Validations and descriptions of European syntaxa of vegetation dominated by lichens, bryophytes and algae

Fourty-two high-rank syntaxa and seven associations of the thallophyte system of syntaxa are either described as new or validated in this paper. Among those, there are the following nine classes: Aspicilietea candidae, Caulerpetea racemosae, Desmococcetea olivacei, Entophysalidetea deustae, Gloeocapsetea sanguineae, Mesotaenietea berggrenii, Naviculetea gregariae, Porpidietea zeoroidis, Roccelletea phycopsis. Eleven orders and ten alliances as well as three associations are described or validated: the Aspicilietalia verruculosae (incl. Aspicilion mashiginensis and Teloschistion contortuplicati), the Caulerpetalia racemosae (incl. Caulerpion racemosae), the Desmococcetalia olivacei (incl. Desmococcion olivacei), the Dirinetalia massiliensis, the Fucetalia vesiculosi (incl. Ascophyllion nodosi), the Gloeocapsetalia sanguineae, the Lecideetalia confluescentis (incl. Lecideion confluescentis), the Mesotaenietalia berggrenii (incl. Mesotaenion berggrenii, Mesotaenietum berggrenii and Chloromonadetum nivalis), the Naviculetalia gregariae (incl. Oscillatorion limosae and Oscillatorietum limosae), the Porpidietalia zeoroidis (incl. Porpidion zeoroidis), and the Roccelletalia fuciformis (incl. Paralecanographion grumulosae). Further, five orders, seven alliances and four associations, classified in known classes, were described as well. These include: the Bacidinetalia phacodis, the Agonimion octosporae and the Dendrographetalia decolorantis (all in the Arthonio radiatae-Lecidelletea elaeochromae), the Staurothelion solventis (in the Aspicilietea lacustris), the Pediastro duplicis-Scenedesmion quadricaudae and the Pediastro duplicis-Scenedesmetum quadricaudae (both in the Asterionelletea formosae), the Peccanion coralloidis and the Peltuletalia euplocae (both in the Collematetea cristati), the Laminarion hyperboreae, the Saccorhizo polyschidi-Laminarietum and the Alario esculenti-Himanthalietum elongatae (all in the Cystoseiretea crinitae), the Delesserietalia sanguinei, the Delesserion sanguinei and the Delesserietum sanguineae (all in the Lithophylletea soluti), as well as the the Rinodino confragosae-Rusavskietalia elegantis and the Rhizocarpo geographici-Rusavskion elegantis (both in the Rhizocarpetea geographici).

“Islands” in Holocene forests: implications for forest openness, landscape clearance and “culture-steppe” species

Human activity has undoubtedly had a major impact on Holocene forested ecosystems, with the concurrent expansion of plants and animals associated with cleared landscapes and pasture, also known as 'culture-steppe'. However, this anthropogenic perspective may have underestimated the contribution of autogenic disturbance (e.g. wind-throw, fire), or a mixture of autogenic and anthropogenic processes, within early Holocene forests. Entomologists have long argued that the north European primary forest was probably similar in structure to pasture woodland. This idea has received support from the conservation biologist Frans Vera, who has recently strongly argued that the role of large herbivores in maintaining open forests in the primeval landscapes of Europe has been seriously underestimated. This paper reviews this debate from a fossil invertebrate perspective and looks at several early Holocene insect assemblages. Although wood taxa are indeed important during this period, species typical of open areas and grassland and dung beetles, usually associated with the dung of grazing animals, are persistent presences in many early woodland faunas. We also suggest that fire and other natural disturbance agents appear to have played an important ecological role in some of these forests, maintaining open areas and creating open vegetation islands within these systems. More work, however, is required to ascertain the role of grazing animals, but we conclude that fossil insects have a significant contribution to make to this debate. This evidence has fundamental implications in terms of how the palaeoecological record is interpreted, particularly by environmental archaeologists and palaeoecologists who may be more interested in identifying human-environment interactions rather than the ecological processes which may be preserved within palaeoecological records.

Pinus and Prostomis: a dendrochronological and palaeoentomological study of a study of a mid-Holocene woodland in Eastern England

Tree-ring analysis of sub-fossil Pinus sylvestris L. and Quercus sp. and their associated sub-fossil insect assemblages from tree rot holes have been used to study a prehistoric forest buried in the basal peats at Tyrham Hall Quarry, Hatfield Moors SSSI, in the Humberhead Levels, eastern England. The site provided a rare opportunity to examine the date, composition, age structure and entomological biodiversity of a mid-Holocene Pinus-dominated forest. The combined approaches of dendrochronology and palaeoentomology have enabled a detailed picture of the forest to be reconstructed, within a precise time frame. The Pinus chronology has been precisely dated to 2921- 2445 BC against the English Quercus master curve and represents the first English Pinus chronology to be dendrochronologically dated. A suite of important xylophilous (wood-loving) beetles that are today very rare and four species that no longer live within the British Isles were also recovered, their disappearance associated with the decline in woodland habitats as well as possible climate change. The sub-fossil insects indicate that the characteristic species of the site's modern-day fauna were already in place 4000 years ago. These findings have important implications in terms of maintaining long-term invertebrate biodiversity of mire sites.

