations resulted in clear geographical gradients for calcareous grasslands, heaths and woodlands but not for mesotrophic grasslands. Significant correlations were shown between some of the vegetation gradients and the climate variables, with the strongest relationships occurring between the calcareous grassland gradients and July temperature, latitude and oceanicity. Some of the vegetation gradients were also inferred to reflect edaphic factors, management and vegetation history.
        • 5 Those gradients that were related to temperature were hypothesized to reflect the influence of a progressively warmer climate on species composition, providing a baseline for further studies on the influence of climate change on species composition.
        • 6 The validity of the literature data was assessed by the collection of an original set of field data for calcareous grasslands and the subsequent ordination of a dataset containing samples from both the literature and the field. The considerable overlap between the samples from the literature and the field suggest that literature data can be used, despite certain limitations. Such preliminary analyses, using readily available data, can thus achieve useful results, thereby saving lengthy and costly field visits.

     

Macrophyte community structure and species occurrence in relation to environmental determinants in the ephemeral aquatic habitats of Gavdos,Greece

The aims of this study were to explore the environmental factors that determine the distribution of plant communities in temporary rock pools and provide a quantitative analysis of vegetation–environment relationships for five study sites on the island of Gavdos, southwest of Crete, Greece. Data from 99 rock pools were collected and analysed using Two-Way Indicator Species Analysis (TWINSPAN), Detrended Correspondence Analysis (DCA) and Canonical Correspondence Analysis (CCA) to identify the principal communities and environmental gradients that are linked to community distribution. A total of 46 species belonging to 21 families were recorded within the study area. The dominant families were Labiatae, Gramineae and Compositae while therophytes and chamaephytes were the most frequent life forms. The samples were classified into six community types using TWINSPAN, which were also corroborated by CCA analysis. The principal gradients for vegetation distribution, identified by CCA, were associated with water storage and water retention ability, as expressed by pool perimeter and water depth. Generalised Additive Models (GAMs) were employed to identify responses of four dominant rock pool species to water depth. The resulting species response curves showed niche differentiation in the cases of Callitriche pulchra and Tillaea vaillantii and revealed competition between Zannichellia pedunculata and Chara vulgaris. The use of classification in combination with ordination techniques resulted in a good discrimination between plant communities. Generalised Additive Models are a powerful tool in investigating species response curves to environmental gradients. The methodology adopted can be employed for improving baseline information on plant community ecology and distribution in Mediterranean ephemeral pools. Handling editor: S. M. Thomaz     

The Balkan wet grassland vegetation: a prerequisite to better understanding of European habitat diversity

The knowledge of broad-scale floristic variation in wet grasslands, which are endangered throughout Europe, is still limited and some regions have remained unexplored so far. In addition, hitherto published phytosociological studies were concentrated at the national level and therefore national vegetation classifications are not consistent with each other. In order to overcome these shortcomings of traditional phytosociology, we gathered original data from Bulgaria and analysed them together with the data from Central Europe. We further analysed major compositional gradients within Bulgarian wet grasslands and changes in species richness along them. We sampled 164 wet grassland vegetation plots throughout Bulgaria. We further prepared a restricted data set of wet grasslands from Central-European phytosociological databases. Both data sets were merged and classified by modified TWINSPAN. Four distinct vegetation types were differentiated. Even if they correspond with traditional alliances, which are primarily drawn as geographically defined units in Western and Central Europe (sub-Mediterranean Trifolion resupinati, sub-continental Deschampsion cespitosae and Molinion caeruleae and sub-oceanic Calthion palustris), they all occur in Bulgaria. When more precise classification was applied, two types of sub-Mediterranean wet grasslands and one high-altitude type of Calthion grasslands were detected solely in Bulgaria. DCA analysis showed that altitude is a dominant gradient controlling variation in Balkan wet grasslands. The second DCA axis was interpreted as the gradient of nutrient availability. Species richness shows skewed-unimodal trends along both major gradients, with the highest species richness in intermittently wet nutrient-limited grasslands. Tukey post-hoc test of altitudinal differences amongst vegetation types is significant for all pairs of clusters, suggesting that altitudinal differentiation is responsible for co-occurrence of nearly all European types of wet grasslands in Bulgaria. Our results suggest that (1) climate is an important factor for the diversity of wet grasslands; (2) Balkan vegetation of middle altitudes matches with that of Central Europe, whereas that of the lowest altitudes corresponds rather to the sub-Mediterranean region and high mountains are specific; (3) upward shift of Central-European vegetation types in southern Europe, so often described in forest vegetation is also evident for grassland vegetation and (4) the high diversity of Balkan vegetation is determined by a diverse relief enabling confluence of habitats possessing different climatic conditions.     

Variations in mountain vegetation use by reindeer (Rangifer tarandus) affects dry heath but not grass heath

Question: Are differences in landscape use of semi‐domesticated reindeer reflected in the vegetation of summer grazing grounds? Location: Alpine heaths, central east Sweden. Methods: Dry heath and grass heath vegetation plots with inferred grazing intensities (high, intermediate and low) were selected a priori from an interpolated pellet count map compiled in 2002. In each plot, faecal pellets were counted, environmental variables measured and vegetation sampled by listing presence and absence. Species composition was compared with a detrended correspondence analysis, and a canonical correspondence analysis was used to infer relations between species composition and environmental variables. Plots were also clustered to provide groupings for an indicator species analysis. Results: Significant differences in faecal pellet count were present between the highest and lowest grazing intensities for both vegetation types, showing that the pattern in the interpolated pellet maps was robust. Differences in species composition between grazing intensities were found for the dry heath only. Here, there was an apparent grazing gradient, with lichens and mosses in the low‐use plots and grasses and herbs in the high‐use plots. No such gradient was found for the grass heath. Conclusions: Within the dry heath vegetation type, grazing levels had a subtle effect on the vegetation, while no effects were seen in the grass heath, probably as a result of the dominance of more grazing‐tolerant graminoids. Even in the dry heath, species richness did not differ between grazing levels, but the relative abundances of species differed.