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.     

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.     

The role of landscape structure in determining palynological and floristic richness

The associations between floristic and palynological richness and landscape structure were studied based on modern pollen?Cvegetation data from a patchy cultural landscape in southern Estonia (northern temperate vegetation zone). Nine study sites (small lakes and their surrounding vegetation) represent land cover gradient from closed forest to semi-open vegetation. Floristic richness (number of species) and floristic richness of pollen types (number of pollen-equivalent taxa) were used to describe the vegetation within the radius of 250?m from the pollen sampling sites. Palynological richness was calculated to describe the modern pollen samples diversity. Landscape structure was estimated on the basis of landscape openness and three landscape diversity measures: richness of community patches, Simpson evenness of community patches and Simpson diversity of community patches. To study the effect of the spatial scale of landscapes on the vegetation?Clandscape and pollen?Clandscape associations, landscape structure was estimated within eight radii (250?C2,000?m) around each lake. The results showed that landscape openness was the most important determinant of both floristic richness and palynological richness in southern Estonia and that landscape diversity estimated by Simpson diversity index was also significantly associated with the richness estimates. Floristic and palynological richness were significantly positively correlated with landscape structure within the radii greater than 1,000?m from the pollen sampling sites, which is similar to the estimated Relevant Source Area of Pollen in southern Estonia. We conclude that within one floristic or climatic region, palynological richness gives reliable estimates about the variation in floristic richness and landscape structure; however, caution must be taken when comparing pollen-inferred vegetation diversities from different regions or when interpreting fossil pollen records from times with highly different vegetation associations.     

Exploring Holocene Changes in Palynological Richness in Northern Europe – Did Postglacial Immigration Matter?

In mid to high latitudes glacial and interglacial cycles have repeatedly changed the area available for plant growth. The speed at which plants are able to colonize areas at the onset of an interglacial is hypothesized to limit their distribution ranges even today (migrational lag). If the spread of plants would have been generally slow then plant diversity in previously glaciated areas would be expected to increase over time. We explore this hypothesis using results from six palynological investigations from two previously glaciated regions: central Sweden and north-eastern Germany. Rarefaction, slope of rank order abundance, and taxa accumulation plots were used to evaluate richness and evenness in pollen data in an attempt to separate richness from evenness. These analyses show little change in palynological richness for the northern sites throughout the Holocene. In contrast, the southern sites show an increase in richness and evenness during the early Holocene; this may be explained by the different initial conditions at the onset of the Holocene. A strong rise in palynological richness around 6000 and 1000 years ago at the southern sites can be attributed to the regional initiation of agriculture and major opening of the forest, respectively. For the northern sites there is no evidence for increased taxonomic diversity through time that could be due to delayed immigration of species.     

Biodiversity and pollen analysis: modern pollen studies and the recent history of a floodplain woodland in S. W. Ireland

One of the problems with biodiversity for palynologists is that their samples come from an unbounded area and source area varies with both pollen/spore type and vegetation type. There have been two broad approaches to the problem of inferring diversity from pollen/spore samples; firstly the use of pollen type richness as a proxy for species richness, and secondly the ‘identification’ of past vegetation communities aided by ecological inference and historical or modern data concerning species affinities and typical diversity. The estimation of biodiversity from pollen analysis depends upon the pollen count, taxonomic precision, source strength for individual types and dispersal/transport of pollen from source areas to the site. A transect of surface pollen samples is used here to test the effect of a vegetation boundary on pollen/spore diversity and compare the pollen/spore diversity with the species richness of the woodland. Palynological richness (at a constant count sum) does, to some extent, reflect changing local vegetation along the transect. Type count curves also reflect the changes in vegetation type along the transect due to the partial influence of species richness on pollen/spore richness. This study suggests that depending upon the woodland composition, the woodland may not entirely drown-out the pollen signal of the surrounding vegetation, and that a small woodland may be more representative (as a sampling site) of the valley floor diversity than a raised mire. The surface transect is also used in the interpretation of Medieval pollen levels from The Gearagh.     