Postglacial formation and dynamics of North Patagonian Rainforest in the Chonos Archipelago, Southern Chile

Pollen analysis of continuous sediment cores from two lakes in the northern Chonos Archipelago (44S) in southern Chile shows a complete postglacial record of vegetation change. The fossil records indicate that deglaciation was complete in the northern Chonos by at least 13,600 14Cyr BP. Ericaceous heath and grassland persisted for more than 600 years after deglaciation under the influence of dry/cold climates and frequent burning. Nothofagus-Pilgerodendron-Podocarpus forest, with modern analogues in the southern Chonos Archipelago, was established across the northern islands by 12,400 14Cyr BP under increasingly warm and wet climates. There is no evidence for a return to cooler climates during the Younger Dryas chronozone. The rise of Tepualia stipularis and Weinmannia trichosperma as important forest components between 10,600 and 6000 14Cyr BP may be associated withclimates that were warmer than present. The collapse of Pilgerodendron communities during this time may have been triggered by a combination of factors related to disturbance frequency including tephra deposition events, fire and climate change. After 6000 14Cyr BP Pilgerodendron recovers and Nothofagus-Pilgerodendron-Tepualia forest persists until the present. European logging and burning activity may have increased the susceptibility of North Patagonian Rainforest to invasion by introduced species and to future collapse of the long-lived Pilgerodendron communities.

DNA from pollen: principles and potential

This paper describes our recent extraction of ancient DNA (aDNA) from Holocene pollen and discusses the potential of the technique for elucidating timescales of evolutionary change. We show that plastid DNA is recoverable and usable from pollen grains of Scots pine Pinus sylvestris from 10 ka and 100 years ago. Comparison of the ancient sequences with modern sequences, obtained from an extant population, establish a first genetic link between modern and fossil samples of Scots pine, providing a genetic continuity through time. One common haplotype is present in each of the three periods investigated, suggesting that it persisted near the lake throughout the postglacial. The retrieval of aDNA from pollen has major implications for palaeoecology by allowing (i) investigation of population level dynamics in time and space, and (ii) tracing ancestry of populations and developing phylogenetic trees that include extinct as well as extant taxa. The method should work over the last glacial oscillation, thus giving access to ancestry of populations over a crucial period of time for the understanding of the relationship between speciation and climate change.

Long-term ecology of native pinewood communities in East Glen Affric, Scotland

Fossil pollen, stomata and charcoal were examined from a lake sedimentary sequence in the Glen Affric National Nature Reserve, one of the largest areas of remnant native pine woodland in Scotland, in order to assess ecosystem dynamics over the last 11 000 years. Results reveal that pinewood communities have been continuously present in East Glen Affric for the last 8300 years. Pinus sylvestris fi rst arrived in the area around 9900 cal. BP, but occurred in only low abundance for the subsequent 1600 years. Pine populations expanded around 8300 cal. BP and remained in relatively constant abundance throughout the remainder of the Holocene. There is no evidence of a hypothesized regional mid-Holocene ‘ pine decline ’ at the site. Charcoal results reveal that pinewood communities in East Glen Affric do not appear to have been dependent on fire for either their establishment or their maintenance as has previously been suggested.

How can a knowledge of the past help to conserve the future? Biodiversity conservation and the relevance of long-term ecological studies

This paper evaluates how long-term records could and should be utilized in conservation policy and practice. Traditionally, there has been an extremely limited use of long-term ecological records (greater than 50 years) in biodiversity conservation. There are a number of reasons why such records tend to be discounted, including a perception of poor scale of resolution in both time and space, and the lack of accessibility of long temporal records to non-specialists. Probably more important, however, is the perception that even if suitable temporal records are available, their roles are purely descriptive, simply demonstrating what has occurred before in Earth’s history, and are of little use in the actual practice of conservation. This paper asks why this is the case and whether there is a place for the temporal record in conservation management. Key conservation initiatives related to extinctions, identification of regions of greatest diversity/threat, climate change and biological invasions are addressed. Examples of how a temporal record can add information that is of direct practicable applicability to these issues are highlighted. These include (i) the identification of species at the end of their evolutionary lifespan and therefore most at risk from extinction, (ii) the setting of realistic goals and targets for conservation ‘hotspots’, and (iii) the identification of various management tools for the maintenance/restoration of a desired biological state. For climate change conservation strategies, the use of long-term ecological records in testing the predictive power of species envelope models is highlighted, along with the potential of fossil records to examine the impact of sea-level rise. It is also argued that a long-term perspective is essential for the management of biological invasions, not least in determining when an invasive is not an invasive. The paper concludes that often inclusion of a long-term ecological perspective can provide a more scientifically defensible basis for conservation decisions than the one based only on contemporary records. The pivotal issue of this paper is not whether long-term records are of interest to conservation biologists, but how they can actually be utilized in conservation practice and policy.

