Fossil pollen as a record of past biodiversity

Quaternary pollen records may contribute uniquely to the understanding of present plant diversity. Pollen assemblages can reflect diversity at community and landscape scales but the time resolution of most studies does not match that of modern ecological studies. Because of the complicating effects of differential pollen productivity and dispersal, pollen records do not directly reflect equitability aspects of vegetation diversity. Vegetation diversity indices other than S (the total number of taxa) are therefore not appropriate for pollen assemblages. As a measure of the species richness palynological richness is biased by the lack of taxonomic precision, by a possible interference on pollen dispersal from vegetation structure and by pollen representation. The nonlinear relationship between species richness and pollen-taxa richness may be used in attempts to estimate past floristic richness from fossil pollen assemblages. Using a hypothetical example the strong effect of cover shifts in the vegetation affecting taxa with different representation (R_rel) values on observed palynological richness is demonstrated. It is suggested that estimates of relative pollen productivity should be used to guide the pollen sum on which pollen-type richness is estimated by rarefaction techniques and this approach is illustrated using a paired site study of late Holocene diversity dynamics. The need for a modern training set relating pollen-type richness to species richness, pollen productivity and vegetation structure is emphasized.     

Challenges in estimating past plant diversity from fossil pollen data: statistical assessment, problems, and possible solutions

Fossil pollen data from sediment cores may be used as a measure for past plant diversity. According to the theory of probability, palynological richness is positively related to the pollen count. In a low pollen count, only common taxa are detected, whereas rare taxa are only detected by chance. The detection of all pollen taxa requires a very high pollen count, which is time-consuming. In regular palynological investigations, the detected richness in pollen spectra varies with the pollen count. Rarefaction analysis estimates palynological richness in an exactly equal-sum count for all samples, so that comparison between samples is meaningful. However, the over-representation of some taxa suppresses the detection probability of rare taxa; low total pollen abundance in a sample enhances the detection probability of rare taxa and long-distance transported pollen grains. These factors bias the observed palynological richness and distort comparisons. Palynological richness in a pollen count proportional to its pollen influx may be one proxy for reconstructing diversity trends through time. The use of this proxy overcomes most problems encountered in rarefaction analysis, but is constrained by inaccuracy in estimating pollen influx due to the imprecise time control of sediment cores. Estimating palynological richness by mathematical methods may be another way of reconstructing pollen diversity. Pollen data tend to reflect diversity on a regional scale. Sites from small basins have the advantage of recording diversity at both local and regional scales, if the detection of each taxon is independent. By associating one site from a large basin with a series of sites from very small basins (e.g. forest-hollows), information about both regional and local diversity may be obtained. Entomophilous pollen taxa may have to be measured using a different strategy than anemophilous taxa.     

Modern pollen and stomate deposition in lake surface sediments from across the treeline on the Kola Peninsula, Russia

We sampled and analyzed surface sediments from 31 lakes along a latitudinal transect crossing the coniferous treeline on the Kola Peninsula, Russia. The major vegetation zones along the transect were tundra, birch-forest tundra, pine-forest tundra, and forest. The results indicate that the major vegetation types in our study area have distinct pollen spectra. Sum-of-squares cluster analysis and principal components analysis (PCA) groupings of pollen sites correspond to the major vegetation zones. PCA ordination of taxa indicates that the first axis separates taxa typical of the forest zone (Pinus, Picea) from taxa typical of tundra and forest-tundra zones (Polypodiaceae, Ericaceae, and Betula). The current position of the coniferous treeline, defined in our region by Pinus sylvestris, occurs roughly where Pinus pollen values reach 35% or greater. Arboreal pollen (AP)/non-arboreal pollen (NAP) ratios were calculated for each site and plotted against geographic distance along the transect. AP/NAP ratios of 7 or greater are found within pine-forest tundra and forest vegetation zones. Pinus stomates (dispersed stomatal guard cells) are absent from sites north of the coniferous treeline and all but two samples from the forested sites contain stomates. Stomate concentrations among the samples are highly variable and range from 10 to 458 per ml and positively correlate with the changing Pinus pollen values.     