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.     

Floristic diversity in the transition from traditional to modern land-use in southern Sweden a.d. 1800–2008

We aim to provide a long-term ecological analysis of land-use and floristic diversity in the transition from traditional to modern land-use management in the time a.d. 1800–2008 in southern Sweden. We use the Regional Estimates of Vegetation Abundance from Large Sites (REVEALS) model to quantify land-cover changes on a regional scale at 20-year intervals, based on the fossil pollen record. Floristic richness and evenness are estimated using palynological richness and the Shannon index applied to the REVEALS output, respectively. We identified a transition period of 60?years between 1880 and 1940 when the total tree cover increased and the tree composition changed from deciduous to coniferous dominance. Within the shrinking area of open land, arable land taxa expanded, while the number and coverage of herbs in the remaining grasslands decreased. The succession from open grasslands to more tree-covered habitats initially favoured palynological richness, which reached its highest values during the first 40?years of the transition period. The highest REVEALS-based evenness was recorded in the time of traditional land-use and at the beginning of the transition period, reflecting higher habitat diversity at these time intervals. Our results support a more dynamic ecosystem management that changes between traditional land-use and phases of succession (<40?years) to promote floristic diversity. We have developed and applied a palaeoecological methodology that contributes realistic estimates to be used in ecosystem management.     

A deep dig––hindsight on Holocene vegetation composition from ancient environmental DNA

Want a glimpse at past vegetation? Studying pollen and other plant remains, which are preserved for example in lake sediments or mires for thousands of years, allows us to document regional occurrences of plant species over radiocarbon‐dated time series. Such vegetation reconstructions derived from optical analyses of fossil samples are inherently incomplete because they only comprise taxa that contribute sufficient amounts of pollen, spores, macrofossil or other evidences. To complement optical analyses for paleoecological inference, molecular markers applied to ancient DNA (aDNA) may help in disclosing information hitherto inaccessible to biologists. Parducci et al. (2013) targeted aDNA from sediment cores of two lakes in the Scandes Mountains with generic primers in a meta‐barcoding approach. When compared to palynological records from the same cores, respective taxon lists show remarkable differences in their compositions, but also in quantitative representation and in taxonomic resolution similar to a previous study (Jørgensen et al. 2012). While not free of assumptions that need critical and robust testing, notably the question of possible contamination, this study provides thrilling prospects to improve our knowledge about past vegetation composition, but also other organismic groups, stored as a biological treasure in the ground.     

Test of an equal taxon–weight modification of Middeldorp's pollen density dating on data from varved sediments of Lake Gościąż, Poland

The cores from the varved sediments of Lake Gościąż enable testing of methods, based on pollen concentration, to reconstruct the time-basis of a palynological data set. The sediments' varves or annual laminae were counted and their thicknesses measured, and the pollen content was recorded at known intervals with regard to depth and time. In the present study, we have treated this pollen record as if the sediments were not annually laminated and as if, consequently, the time-basis of the pollen record was unknown. We reconstructed several different time-bases by means of two methods using pollen concentration, namely pollen density dating (PDD) according to Middeldorp and a modification of that method, which gives all included arboreal pollen-taxa the same weight: equal taxon–weight modification (ETW). After reconstructing these time-bases we compared them with the varve time-basis. We conclude that both PDD and ETW result in good time-bases. Our modification improves the performance of Middeldorp's method, largely because of the use of spline interpolation instead of a polynomial regression of varve age on pollen variables. An advantage of ETW over PDD is that ETW using a spline produces a good approximation of arboreal influx, preserving both trends and short-term variability, whereas PDD results in an arboreal influx being an unnaturally smooth function of time.     

