Range-wide phylogeography of the European temperate-montane herbaceous plant Meum athamanticum Jacq.: evidence for periglacial persistence

Aim  The aim of this study is to analyse the genetic population structure of Meum athamanticum Jacq. in order to explore the alternative hypotheses (1) that the central and northern highland populations are the result of post-glacial recolonization from southern refugia, and the disjunct distribution of M. athamanticum can be explained by modern ecological conditions, or (2) that extant populations north of the Alps and Pyrenees persisted in situ during glacial periods.
Location  Europe.
Methods  Variation of amplified fragment length polymorphisms (AFLPs) was analysed for 23 populations from the entire range of the species. We used band-based approaches and methods based on allele frequencies to measure genetic diversity and to identify population structure.
Results  Our analyses reveal a north–south differentiation within M. athamanticum . High levels of genetic diversity, as well as private fragments, are found in populations both north and south of the Alps. Differentiation among populations is lower in the northern than in the southern population group, and significant isolation-by-distance is found only in the latter group.
Main conclusions  Our results indicate that M. athamanticum survived the last ice age in multiple refugia throughout its contemporary range and did not expand into areas north of the Alps from southern refugia. We found evidence that regional-scale migration in northern, formerly periglacial, parts of the species range has resulted in the intermingling of populations. In contrast, southern populations are characterized by long-term isolation. The south-west Alps represent an area where immigration and mixing of populations from northern and southern refugia appears to have taken place.     

Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica)

Aim  Establishing possible relationships between the magnitudes of the glacial distribution of the European beech, Fagus sylvatica L., and its post-glacial spread.
Location  Europe.
Methods  A database of over 400 pollen records has been used to locate Fagus populations at the end of the last glacial and during the post-glacial in Europe and to assess the areal extent of their past distribution.
Results  The rate of late-glacial and post-glacial increase in the number of pollen sites where Fagus was locally present conforms well to a logistic model of population growth. This suggests that the area occupied by beech populations expanded exponentially from the glacial refugia for a duration of over 10,000 years, until about 3500 yr bp . In the past three millennia beech populations increased at a slower rate, tending towards an equilibrium value.
Main conclusions  The conformity of the increase in beech distribution to the classical logistic model of population growth indicates that: (1) a multiplicative biological process was the main factor shaping the pattern of the post-glacial expansion of F. sylvatica in Europe, (2) climate conditions, human activity and competition may have influenced its rate of spread, and (3) beech populations did not expand with a moving closed front, but with a diffuse spread from scattered nuclei. The distribution of Fagus in Europe at the end of the last glacial appears to have been of two orders of magnitude less extensive than at present. Pleistocene refugia were likely to have been a mosaic of sparse stands of small populations scattered in multiple regions. Fagus populations appear to have increased very slowly and to a moderate extent in southern Europe, where they are now declining slightly. The central European populations increased quickly and extensively, reaching northern Europe, and are now approaching their carrying capacity.     

Long-term forest fire ecology and dynamics in southern Switzerland

1 Pollen and charcoal analysis at two lakes in southern Switzerland revealed that fire has had a prominent role in changing the woodland composition of this area for more than 7000 years.
2 The sediment of Lago di Origlio for the period between 5100 and 3100  bc cal. was sampled continuously with a time interval of about 10 years. Peaks of charcoal particles were significantly correlated with repeated declines in pollen of Abies , Hedera , Tilia , Ulmus , Fraxinus excelsior t., Fagus and Vitis and with increases in Alnus glutinosa t., shrubs (e.g. Corylus , Salix and Sambucus nigra t.) and several herbaceous species. The final disappearance of the lowland Abies alba stands at around 3150  bc cal. may be an example of a fire-caused local extinction of a fire-intolerant species.
3 Forest fires tended to diminish pollen diversity. The charcoal peaks were preceded by pollen types indicating human activity. Charcoal minima occurred during periods of cold humid climate, when fire susceptibility would be reduced.
4 An increase of forest fires at about 2100  bc cal. severely reduced the remaining fire-sensitive plants: the mixed-oak forest was replaced by a fire-tolerant alder–oak forest. The very strong increase of charcoal influx, and the marked presence of anthropogenic indicators, point to principally anthropogenic causes.
5 We suggest that without anthropogenic disturbances Abies alba would still form lowland forests together with various deciduous broadleaved tree taxa.     