Pollen-vegetation richness and diversity relationships in the tropics

Tracking changes in biodiversity through time requires an understanding of the relationship between modern diversity and how this diversity is preserved in the fossil record. Fossil pollen is one way in which past vegetation diversity can be reconstructed. However, there is limited understanding of modern pollen-vegetation diversity relationships from biodiverse tropical ecosystems. Here, pollen (palynological) richness and diversity (Hill N₁) are compared with vegetation richness and diversity from forest and savannah ecosystems in the New World and Old World tropics (Neotropics and Palaeotropics). Modern pollen data were obtained from artificial pollen traps deployed in 1-ha vegetation study plots from which vegetation inventories had been completed in Bolivia and Ghana. Pollen counts were obtained from 15 to 22 traps per plot, and aggregated pollen sums for each plot were >?2,500. The palynological richness/diversity values from the Neotropics were moist evergreen forest?=?86/6.8, semi-deciduous dry forest?=?111/21.9, wooded savannah?=?138/31.5, and from the Palaeotropics wet evergreen forest?=?144/28.3, semi-deciduous moist forest?=?104/4.4, forest-savannah transition?=?121/14.1; the corresponding vegetation richness/diversity was 100/36.7, 80/38.7 and 71/39.4 (Neotropics), and 101/54.8, 87/45.5 and 71/34.5 (Palaeotropics). No consistent relationship was found between palynological richness/diversity, and plot vegetation richness/diversity, due to the differential influence of other factors such as landscape diversity, pollination strategy, and pollen source area. Palynological richness exceeded vegetation richness, while pollen diversity was lower than vegetation diversity. The relatively high global diversity of tropical vegetation was found to be reflected in the pollen rain.     

Seasonality,habitat type and locality influenced bird assemblage structure in Nigeria

This is the first report of the avian assemblage in the study area of Dutse, Nigeria. In addition to recording bird species, the effects of season, dominant vegetation structure, locality and anthropogenic activities on bird abundance, species richness and diversity were investigated. Using the point transect method, 264 points on 48 km of transect were used to count birds between 06:30 and 11:00 from August 2015 to February 2016. A total of 122 bird species of 41 families were recorded. Highest bird species richness was recorded in Warwade, highest abundance in Model, and highest diversity in Malamawa. The dry season and woodland habitat showed higher bird species richness, abundance and diversity than the wet season and shrubland habitat. Tree density was more important in increasing bird abundance than shrub density. Small-scale anthropogenic activities and habitat modification, such as farming, grazing, wood removal and human interference did not appear to have impacted the birds; however, loss of high tree-density woodland habitats may pose a major threat to the bird community in Dutse. The presence of birds of concern in the area suggests the need for conservation efforts of avifauna and as well as the forested habitats in Dutse.     

Vegetation patterns,pollen deposition and distribution of non-pollen palynomorphs in an ice-wedge polygon near Kytalyk (NE Siberia), with some remarks on Arctic pollen morphology

In ice-wedge polygon mires, small-scaled microrelief of ridges enclosing small depressions results in a short-distance vegetation mosaic. The correct recognition of these landscape elements in palaeoecological studies of peat sections in order to reconstruct their patterns and dynamics requires insight in the short-distance relationship between vegetation and pollen deposition. This paper presents an analysis of pollen surface samples in a high-resolution (1 m) transect across an ice-wedge polygon near Kytalyk (NE Siberia), including a discussion on the morphology of some critical pollen types and non-pollen palynomorphs (NPPs). We found a strong correlation between vegetation and surface elevation and a fair correspondence between pollen deposition and vegetation. Distribution of NPPs reflects surface elevation well, with algal spores dominating deep spots and testate amoebae prevailing on higher spots. Peak pollen/spore values unrelated to high species coverages (e.g. of Salix, Betula, Sphagnum, Poaceae) indicate that single plants within a population may cause the bulk of the pollen production. The absence of pollen of taxa with an important presence in the vegetation (e.g. Utricularia) must be attributable to low pollen productivity. Distributional patterns point at pollen transport by water in the polygon troughs/depressions. Our study shows that Arctic pollen records mainly reflect short-distance vegetation patterns. Palaeosequences consequently allow accurate reconstruction of local microtopography and its dynamics, but should not be over-interpreted in terms of changing (over)regional vegetation patterns and associated drivers.     