The VLT-FLAMES Survey of massive stars: Rotation and nitrogen enrichment as the key to understanding massive star evolution

Rotation has become an important element in evolutionary models of massive stars, specifically via the prediction of rotational mixing. Here we study a sample of stars, including rapid rotators, to constrain such models and use nitrogen enrichments as a probe of the mixing process. Chemical compositions (C, N, O, Mg, and Si) have been estimated for 135 early B-type stars in the Large Magellanic Cloud with projected rotational velocities up to similar to 300 km s(-1) using a non-LTE TLUSTY model atmosphere grid. Evolutionary models, including rotational mixing, have been generated attempting to reproduce these observations by adjusting the overshooting and rotational mixing parameters and produce reasonable agreement with 60% of our core hydrogen burning sample. We find (excluding known binaries) a significant population of highly nitrogen-enriched intrinsic slow rotators (nu sin i less than or similar to 50 km s(-1)) incompatible with our models (similar to 20% of the sample). Furthermore, while we find fast rotators with enrichments in agreement with the models, the observation of evolved (dex) fast rotators (log g < 3.7 dex) that are relatively unenriched (a further similar to 20% of the sample) challenges the concept of rotational mixing. We also find that 70% of our blue supergiant sample cannot have evolved directly from the hydrogen-burning main sequence. We are left with a picture where invoking binarity and perhaps fossil magnetic fields is required to understand the surface properties of a population of massive main- sequence stars.

Impacts of climate change on species, populations and communities: palaeobiogeographical insights and frontiers

Understanding climate change and its potential impact on species, populations and communities is one of the most pressing questions of twenty-fi rst-century conservation planning. Palaeobiogeographers working on Cenozoic fossil records and other lines of evidence are producing important insights into the dynamic nature of climate and the equally dynamic response of species, populations and communities. Climatic variations ranging in length from multimillennia to decades run throughout the palaeo-records of the Quaternary and earlier Cenozoic and have been shown to have had impacts ranging from changes in the genetic structure and morphology of individual species, population sizes and distributions, community composition to large-scale bio-diversity gradients. The biogeographical impacts of climate change may be due directly to the effects of alterations in temperature and moisture on species, or they may arise due to changes in factors such as disturbance regimes. Much of the recent progress in the application of palaeobiogegraphy to issues of climate change and its impacts can be attributed to developments along a number of still advancing methodological frontiers. These include increasingly finely resolved chronological resolution, more refi ned atmosphere-biosphere modelling, new biological and chemical techniques in reconstructing past species distributions and past climates, the development of large and readily accessible geo-referenced databases of biogeographical and climatic information, and new approaches in fossil morphological analysis and new molecular DNA techniques.

Phylogeographic insights into cryptic glacial refugia

The glacial episodes of the Quaternary (2.6 million years ago–present) were a major factor in shaping the present-day distributions of extant flora and fauna, with expansions and contractions of the ice sheets rendering large areas uninhabitable for most species. Fossil records suggest that many species survived glacial maxima by retreating to refugia, usually at lower latitudes. Recently, phylogeographic studies have given support to the existence of previously unknown, or cryptic, refugia. Here we summarise many of these insights into the glacial histories of species in cryptic refugia gained through phylogeographic approaches. Understanding such refugia might be important as the Earth heads into another period of climate change, in terms of predicting the effects on species distribution and survival.

Variability in thermal and UV-B energy fluxes through time and their influence on plant diversity and speciation

Throughout Earth's history there have been temporal and spatial variations in the amount of visible and ultraviolet radiation received by ecosystems. This paper examines if temporal changes in these forms of energy receipt could have influenced the tempo and mode of plant diversity and speciation, focusing in particular upon Cenozoic time-scales. Evidence for changing patterns of plant diversity and speciation apparent in various fossil records and molecular phylogenies are considered alongside calculated changes in thermal and solar ultraviolet energy (specifically UV-B) over the past 50 Myr. We suggest that changes in thermal energy influx (amount and variability) affected the tempo of evolution through its influence upon community dynamics (e.g. population size, diversity, turnover, extinctions). It was not only the amount of thermal energy but also variability in its flux that may have influenced these processes, and ultimately the rate of diversification. We suggest that variations in UV-B would have influenced the mode and tempo of speciation through changes to genome stability during intervals of elevated UV-B. We argue, therefore, that although variability in thermal energy and UV-B fluxes through time may lead to the same end-point (changing the rate of diversification), the processes responsible are very different and both need to be considered when linking evolutionary processes to energy flux.