Vegetation history of Abies alba Mill. (silver fir) in Switzerland – pollen analytical and genetic surveys related to aspects of vegetation history of Picea abies (L.) H. Karsten (Norway spruce)

This contribution deals with some new aspects of the relationship between the vegetation history of Abies alba Mill. (silver fir) and genetic studies of this tree species in Switzerland. The results of the present study confirm the pollen analytical hypothesis that A. alba re-immigrated into Switzerland mainly from glacial refugia located in northern and central Italy. In particular, some distinct immigration routes of silver fir into the Ticino Alps, Valais, the Bernese Oberland (northwestern Alps), and Graubünden (eastern Alps) could be confirmed by genetic studies. Furthermore, the occurrence of other area-specific alleles indicates an additional influence from eastern European refugia on the Swiss gene pool. Moreover, genetic studies on Picea abies (L.) H. Karsten (Norway spruce) confirmed the general immigration routes from the eastern to the western Alps and from Savoie to the Jura Mountains. The combination of tree pollen and macrofossil analyses of Quaternary sediments with genetic studies of the same tree species represents a considerable research potential and is a new approach of floristic and genetic research. Received November 17, 2000 / Accepted April 27, 2001     

Palaeostructures of vegetation at the upper limit of forests in the inner French Alps

The comparison of six pollen diagrams from French Alps allows us to reconstruct the past changes of vegetation structure at the upper limit of Subalpine range. Dense populations of Pinus cembra developed between 6500 and 2400 cal. BP, both in the southern Alps and the northern ones. Southern Alps seem however to be characterised by higher altitudinal limits, as shown by the past development of fir forests at 2080 m a.s.l. in the Ubaye valley. This study highlights the importance of taking in account local parameters in regional or continental reviews.     

A regional study of Holocene climate change and human occupation in Peruvian Amazonia

Aim  To investigate the influence of Holocene climatic and human-induced changes on a region of high biodiversity in southern Peruvian Amazonia.
Location  Four palaeoecological records from separate lakes within a lake district close to the modern city of Puerto Maldonado, Peru.
Results  The lakes provide a palaeoecological record spanning the last 8200 years. A mid-Holocene dry event is documented in all of the records that extend back > 6000 years. The dry event appears to have lasted from c . 7200 yr bp until c . 3300 yr bp . The onset of wetter conditions coincides with the formation of the youngest of the four lakes. The earliest occupation of these sites is inferred from the presence of charcoal at 7200 yr bp , and the first crop pollen is found at 3630 yr bp . Lakes that were regularly occupied were colonized soon after they formed. A reduction in charcoal concentration and the absence of crop pollen after c . 500  bp in all lakes is consistent with site abandonment following conquest.
Main conclusions  The mid-Holocene dry event is suggested to be part of a time-transgressive drying that tracked from north to south in both the Andes and the Amazon lowlands. The last millennium may represent the period of highest sustained lake levels within the Holocene. The proximity of the four lakes allows a landscape-scale analysis of the spatial extent of human disturbance centred on a known site of human occupation and reveals the highly localized nature of pre-Columbian anthropogenic disturbance in Amazonian landscapes. Inferences regarding widespread pre-Columbian landscape modification by indigenous peoples must take into account key site attributes, such as seasonality and proximity to rivers.     