Environmental implications of pollen spectra in bat droppings from southeastern Spain and potential for palaeoenvironmental reconstructions

Pollen was analysed from bat guano from nine caves in southeastern Spain and surface soils in their immediate surroundings. We compare the pollen spectra of 34 modern dung samples from the nine caves with one modern surface pollen sample from each cave. The contents suggest reasonable pollen diversity and richness, including anemophilous and zoophilous pollen types. Since the latter is usually under-represented in atmospheric pollen, the guano spectra therefore appear to reflect the vegetation more effectively than normal surface soil samples. Despite health hazards such as histoplasmosis, the difficulties of obtaining bat guano in deep caves and possible interpretational concerns relating to behaviour and feeding habits of different bat species, this material can be very useful in palaeoecological research provided that the dung was fossilized under favourable environmental conditions that allowed the preservation of pollen.     

Occurrence of pollen and spores in relation to present-day vegetation in a Dutch heathland area

Abstract. Pollen, spores and fungal remains in moss cushions along a transect, traversing a Betula-Quercus forest and a surrounding heathland, were analysed in order to study the relation between present-day vegetation and recent pollen deposition. Pollen and spores are divided into local types from plant species encountered along the transect and regional types, not present along the transect. Relative percentages and absolute concentration of the palynomorphs were estimated. Radiocaesium activity in the moss was measured to assess the minimal duration of palynomorph accumulation (i.e. the age of the moss samples). The absence of simultaneous trends in the regional pollen types indicates that the samples do not have major differences in age. The moss represents the later stages in the heath mosaic cycle. This is supported by the radiocaesium content of the moss. Thus, in this case the concentration values of regional pollen approach deposition values. The values of the local pollen types calculated on such a rather constant regional pollen flux is considered to represent real differences in the local deposition. The local pollen types can be arranged topographically into five groups characteristic of the local vegetation types and their ecotones. The arrangement of the curves of types from phyco-, zoo-, and myco-coenoses suggests their possible origin, either locally from the heath area or from the Betula-Quercus forest, or regionally from elsewhere. Analyses and topographical arrangement of the curves of unknown types contribute in this way to their identification and their possible source of origin.     

A survey of modern pollen and vegetation along a south–north transect in Mongolia

Aim This modern pollen‐rain study documents the spatial and quantitative relationships between modern pollen and vegetation in Mongolia, and explores the potential for using this relationship in palaeoclimatic reconstructions. Location East‐central Mongolia. Methods We collected 104 pollen surface samples along a south–north transect across five vegetation zones in Mongolia. Discriminant analysis was used to classify the modern pollen spectra into five pollen assemblages corresponding to the five vegetation zones. Hierarchical cluster analysis was used to divide the main pollen taxa into two major groups and seven subgroups representing the dry and moist vegetation types and the main vegetation communities within them. Results Each vegetation zone along the transect can be characterized by a distinctive modern pollen assemblage as follows: (1) desert zone: Chenopodiaceae–Zygophyllaceae–Nitraria–Poaceae pollen assemblage; (2) desert‐steppe zone: Poaceae–Chenopodiaceae pollen assemblage; (3) steppe zone: Artemisia–Aster‐type–Poaceae–Pinus Haploxylon‐type pollen assemblage; (4) forest‐steppe zone: Pinus Haploxylon‐type–Picea–Artemisia–Betula, montane forb/shrub and pteridophyte pollen assemblage; and (5) mountain taiga zone: Pinus Haploxylon‐type–Picea–Poaceae–Cyperaceae, montane forb/shrub and Pteridophyte pollen assemblage. Main conclusions Based on the ratio between the major pollen taxon groups and subgroups, we propose two pollen–climate indices that represent the precipitation and temperature conditions in the study region. When plotted along our south–north transect, the moisture indices (M) and temperature indices (T) mimic the regional gradients of precipitation and temperature across Mongolia very closely. These pollen–climate indices can be used for palaeoclimatic reconstruction based on fossil pollen data.     

