Shinya Sugita

Pollen-based quantitative reconstructions of Holocene regional vegetation cover (plant-functional types and land-cover types) in Europe suitable for climate modelling

We present quantitative reconstructions of regional vegetation cover in north-western Europe, western Europe north of the Alps, and eastern Europe for five time windows in the Holocene [around 6k, 3k, 0.5k, 0.2k, and 0.05k calendar years before present (bp)] at a 1° × 1° spatial scale with the objective of producing vegetation descriptions suitable for climate modelling. The REVEALS model was applied on 636 pollen records from lakes and bogs to reconstruct the past cover of 25 plant taxa grouped into 10 plant-functional types and three land-cover types [evergreen trees, summer-green (deciduous) trees, and open land]. The model corrects for some of the biases in pollen percentages by using pollen productivity estimates and fall speeds of pollen, and by applying simple but robust models of pollen dispersal and deposition. The emerging patterns of tree migration and deforestation between 6k bp and modern time in the REVEALS estimates agree with our general understanding of the vegetation history of Europe based on pollen percentages. However, the degree of anthropogenic deforestation (i.e. cover of cultivated and grazing land) at 3k, 0.5k, and 0.2k bp is significantly higher than deduced from pollen percentages. This is also the case at 6k in some parts of Europe, in particular Britain and Ireland. Furthermore, the relationship between summer-green and evergreen trees, and between individual tree taxa, differs significantly when expressed as pollen percentages or as REVEALS estimates of tree cover. For instance, when Pinus is dominant over Picea as pollen percentages, Picea is dominant over Pinus as REVEALS estimates. These differences play a major role in the reconstruction of European landscapes and for the study of land cover-climate interactions, biodiversity and human resources.

Phylogeography of Douglas‐fir based on mitochondrial and chloroplast DNA sequences: testing hypotheses from the fossil record

The integration of fossil and molecular data can provide a synthetic understanding of the ecological and evolutionary history of an organism. We analysed range‐wide maternally inherited mitochondrial DNA and paternally inherited chloroplast DNA sequence data with coalescent simulations and traditional population genetic methods to test hypotheses of population divergence generated from the fossil record of Douglas‐fir (Pseudotsuga menziesii), an ecologically and economically important western North American conifer. Specifically, we tested (i) the hypothesis that the Pliocene orogeny of the Cascades and Sierra Nevada caused the divergence of coastal and Rocky Mountain Douglas‐fir varieties; and (ii) the hypothesis that multiple glacial refugia existed on the coast and in the Rocky Mountains. We found that Douglas‐fir varieties diverged about 2.11u2003Ma (4.37u2003Ma–755u2003ka), which could be consistent with a Pliocene divergence. Rocky Mountain Douglas‐fir probably resided in three or more glacial refugia. More variable molecular markers would be required to detect the two coastal refugia suggested in the fossil record. Comparison of mitochondrial DNA and chloroplast DNA variation revealed that gene flow via pollen linked populations isolated from seed exchange. Postglacial colonization of Canada from coastal and Rocky Mountain refugia near the ice margin at the Last Glacial Maximum produced a wide hybrid zone among varieties that formed almost exclusively by pollen exchange and chloroplast DNA introgression, not seed exchange. Postglacial migration rates were 50–165u2003m/year, insufficient to track projected 21st century warming in some regions. Although fossil and genetic data largely agree, each provides unique insights.

Southward Pleistocene migration of Douglas‐fir into Mexico: phylogeography, ecological niche modeling, and conservation of ‘rear edge’ populations

• Poleward Pleistocene plant migration has been an important process structuring modern temperate and boreal plant communities, but the contribution of equatorward migration remains poorly understood. Paleobotanical evidence suggests Miocene or Pleistocene origin for temperate sky island plant taxa in Mexico. These rear edge populations situated in a biodiversity hotspot may be an important reserve of genetic diversity in changing climates. • We used mtDNA sequences, cpDNA sequences and chloroplast microsatellites to test hypotheses of Miocene vs Pleistocene colonization of temperate Douglas-fir in Mexico, explore geographic patterns of molecular variation in relation to Pleistocene climate history using ecological niche models, and assess the taxonomic and conservation implications. • We found strong evidence for Pleistocene divergence of Douglas-fir in Mexico (958 thousand yr before present (ka) with the 90% highest posterior density interval ranging from 1.6 million yr before present (Ma) to 491 ka), consistent with the southward Pleistocene migration hypothesis. Genetic diversity was high and strongly partitioned among populations. Spatial patterns of molecular variation and ecological niche models suggest a complex late Pleistocene history involving periods of isolation and expansion along mountain corridors. • These results highlight the importance of southward Pleistocene migration in establishing modern high-diversity plant communities and provide critical insights into proposals to conserve the unique biodiversity of Mexican Douglas-fir and associated taxa.

Estimating pollen productivity and relevant source area of pollen using lake sediments in Norway: How does lake size variation affect the estimates?