The Holocene vegetation history of northern West Jutland, Denmark

Holocene sediments of three closed Danish lake basins (Solsø, Skånsø, Kragsø) were used for the inference of post-glacial vegetational dynamics in former heathland areas in northern West Jutland, Denmark. The sites were selected to represent the major geomorphological units of West Jutland. The Holocene history of each lake basin was investigated by mapping of sediment distribution, analysis of loss-on-ignition, coarse inorganic matter, humus content, mineral magnetics, δ¹³C, pollen and selected other microfossils. These techniques were supplemented by plant macrofossil analysis at one site. Holocene terrestrial vegetational development was inferred at each site from analyses of pollen and microscopical charred particles. Chronologies were provided by numerous ¹⁴C dates. Stratigraphies of wet ground and terrestrial pollen and spore types were zooned by stratigraphically constrained cluster analysis. Based on the resultant site pollen asemblage zones (site PAZ), regional PAZ were proposed. Using modern analogues, Holocene floristic richness was estimated from pollen richness in the microfossil assemblages. The results support the hypothesis that disturbance is one of the most important mechanisms behind the maintenance of floristic richness. In particular, the response of estimated floristic richness to the intensity of vegetational fires followed the predictions of the Intermediate Disturbance Hypothesis. A period of elevated palynological richness and inferred vegetational disturbance was identified at all sites between 6000 and 5200 BC (calendar years). Using correspondence analysis (CA), the major gradient in the terrestrial pollen sequences was identified as a light-shade gradient, and CA first axis sample scores were used as a supplement to standard AP/NAP pollen ratios as an indicator of the shade-tolerance/light-demand of Holocene terrestrial plant communities. In spite of different vegetational developments since 4000 BC, the timing of major changes towards more light-demanding vegetation types were broadly synchronous at the three sites. Using chord distance as a dissimilarity index, rates of palynological change suggest that the interval between 8000 and 7500 BC (calendar years) was the period of most rapid vegetational change during the Holocene, both in terrestrial as well as lacustrine ecosystems. While climatic forcing of the rapid events around 8000 BC is hypothesised, the synchronous timing of relatively rapid inferred change in lake and terrestrial vegetation around AD 600 may reflect changes in climate as well as in land-use. Reducdancy analysis was used to develop a model between fire intensity (inferred from microscopical charred particles) and vegetational response, as reflected by pollen assemblages. Formulated at one site and tested at the two other sites, the model explains regional Calluna-heathland expansions as a result of vegetational burning. Similarly, declines in heathland cover are explained by lack of maintenance by fire. Regional vegetational development in northern West Jutland is reconstructed and special consideration is given to heathland history. The Holocene heathland development is interpreted as resulting from its importance for grazing. It is hypothesized that on poor soils, Calluna-dominated heathland was a better grazing resource than grass-dominated pasture, due to the winter-grazing offered by Calluna and the low palatibility of dominant grasses on poor soils. This hypothesis is relevant for the explanation of the variation in timing of heathland expansions on the different soil types represented by the study sites.     

Diverse invertebrate fauna using dry sediment as a refuge in semi-arid and temperate Australian rivers

Dormant aquatic invertebrates can remain viable in riverbed sediment during dry phases, forming a source for recolonisation during wet periods. Regional differences in capacity for invertebrates to survive drying in this way are poorly understood, but may indicate regional differences in vulnerability to altered flow regimes. We compared diversity of invertebrates in dry sediment from intermittent rivers in temperate and semi-arid Australia after 4–8 weeks of drying. We predicted adaptations of semi-arid biota to severe and unpredictable drying would make dry sediment a more significant recolonisation source, with higher relative diversity when compared with temperate rivers. Emerging aquatic invertebrate assemblages were compared to those sampled in nearby pools, as a common drying refuge. Relative taxa richness in rehydrated sediments was higher in the semi-arid region (83 ± 16% of pool taxa) than the temperate (47 ± 6% of pool taxa), despite lower overall richness (24 taxa in semi-arid, 32 taxa in temperate). Semi-arid rivers had greater potential for dry riverbeds to act as a source for recolonisation, given high relative diversity and abundance in dry sediment, combined with the frequent absence of alternative refuges. However, dry riverbeds in both regions provided a significant short-term refuge for aquatic invertebrates.     