Protostelids from tropical forests, woodlands and deserts in Australia

Most of what is known about the distribution of protostelids is limited to results from surveys carried out in North and Central America. To increase our knowledge about protostelid diversity and distribution we surveyed protostelids from 12 study sites in northern Queensland and the Northern Territory of Australia during May-Jun 2003. Aerial litter and ground litter samples were randomly collected along a 200 m transect at each site. Study sites ranged from tropical forests to deserts. We recovered 10 species and two apparently undescribed species from samples of aerial (dead but still attached plant parts) and ground litter. Samples from a woodland site characterized by intermediate moisture conditions had the greatest species richness, followed by samples from dry woodland, tropical forest and desert sites. When species richness for a particular microhabitat was considered, samples of aerial litter yielded more species than samples of ground litter. Percentages of samples colonized with protostelids were similar for the aerial and ground litter microhabitats within a given habitat type except for dry woodlands, in which aerial litter samples were characterized by higher numbers of species than ground litter samples. Two species (Protostelium mycophaga and Soliformovum irregularis) that in temperate North America are associated with aerial litter microhabitats also were recovered from aerial litter in dry habitats in Australia. Schizoplasmodiopsis pseudoendospora, a North American temperate ground litter species, was equally abundant in aerial litter and ground litter in Australia. This study is the first of its kind for protostelid ecology in Australia and the most extensive study of protostelids in the southern hemisphere. These data complement ongoing research on protostelid distribution from around the world.     

Mid- and late-Holocene vegetation history of Quidenham Mere,Norfolk, UK interpreted using recurrent groups of taxa

The results of pollen analyses from the top 8 m of sediment retrieved from Quidenham Mere, Norfolk, UK are presented. These, together with the results of charcoal analyses and the numerical classification of the pollen and spore taxa into ‘recurrent groups’ using simple discriminant function analysis, are used to reconstruct the vegetation history of the Quidenham Mere catchment over approximately the last 8000 years and to infer anthropogenic effects on the landscape. After some minor woodland clearance in the Bronze Age and further clearance in the Iron Age, it was only in the Anglo-Saxon period that most of the woodland was cleared. Although arable farming was important, it appears that there was also a considerable amount of land used for grazing, with grazed woodland and wood pasture. Towards the top of the sequence, within the past 200 years, there is evidence of the foundation and development of the Quidenham Hall parkland adjacent to the site, with the planting of native and exotic species of trees. A comparison is made between the vegetation histories and anthropogenic effects on the landscapes of Quidenham Mere and Diss Mere, a similar lake about 10 km from Quidenham. Both sites reflect the agricultural practices occurring in their catchments from early times. Exceptionally high values of Cannabaceae grains are indicative of retting of hemp during the past 1500 years. Rarefaction analysis is used to compare the pollen richness of the two sites.     

Spatial patterns of plant diversity and communities in Alpine ecosystems of the Hengduan Mountains,Northwest Yunnan,China

Aims Conduct a quantitative, but rapid, regional-level assessment of the alpine flora across northwest Yunnan (NWY) to provide a broad-based understanding of local and regional patterns of the composition, diversity and health of alpine ecosystems across NWY.Methods A stratified random sampling design was employed to select sites across the different mountain ranges of NWY. Vegetation was sampled by stratifying each site by the three major alpine vegetation community types: meadow, dwarf shrub and scree. Two 50-m transects were randomly located within each community type at each sampling site with 10 1-m 2 subplots systematically placed along each transect. Environmental variables were recorded at each transect. Multivariate analyses were used to classify the major plant community assemblages and link community patterns to environmental and habitat variables.Important findings Forb species richness varied from 19 to 105 species per site (21 sites total) with an average of 59 species per site (60 m 2 sampled per site). Most species were patchily distributed with narrow distributions and/or small population sizes; over half the species occurred at only one or two sites. Distinct species assemblages were identified in the meadow vegetation that was strongly aggregated by geographic location suggesting the presence of distinct phytogeographic zones of the meadow alpine flora. Elevation and geographic location were the dominant environmental gradients underlying the variations in species composition. Jaccard's coefficient of similarity averaged only 10% among sites indicating there was little similarity in the alpine flora across the region. The alpine vegetation is highly heterogeneous across the complex landscape of the Hengduan Mountains of NWY. Conservation strategies need to take into account the large geographic differences in the flora to maximize protection of biodiversity.     