This study aims to estimate relative pollen productivity (PPEs) of major pollen types and the relevant source area of pollen (RSAP) in a semi-open landscape in western Norway. Extended R-value (ERV) models are used to analyse a data set consisting of surface pollen assemblages from 34 lakes and vegetation survey around each site. Ordination techniques indicate relatively short gradients (<2.3 SD) both in the vegetation and pollen data sets. The lake sizes vary from 0.4 ha to 19.0 ha (mean=5.7 ha, sd=4.3 ha) but follow a normal distribution – a departure from the assumption of the ERV models that the lake size should be constant among sites. Simulations demonstrate that, if the sizes of circular-shaped lakes follow a normal distribution, the ERV model-based methods provide the expected PPE and RSAP values using a standardized lake radius. If the distribution is highly skewed or equally random around the mean, the results are not reliable. We apply the analytical strategy implied from the simulations to obtain relative PPEs and the RSAP in western Norway. PPEs for ten taxa (Alnus, Fagus, Picea, Pinus, Quercus, Juniperus, Salix, Calluna, Cyperaceae, Rumex) relative to Poaceae are comparable with some of those previously obtained in different parts of Europe, indicating that there are general patterns of high and low pollen producers that will be useful for reconstruction purposes. The RSAP estimate is the area within a radius of 900–1100 m. This study demonstrates the importance of careful evaluation of the extent to which the departures from the model assumptions affect the outcomes from the ERV model-based analysis.

The role of tree composition in Holocene fire history of the hemiboreal and southern boreal zones of southern Sweden, as revealed by the application of the Landscape Reconstruction Algorithm: Implications for biodiversity and climate-change issues

We present a quantitative reconstruction of local forest history at two sites, Stavsåkra (hemiboreal zone) and Storasjö (southern boreal zone), in southern Sweden (province of Småland) to evaluate possible causes of contrasting Holocene fire histories in mid- and late Holocene. The Landscape Reconstruction Algorithm (LRA) is applied to evaluate between-site differences in the relative abundance of deciduous trees and Pinus (pine) and landscape/woodland openness during the Holocene. The LRA estimates of local vegetation abundance are compared with other proxies of local vegetation, that is, plant and beetle remains. The LRA results suggest that Pinus was a major tree taxon in the woodlands of Storasjö during mid- and late Holocene, while Tilia (linden) and Betula (birch) were dominant at Stavsåkra. The contrasting fire histories are shown to be strongly related to between-site differences in tree composition during mid-Holocene, 4000–2000 bc in particular. The archaeological/historical and beetle data indicate contrasting land uses from c. 1000 bc (late Bronze Age/early Iron Age), grazing in open Calluna heaths at Stavsåkra and woodland grazing at Storasjö. Between-site differences in fire history during late Holocene were likely due to different land-use practices. Between-site differences in tree composition in mid-Holocene are best explained by local climatic and geological/geomorphological differences between the hemiboreal and southern boreal zones of Småland, which might also be the primary cause of between-site differences in land-use histories during late Holocene. Maintenance of biodiversity at the landscape scale in the study area requires that existing old pine woodlands and Calluna heath are managed with fire and cattle grazing. Further climate warming might lead to higher probabilities of climate-induces fire, in particular in pine-dominated woodlands.

Historical land-use and landscape change in southern Sweden and implications for present and future biodiversity.

The two major aims of this study are (1) To test the performance of the Landscape Reconstruction Algorithm (LRA) to quantify past landscape changes using historical maps and related written sources, and (2) to use the LRA and map reconstructions for a better understanding of the origin of landscape diversity and the recent loss of species diversity. Southern Sweden, hemiboreal vegetation zone. The LRA was applied on pollen records from three small bogs for four time windows between AD 1700 and 2010. The LRA estimates of % cover for woodland/forest, grassland, wetland, and cultivated land were compared with those extracted from historical maps within 3-km radius around each bog. Map-extracted land-use categories and pollen-based LRA estimates (in % cover) of the same land-use categories show a reasonable agreement in several cases; when they do not agree, the assumptions used in the data (maps)-model (LRA) comparison are a better explanation of the discrepancies between the two than possible biases of the LRA modeling approach. Both the LRA reconstructions and the historical maps reveal between-site differences in landscape characteristics through time, but they demonstrate comparable, profound transformations of the regional and local landscapes over time and space due to the agrarian reforms in southern Sweden during the 18th and 19th centuries. The LRA was found to be the most reasonable approach so far to reconstruct quantitatively past landscape changes from fossil pollen data. The existing landscape diversity in the region at the beginning of the 18th century had its origin in the long-term regional and local vegetation and land-use history over millennia. Agrarian reforms since the 18th century resulted in a dramatic loss of landscape diversity and evenness in both time and space over the last two centuries leading to a similarly dramatic loss of species (e.g., beetles).