The Holocene vegetation history of northern West Jutland, Denmark

Holocene sediments of three closed Danish lake basins (Solsø, Skånsø, Kragsø) were used for the inference of post-glacial vegetational dynamics in former heathland areas in northern West Jutland, Denmark. The sites were selected to represent the major geomorphological units of West Jutland. The Holocene history of each lake basin was investigated by mapping of sediment distribution, analysis of loss-on-ignition, coarse inorganic matter, humus content, mineral magnetics, δ¹³C, pollen and selected other microfossils. These techniques were supplemented by plant macrofossil analysis at one site. Holocene terrestrial vegetational development was inferred at each site from analyses of pollen and microscopical charred particles. Chronologies were provided by numerous ¹⁴C dates. Stratigraphies of wet ground and terrestrial pollen and spore types were zooned by stratigraphically constrained cluster analysis. Based on the resultant site pollen asemblage zones (site PAZ), regional PAZ were proposed. Using modern analogues, Holocene floristic richness was estimated from pollen richness in the microfossil assemblages. The results support the hypothesis that disturbance is one of the most important mechanisms behind the maintenance of floristic richness. In particular, the response of estimated floristic richness to the intensity of vegetational fires followed the predictions of the Intermediate Disturbance Hypothesis. A period of elevated palynological richness and inferred vegetational disturbance was identified at all sites between 6000 and 5200 BC (calendar years). Using correspondence analysis (CA), the major gradient in the terrestrial pollen sequences was identified as a light-shade gradient, and CA first axis sample scores were used as a supplement to standard AP/NAP pollen ratios as an indicator of the shade-tolerance/light-demand of Holocene terrestrial plant communities. In spite of different vegetational developments since 4000 BC, the timing of major changes towards more light-demanding vegetation types were broadly synchronous at the three sites. Using chord distance as a dissimilarity index, rales of palynological change suggest that the interval between 8000 and 7500 BC (calendar years) was the period of most rapid vegetational change during the Holocene, both in terrestrial as well as lacustrine ecosystems. While climatic forcing of the rapid events around 8000 BC is hypothesised, the synchronous timing of relatively rapid inferred change in lake and terrestrial vegetation around AD 600 may reflect changes in climate as well as in land-use. Reducdancy analysis was used to develop a model between fire intensity (inferred from microscopical charred particles) and vegetational response, as reflected by pollen assemblages. Formulated at one site and tested at the two other sites, the model explains regional Colluna-heathland expansions as a result of vegetational burning. Similarly, declines in heathland cover are explained by lack of maintenance by fire. Regional vegetational development in northern West Jutland is reconstructed and special consideration is given to heathland history. The Holocene heathland development is interpreted as resulting from its importance for grazing. It is hypothesized that on poor soils, Calluna-dominated heathland was a better grazing resource than grass-dominated pasture, due to the winter-grazing offered by Calluna and the low palatibility of dominant grasses on poor soils. This hypothesis is relevant for the explanation of the variation in timing of heathland expansions on the different soil types represented by the study sites.     

Species turnover in lake littorals: spatial and temporal variation of benthic macroinvertebrate diversity and community composition

Aims Despite wide consensus that ecological patterns and processes should be studied at multiple spatial scales, the temporal component of diversity variation has remained poorly examined. Specifically, rare species may exhibit patterns of diversity variation profoundly different from those of dominant taxa. Location Southern Finland. Methods We used multiplicative partitioning of true diversities (species richness, Shannon diversity) to identify the most important scale(s) of variation of benthic macroinvertebrate communities across several hierarchical scales, from individual samples to multiple littorals, lakes and years. We also assessed the among‐scale variability of benthic macroinvertebrate community composition by using measures of between‐ and within‐group distances at hierarchical scales. Results On average, a single benthic sample contained 23% of the total regional macroinvertebrate species pool. For both species richness and Shannon diversity, beta‐diversity was clearly the major component of regional diversity, with within‐littoral beta‐diversity (β₁) being the largest component of gamma‐diversity. The interannual component of total diversity was small, being almost negligible for Shannon index. Among‐sample (within‐littoral) diversity was related to variation of substratum heterogeneity at the same scale. By contrast, only a small proportion of rare taxa was found in an average benthic sample. Thus, dominant species among lakes and years were about the same, whereas rare species were mostly detected in a few benthic samples in one lake (or year). For rare species, the temporal component of diversity was more important than spatial turnover at most scales. Main conclusions While individual species occurrences and abundances, particularly those of rare taxa, may vary strongly through space and time, patterns of dominance in lake littoral benthic communities are highly predictable. Consequently, many rare species will be missed in temporally restricted samples of lake littorals. In comprehensive biodiversity surveys, interannual sampling of littoral macroinvertebrate communities is therefore needed.     