The distribution and diversity of Chironomidae (Insecta: Diptera) in western Finnish Lapland, with special emphasis on shallow lakes

Aim Predictions of aquatic ecosystem change with global warming require basic data that accurately reflect the environmental conditions underlying species distributions. However, in remote arctic areas such baseline data are scarce. We assess the influence of environmental variables on chironomid distribution and taxon richness in shallow, isothermal lakes in a poorly studied arctic region. We pay particular attention to community variation along the treeline ecotonal zone where many environmental variables change abruptly in a relatively small area. Location Lake transect in Finnish Lapland spanning from boreal coniferous forest to arctic tundra. Methods Chironomid assemblages were determined from surface‐sediment samples of 50 shallow (< 10 m) natural lakes. Abundance and taxon richness data were related to 24 limnological variables using canonical ordination techniques (DCA, CCA, RDA). A Monte Carlo permutation procedure was used to assess the explanatory power of single variables. Between‐vegetation zone differences of richness were tested for statistical significance using one‐way anova . Results In total, 7771 chironomid head capsules were identified, consisting of 13 species, 10 species groups, four subgenera, 41 genera, four genus groups, five types and three with uncertain taxonomic affiliation. A hump‐shaped relationship between taxon richness and elevation was noted along the study transect with a peak in taxon richness occurring in mountain birch woodland lakes at middle elevations, decreasing then towards both warmer and colder ends of the elevation/temperature gradient. Of the individual parameters, sediment organic content, total organic carbon, pH, and lake‐specific air temperature accounted for the greatest amount of variation in the chironomid data. Main conclusions Maximum taxon richness occurred at mid‐elevations where aquatic algae also reached their maximum diversity. This area coincides with an ecotonal transitional zone, which seems more likely to account for the peak in species richness. Our study demonstrates that the factors most strongly affecting chironomids in Finnish Lapland (i.e. temperature, and ecosystem features) are those that with great probability will also change as a result of future climate change. This will likely have an effect on the distribution of chironomids in subarctic and arctic areas.     

Species–area relationships in the Hantam-Tanqua-Roggeveld,Succulent Karoo,South Africa

The Hantam-Tanqua-Roggeveld subregion is part of the Succulent Karoo hotspot of biodiversity which stretches along the southwestern side of South Africa and Namibia. Forty Whittaker plots were surveyed in the spring of 2005, in eight vegetation associations, to gather diversity data for the Hantam, Tanqua Karoo and Roggeveld areas. Seven plot sizes were used to construct species–area curves using three different models namely: the untransformed linear function, the power function and the exponential function. In general, the power and exponential functions produced a more significant fit to the data than the untransformed linear function. Linear regressions using environmental parameters indicated that altitude, mean annual precipitation and mean annual temperature were significant predictors of species richness at the 1, 10, 100 and 1000 m² scales. To illustrate the variation in species–area curves and species richness across the landscape, a transect through the study area is discussed. The transect stretches eastwards from the Tanqua Karoo across the escarpment into the Roggeveld and crosses five different vegetation associations. Differences between associations were found in species richness in the 1000 m² plots. Each association also produced species–area curves with their own characteristics. Slope values for the samples within an association did not differ significantly, although the intercept value often did. Comparisons between associations along the transect revealed significant differences in the slope value between the associations, except for the Dicerothamnus rhinocerotis Mountain Renosterveld which did not differ significantly from the associations bordering it on either side.     