Two hundred years of land-use change in the South Swedish Uplands: comparison of historical map-based estimates with a pollen-based reconstruction using the landscape reconstruction algorithm

Long-term records of environmental history at decadal to millennial time-scales enable an assessment of ecosystem variability and responses to past anthropogenic disturbances and are fundamental for the development of environmental management strategies. This study examines the local variability of land-use history in the South Swedish Uplands over the last 200xa0years based on pollen records from three lake-sediment successions. Temporal changes in the proportional cover of 14 plant taxa were quantified as percentages using the landscape reconstruction algorithm (LRA). The LRA-based estimates of the extent of four land-use categories (cropland, meadows/grassland, wetland, outland/woodland) were compared to corresponding estimates based on historical maps and aerial photographs from ad 1769–1823, 1837–1895, 1946 and 2005. Although the LRA approach tends to overestimate grassland cover by 10–30xa0% for the two earliest time periods, the reconstructed vegetation composition is generally in good agreement with estimates based on the historical records. Subsequently, the LRA approach was used to reconstruct the 200-year history of local land-use dynamics at 20-year intervals around two small lakes. The qualitative assessment of difference approach, which requires fewer assumptions and parameters than LRA for objective evaluation of between-site differences in plant abundances, provides consistent results in general. Significant differences exist in the land-use history between the sites. Local catchment characteristics, such as soil conditions and wetland cover, appear important for the development of human impact on the landscape. Quantifications of past vegetation dynamics provide information on the amplitude, frequency and duration of the land-use changes and their effects on terrestrial and aquatic ecosystems, and should be taken into account when nature conservation strategies are developed.

Are pollen records from small sites appropriate for REVEALS model-based quantitative reconstructions of past regional vegetation? An empirical test in southern Sweden

In this paper we test the performance of the Regional Estimates of VEgetation Abundance from Large Sites (REVEALS) model using pollen records from multiple small sites. We use Holocene pollen records from large and small sites in southern Sweden to identify what is/are the most significant variable(s) affecting the REVEALS-based reconstructions, i.e. type of site (lakes and/or bogs), number of sites, site size, site location in relation to vegetation zones, and/or distance between small sites and large sites. To achieve this objective we grouped the small sites according to (i) the two major modern vegetation zones of the study region, and (ii) the distance between the small sites and large lakes, i.e. small sites within 50, 100, 150, or 200xa0km of the large lakes. The REVEALS-based reconstructions were performed using 24 pollen taxa. Redundancy analysis was performed on the results from all REVEALS-model runs using the groups within (i) and (ii) separately, and on the results from all runs using the groups within (ii) together. The explanatory power and significance of the variables were identified using forward selection and Monte Carlo permutation tests. The results show that (a) although the REVEALS model was designed for pollen data from large lakes, it also performs well with pollen data from multiple small sites in reconstructing the percentage cover of groups of plant taxa (e.g. open land taxa, summer-green trees, evergreen trees) or individual plant taxa; however, in the case of this study area, the reconstruction of the percentage cover of Calluna vulgaris, Cyperaceae, and Betula may be problematic when using small bogs; (b) standard errors of multiple small-site REVEALS estimates will generally be larger than those obtained using pollen records from large lakes, and they will decrease with increasing size of pollen counts and increasing number of small sites; (c) small lakes are better to use than small bogs if the total number of small sites is low; and (d) the size of small sites and the distance between them do not play a major role, but the distance between the small sites and landscape/vegetation boundaries is a determinant factor for the accuracy of the vegetation reconstructions.

Long-term changes in regional vegetation cover along the west coast of southern Norway: The importance of human impact

Special Feature “Millennial to centennial vegetation change” (Eds. Thomas Giesecke, Petr Kuneš & Triin Reitalu). 1Department of Natural History, University Museum, University of Bergen, Bergen, Norway 2Department of Biological Sciences, University of Bergen, Bergen, Norway 3Museum of Archaeology, University of Stavanger, Stavanger, Norway 4Institute of Ecology, Tallinn University, Tallinn, Estonia 5Museum of Cultural History, University of Oslo, Oslo, Norway

The effects of ecotope, microtopography and environmental variables on diatom assemblages in hemiboreal bogs in Northern Europe

Bogs comprise various niches that are ideal habitats for diatoms. Even though diatoms are widely used bioindicators in many aquatic ecosystems, little is known about their within-site distribution and versatility in bogs. The abundance of bogs in hemiboreal region offers a good opportunity to study the variability of diatom assemblages in different bog habitats and factors influencing it. This study aims to describe the diatom assemblages (1) in the most common bog ecotopes and (2) along the microtopography, and (3) to evaluate the extent to which abiotic environmental parameters (i.e. water table, electric conductivity, pH, moisture and shading) affect the diatom assemblages. According to our results, the most widespread and abundant species are typical bog-inhabiting acidophilic taxa, such as Eunotia paludosa, Kobayasiella parasubtillissima and E. bilunaris. Multivariate analyses showed that some patterns in different microhabitats and ecotopes can be distinguished according to diatom assemblages. In this study, the diatom assemblages were affected by moisture, shading and pH. The study demonstrates the potential usefulness of diatom assemblages as bioindicators to provide additional information about the microsite conditions in bogs.