Les palynomorphes du Bajocien supérieur : un premier PALYNO-DATA pour les Monts de Rhar Roubane (Algérie nord occidentale)

Apart from the few works carried out in the North Tethyan basins, the palynological study of Mesozoic outcrops is still rare. Thus, the present study devoted to the Zahra Marls Formation (upper Bajocian) of the Rhar Roubane Mountains (northwestern Algeria), allowed us for the first time to inventory 44 different taxa. The palynological assemblages studied are strongly dominated by trilete spores (17 genera and 26 species), the most abundant of which are Concavissimisporites variverrucatus, Cyathidites australis and Cyathidites minor. Pollen grains are relatively frequent (10 genera and 16 species), while acritarchs and algae are represented by only two taxa (Micrhystridium sp. and Botryococcus sp.). In addition, the sporopollinic spectra dominated by the terrestrial elements are represented mainly by the spores of Pteridophyta and Bryophyta and gymnosperm pollen.     

A taphonomic study of modern pollen assemblages from dung and surface sediments in arid environments of Spain

Coastal plant communities in arid southeastern Spain are characterized by insect-pollinated scrub species, which fail to occur in Quaternary pollen sequences from valleys, marshlands and marine cores. We investigate pollen–vegetation relationships in samples from soil surfaces, animal dung, and sediments in depressions or basins that, in theory, should have pollen spectra that are comparable to those from sedimentary basins elsewhere. Pollen spectra from basins or depressions are very susceptible to long-distance wind and water transport. This can mask representation of pollen from the surrounding insect-pollinated vegetation, as can over-representation of basin-margin halophilous and hydrophilous pollen. Pollen spectra from biogenic materials of animal origin are the best analogues of local and regional vegetation, and show the best analytical potential in terms of pollen concentration and taxon diversity. Pollination properties of the species studied indicate they will rarely be found in most conventional pollen records. It cannot be stressed too strongly that insight into Quaternary vegetation of arid regions demands complementary pollen analysis of coprolites, urine-cemented deposits, and cave sediments with preserved biotic remains, in addition to water-lain sediments.     

The relationship between local and regional diversity of indigenous forest fauna in KwaZulu-Natal Province, South Africa

The relationship between local and regional diversity was tested by regressing local community richness against regional species diversity for three taxa, birds, butterflies and mammals, in subtropical forest. The quadratic model best fits the relationship between local and regional diversity for birds. Local bird species richness is theoretically independent of the size of the regional pool of species and may represent saturated communities. A linear model best describes the relationship for mammals and butterflies. For mammals, the slope is shallow (0.264) and regional richness overestimates local species richness, suggesting communities are undersaturated. Extinction filtering may explain this pattern. Past climatic changes have filtered out many mammalian species, these changes have been too recent for autochthanous speciation, and the relatively low vagility of mammals has prevented extensive recolonisation. Differences in the nature of the diversity relationship between taxa are as much due to independent evolutionary histories as to differences in vagility and colonising potential. A pervasive role is suggested for regional biogeographic processes in the development of faunal assemblage structure. Large-scale processes are not considered in current conservation plans. We encourage the shift of conservation emphasis from local ecological processes and species interactions, to whole communities and consideration of regional processes.     