Water availability drives aboveground biomass and bird richness in forest restoration plantings to achieve carbon and biodiversity cobenefits

To combat global warming and biodiversity loss, we require effective forest restoration that encourages recovery of species diversity and ecosystem function to deliver essential ecosystem services, such as biomass accumulation. Further, understanding how and where to undertake restoration to achieve carbon sequestration and biodiversity conservation would provide an opportunity to finance ecosystem restoration under carbon markets. We surveyed 30 native mixed‐species plantings in subtropical forests and woodlands in Australia and used structural equation modeling to determine vegetation, soil, and climate variables most likely driving aboveground biomass accrual and bird richness and investigate the relationships between plant diversity, aboveground biomass accrual, and bird diversity. We focussed on woodland and forest‐dependent birds, and functional groups at risk of decline (insectivorous, understorey‐nesting, and small‐bodied birds). We found that mean moisture availability strongly limits aboveground biomass accrual and bird richness in restoration plantings, indicating potential synergies in choosing sites for carbon and biodiversity purposes. Counter to theory, woody plant richness was a poor direct predictor of aboveground biomass accrual, but was indirectly related via significant, positive effects of stand density. We also found no direct relationship between aboveground biomass accrual and bird richness, likely because of the strong effects of moisture availability on both variables. Instead, moisture availability and patch size strongly and positively influenced the richness of woodland and forest‐dependent birds. For understorey‐nesting birds, however, shrub cover and patch size predicted richness. Stand age or area of native vegetation surrounding the patch did not influence bird richness. Our results suggest that in subtropical biomes, planting larger patches to higher densities, ideally using a diversity of trees and shrubs (characteristics of ecological plantings) in more mesic locations will enhance the provision of carbon and biodiversity cobenefits. Further, ecological plantings will aid the rapid recovery of woodland and forest bird richness, with comparable aboveground biomass accrual to less diverse forestry plantations.     

Do spider diversity and assemblages change in different contiguous habitats? A case study in the protected habitats of the humid chaco ecoregion, northeast Argentina

Spiders are a megadiverse group that can be useful indicators of the overall species richness and health of biotic communities. The spider diversity in subtropical forests of the Neotropical region are not yet well known, especially in Argentinean subtropical forests where systematic fieldwork has not been done until recently. The Great Chaco is very important as the unique dry subtropical forest of the earth, but it is suffering increasing degradation by the advance of agriculture. Spider communities have been shown to be more directly influenced by vegetation architecture than vegetation species composition. In this study, we aim to assess whether spider diversity and assemblages change in adjacent habitats with different types of vegetation. We compare the diversity and spider assemblages in two different contiguous protected habitats (hygrophilous woodland and savannah parkland) of the Mburucuyá National Park, (Humid Chaco ecoregion). Seasonal samples were obtained using three types of sampling methods: pitfall trapping, beating, and manual litter extraction. The spider assemblages were different in the studied areas, and the abundance, diversity, evenness, and species richness were higher in the hygrophilous woodland than the savannah parkland. These differences in spider diversity and assemblages indicate that both types of habitats are important if the biodiversity is to be conserved in the Chaco ecoregion, where different types of habitat are shown as a patchy distribution.     

Numerical analysis of vegetation complexes and community diversity of major coastal dinaric mountains

Abstract. This study includes 24 transects on slopes of the Velebit, Biokovo and Orjen mountains, the tallest coastal mountains along the Adriatic Sea, with a very high species richness according to European standards and having the largest regional concentrations of endemics. Within each transect the presence or absence of phytosociological alliances, according to the Zürich-Montpellier School, was interpreted from local studies. The alliance composition of the transects was compared with methods used in numerical syntaxonomy. Three types of vegetation complexes (N. Dinaric inland, transitional and S. Dinaric coastal) were derived from a numerical classification of the transects; they were defined on the basis of their characteristic alliances. The maritime impact affecting the mountains through the river canyons reduces the contrast in vegetation between inland and coastal slopes. The existence of a climatic macrogradient is indicated by seriation of the basic alliance/transect data matrix leading to a diagonal structure, by a gradual shift of the alliance composition within the same vegetational type, and by the distribution of the transects and alliances along the first axis of a Correspondence Analysis ordination. Transect scores on the second ordination axis are negatively correlated with transect vegetational diversity (number of alliances per transect); this axis may reflect a decrease in humidity and temperature range. This interpretation is supported by the result of a comparison of the total mountain vegetational diversity (Mann-Whitney test), which showed that Orjen is the most diverse mountain ridge. The expected correlation between plant community diversity and altitude of the transects is blurred, probably due to the impact of the cold Bora wind and subsequent compression of vegetational zones.     