Can we characterise ‘openness’ in the Holocene palaeoenvironmental record? Modern analogue studies of insect faunas and pollen spectra from Dunham Massey deer park and Epping Forest, England

This paper examines the degree to which tree-associated Coleoptera (beetles) and pollen could be used to predict the degree of ‘openness’ in woodland. The results from two modern insect and pollen analogue studies from ponds at Dunham Massey, Cheshire and Epping Forest, Greater London are presented. We explore the reliability of modern pollen rain and sub-fossil beetle assemblages to represent varying degrees of canopy cover for up to 1000m from a sampling site. Modern woodland canopy structure around the study sites has been assessed using GIS-based mapping at increasing radial distances as an independent check on the modern insect and pollen data sets. These preliminary results suggest that it is possible to use tree-associated Coleoptera to assess the degree of local vegetation openness. Additionally, it appears that insect remains may indicate the relative intensity of land use by grazing animals. Our results also suggest most insects are collected from within a 100m to 200m radius of the sampling site. The pollen results suggest that local vegetation and density of woodland in the immediate area of the sampling site can have a strong role in determining the pollen signal.

Carbon accumulation in peatlands of West Siberia over the last 2000 years

We use a network of cores from 77 peatland sites to determine controls on peat C content and peat C accumulation over the last 2000 years (since 2 ka) across Russia's West Siberian Lowland (WSL), the world's largest wetland region. Our results show a significant influence of fossil plant composition on peat C content, with peats dominated by Sphagnum having a lower C content. Radiocarbon-derived C accumulation since 2 ka at 23 sites is highly variable from site to site, but displays a significant N–S trend of decreasing accumulation at higher latitudes. Northern WSL peatlands show relatively small C accumulation of 7 to 35 kg C m-2 since 2 ka. In contrast, peatlands south of 60°N show larger accumulation of 42 to 88 kg C m-2. Carbon accumulation since 2 ka varies significantly with modern mean annual air temperature, with maximum C accumulation found between -1 and 0°C. Rates of apparent C accumulation since 2 ka show no significant relationship to long-term Holocene averages based on total C accumulation. A GIS-based extrapolation of our site data suggests that a substantial amount (~40%) of total WSL peat C has accumulated since 2 ka, with much of this accumulation south of 60°N. The large peatlands in the southern WSL may be an important component of the Eurasian terrestrial C sink, and future warming could result in a shift northward in long-term WSL C sequestration.

4 ◦C and beyond: what did this mean for biodiversity in the past?

How do the predicted climatic changes (IPCC, 2007) for the next century compare in magnitude and rate to those that Earth has previously encountered? Are there comparable intervals of rapid rates of temperature change, sea-level rise and levels of atmospheric CO2 that can be used as analogues to assess possible biotic responses to future change? Or are we stepping into the great unknown? This perspective article focuses on intervals in time in the fossil record when atmospheric CO2 concentrations increased up to 1200 ppmv, temperatures in mid- to high-latitudes increased by greater than 4 ?C within 60 years, and sea levels rose by up to 3 m higher than present. For these intervals in time, case studies of past biotic responses are presented to demonstrate the scale and impact of the magnitude and rate of such climate changes on biodiversity. We argue that although the underlying mechanisms responsible for these past changes in climate were very different (i.e. natural processes rather than anthropogenic), the rates and magnitude of climate change are similar to those predicted for the future and therefore potentially relevant to understanding future biotic response. What emerges from these past records is evidence for rapid community turnover, migrations, development of novel ecosystems and thresholds from one stable ecosystem state to another, but there is very little evidence for broad-scale extinctions due to a warming world. Based on this evidence from the fossil record, we make four recommendations for future climate-change integrated conservation strategies.

Steady-State and Transient Performance of Biodiesel-Fueled Compression-Ignition-Based Electrical Generation

This paper investigates the performance characteristics of rapeseed methyl ester, EN 14214 biodiesel, when used for electrical generation in compression ignition engines. The work was inspired by the need to replace fossil diesel fuel with a sustainable low carbon alternative while maintaining generator performance, power quality, and compliance with ISO 8528-5. A 50-kVA Perkins diesel engine generator was used to assess the impact of biodiesel with particular regard to gen-set fuel consumption, load acceptance, and associated standards. Tests were performed on the diesel gen-set for islanded and grid-connected modes of operation, hence both steady-state and transient performance were fully explored. Performance comparisons were made with conventional fossil diesel fuel, revealing minimal technical barriers for electrical generation from this sustainable, carbon benign fuel. Recommendations for improved transient performance are proposed and validated through tests.