Holocene tree‐line fluctuations and climate in central troms,Northern Norway

Based mainly on pollen influx data, an attempt is made at reconstructing Holocene tree‐line fluctuations and palaeotemperatures in central Troms, North Norway. Both past and present‐day (Tauber trap) pollen influx data suggest that influx rates exceeding 250–300 pollen/cm²/year for each of the major arboreal taxa (Betula pubescens and Pinus sylvestris) are only found at sites with corresponding forest types in the immediate surroundings. High influx rates, raised tree‐lines and favourable climatic periods are recorded at 7500–4500 and 3000–2600 B.P. Betula and Pinus woodlands may have reached more than 200 m above their present altitude limits, suggesting a July mean temperature 2°C higher than at present during the Holocene optimum.     

Assessing Paleo-Biodiversity Using Low Proxy Influx

We developed an algorithm to improve richness assessment based on paleoecological series, considering sample features such as their temporal resolutions or their volumes. Our new method can be applied to both high- and low-count size proxies, i.e. pollen and plant macroremain records, respectively. While pollen generally abounds in sediments, plant macroremains are generally rare, thus leading to difficulties to compute paleo-biodiversity indices. Our approach uses resampled macroremain influxes that enable the computation of the rarefaction index for the low influx records. The raw counts are resampled to a constant resolution and sample volume by interpolating initial sample ages at a constant time interval using the age∼depth model. Then, the contribution of initial counts and volume to each interpolated sample is determined by calculating a proportion matrix that is in turn used to obtain regularly spaced time series of pollen and macroremain influx. We applied this algorithm to sedimentary data from a subalpine lake situated in the European Alps. The reconstructed total floristic richness at the study site increased gradually when both pollen and macroremain records indicated a decrease in relative abundances of shrubs and an increase in trees from 11,000 to 7,000 cal BP. This points to an ecosystem change that favored trees against shrubs, whereas herb abundance remained stable. Since 6,000 cal BP, local richness decreased based on plant macroremains, while pollen-based richness was stable. The reconstructed richness and evenness are interrelated confirming the difficulty to distinguish these two aspects for the studies in paleo-biodiversity. The present study shows that low-influx bio-proxy records (here macroremains) can be used to reconstruct stand diversity and address ecological issues. These developments on macroremain and pollen records may contribute to bridge the gap between paleoecology and biodiversity studies.     

Human impacts in soft‐sediment assemblages at Casey Station,East Antarctica: Spatial variation,taxonomic resolution and data transformation

Abstract Differences between marine soft‐sediment assemblages at disturbed (two waste dumps, a sewage outfall and a wharf) and control locations were found at Casey Station, Antarctica. These differences were significant against considerable background spatial variability. Core samples were collected by divers using a hierarchical, spatially nested sampling design incorporating four scales: (i) locations (thousands of metres apart); (ii) sites (hundreds of metres apart); (iii) plots (tens of metres apart); and (iv) among replicates within plots (approximately 1 metre apart). Control locations had greater species richness and diversity than disturbed locations and there were many taxa found at control locations that were not recorded at disturbed locations. Assemblages at disturbed locations were less variable than those at control locations. In contrast, populations of some dominant species were more variable at disturbed locations than at control locations. Significant variation in populations of individual taxa was also found at all scales and although greatest at the level of location, variation was also large at the smallest scale, between replicate cores, indicating significant small‐scale patchiness in populations of taxa. Patterns of assemblage structure were similar at fine (77 taxa) and medium (aggregated to 33 taxa) levels of taxonomic resolution, but changed at coarse levels of resolution (nine phyla). Soft‐sediment assemblages at Casey Station are markedly different from those reported from other areas of Antarctica. Assemblages at Casey are almost completely dominated by crustaceans (up to 99% total abundance) and polychaetes are rare or in very low abundances in most areas investigated in the region. This is also the first demonstration that small Antarctic research stations (population 20–50 people) may cause impacts that are detectable in the adjacent marine environment.