Pollen-rain-vegetation relationships along a forest-savanna transect in southeastern Cameroon

Modern soil and litter samples from southeastern Cameroon, collected along a continuous forest–savanna transect were analysed for pollen content to define modern pollen–vegetation relationships. The pollen results, completed and compared with botanical inventories, leaf area index and basal area measurements performed in the same area, clearly registered the physiognomy, the main floristic composition and floral richness of the two sampled ecosystems. Distortions were observed between sampled vegetations and their pollen rain, related to important differences in pollen production and dispersal of plant species: this is a general feature in many tropical regions. The pollen data in the area studied reflected well the recent transgression of forest versus savanna. This permitted us to define inside the forest ecosystem more successional vegetation communities than the botanical surveys allowed.     

Palynological richness and evenness: insights from the taxa accumulation curve

Palynology provides the opportunity to make inferences on changes in diversity of terrestrial vegetation over long time scales. The often coarse taxonomic level achievable in pollen analysis, differences in pollen production and dispersal, and the lack of pollen source boundaries hamper the application of diversity indices to palynology. Palynological richness, the number of pollen types at a constant pollen count, is the most robust and widely used diversity indicator for pollen data. However, this index is also influenced by the abundance distribution of pollen types in sediments. In particular, where the index is calculated by rarefaction analysis, information on taxonomic richness at low abundance may be lost. Here we explore information that can be extracted from the accumulation of taxa over consecutive samples. The log-transformed taxa accumulation curve can be broken up into linear sections with different slope and intersect parameters, describing the accumulation of new taxa within the section. The breaking points may indicate changes in the species pool or in the abundance of high versus low pollen producers. Testing this concept on three pollen diagrams from different landscapes, we find that the break points in the taxa accumulation curves provide convenient zones for identifying changes in richness and evenness. The linear regressions over consecutive samples can be used to inter- and extrapolate to low or extremely high pollen counts, indicating evenness and richness in taxonomic composition within these zones. An evenness indicator, based on the rank-order-abundance is used to assist in the evaluation of the results and the interpretation of the fossil records. Two central European pollen diagrams show major changes in the taxa accumulation curves for the Lateglacial period and the time of human induced land-use changes, while they do not indicate strong changes in the species pool with the onset of the Holocene. In contrast, a central Swedish pollen diagram shows comparatively little change, but high richness during the early Holocene forest establishment. Evenness and palynological richness are related for most periods in the three diagrams, however, sections before forest establishment and after forest clearance show high evenness, which is not necessarily accompanied by high palynological richness, encouraging efforts to separate the two.     

Effects of three flower field types on bumblebees and their pollen diets

Agri-environment schemes, like flower fields, have been implemented in the EU to counteract the dramatic decline of farmland biodiversity. Farmers in Lower Saxony, Germany, may receive payments for three flower field types: annual, perennial (five years old), and mixed flower fields composed of yearly alternating annual and biannual parts. We assessed the effectiveness of these flower field types in providing bumblebee foraging habitat compared to control cereal fields. We sampled bumblebees with transect walks and assessed the richness of exploited pollen plants using DNA meta-barcoding and direct observations.All flower field types enhanced bumblebee abundance and species richness compared to control fields but attracted mostly three generalist species. Although we expected highest benefits from the more heterogeneous mixed flower fields, abundance was highest in annual, only intermediate in mixed, and lowest in perennial flower fields. Bumblebee species richness did not differ between flower field types.Overall, the proportion of sown plants in pollen loads was surprisingly low (< 50%). Bombus pascuorum, but not B. terrestris agg., exploited 10% of the sown plant species in perennial, 36% in annual and 45% in mixed flower fields, respectively. Compared to direct observations, pollen samples revealed 4.5 times more visited plant species and thus assessed floral resource use more reliably. Plant species richness in pollen loads decreased with local flowering plant species richness and increased with proportion of annual crops in the landscape, potentially due to the exploitation of more diverse and scattered resources, including flowering crops, in homogenized landscapes to fulfil dietary requirements.Our results indicate that under the current management, both annual and mixed flower fields provide the most attractive food resources, while perennial flower fields offered the poorest foraging habitats. Conclusively, flower fields seem important but resources from the surrounding landscape are still needed to sustain bumblebees in agricultural landscapes.