Tree Migration-Rates: Narrowing the Gap between Inferred Post-Glacial Rates and Projected Rates

Faster-than-expected post-glacial migration rates of trees have puzzled ecologists for a long time. In Europe, post-glacial migration is assumed to have started from the three southern European peninsulas (southern refugia), where large areas remained free of permafrost and ice at the peak of the last glaciation. However, increasing palaeobotanical evidence for the presence of isolated tree populations in more northerly microrefugia has started to change this perception. Here we use the Northern Eurasian Plant Macrofossil Database and palaeoecological literature to show that post-glacial migration rates for trees may have been substantially lower (60–260 m yr^–1) than those estimated by assuming migration from southern refugia only (115–550 m yr^–1), and that early-successional trees migrated faster than mid- and late-successional trees. Post-glacial migration rates are in good agreement with those recently projected for the future with a population dynamical forest succession and dispersal model, mainly for early-successional trees and under optimal conditions. Although migration estimates presented here may be conservative because of our assumption of uniform dispersal, tree migration-rates clearly need reconsideration. We suggest that small outlier populations may be a key factor in understanding past migration rates and in predicting potential future range-shifts. The importance of outlier populations in the past may have an analogy in the future, as many tree species have been planted beyond their natural ranges, with a more beneficial microclimate than their regional surroundings. Therefore, climate-change-induced range-shifts in the future might well be influenced by such microrefugia.     

Buildings and wood trade in Aix-en-Provence (South of France) during the Modern period

Many buildings are being restored in the city of Aix-en-Provence, southeastern France and allowed systematic dendrochronological analyses. 144 fir timbers (Abies alba Mill.) were examined and dated between 1303 and 1933. They highlight three felling phases (Medieval period, 17th-18th century, and 19th century). The best documented one corresponds to the 17th-18th centuries and coincides with significant building activity connected to the status of Aix, which is the parliamentary capital of Provence. Around Aix-en-Provence, it is difficult to find timbers with good dimensions. Our results show that timbers were transported from the Alps to Aix’s buildings. Potential testimonies of this timber trade have been found, such as raft assemblage marks, and carved signs and letters. A dendroprovenancing case study shows that between 1810 and 1890, some of Aix’s timbers could have come from the Northern French Alps or the Verdon-Durance region. Finally, the lack of timbers with good dimension and the difficulties of timber supply contributed to the development of reuse economy and technical innovation. These results are the foundation of a future well-established dendroprovenancing approach in the Southern Alps.     

Influence of rainfall seasonality on African lowland vegetation during the Late Quaternary: pollen evidence from Lake Masoko, Tanzania

Aim  To determine if changes in the key bioclimatic parameters associated with rainfall seasonality can be quantified from fossil sequences and to distinguish them from changes in the total annual rainfall.
Location  Lake Masoko, southern Tanzania (9°20' S, 33°45' E, 840 m a.s.l.).
Methods  Fossil pollen was extracted from a long and well-dated sedimentary sequence spanning the period 45,000–4400 cal. yr bp . The modern habitat, geographical distribution and climatic range (mean annual rainfall and temperature, and length of the dry season) of selected East African plants were used to infer past rainfall attributes for the most characteristic fossil pollen taxa identified from the sediments of Lake Masoko.
Results  In the Masoko pollen sequence it has been possible to identify changes in the length/severity of the dry season during the last 45,000 cal. yr bp , which are interpreted to reflect shifts in the mean position of the Intertropical Convergence Zone (ITCZ) over the southern tropics. We suggest that this bioclimatic parameter has been the main driver of the vegetation dynamics in this area. The major inferred change occurred at the Younger Dryas–Holocene transition c . 11,800 cal. yr bp , when semi-deciduous forest disappeared, being replaced in the lake catchment by woodland, which persists to the present day in spite of locally high rainfall.
Main conclusions  In tropical regions under the influence of the ITCZ, the position and strength of which determine the distribution of rainfall through the year, more attention must be paid to the impact of the length and intensity of the dry season on the modern and past distribution and dynamics of the vegetation. This climatic parameter is as important as the total annual amount of rainfall, and probably one of the most relevant in lowland areas.     

Glacial refugia for summer-green trees in Europe and south-west Asia as proposed by ECHAM3 time-slice atmospheric model simulations

Aim  To generate maps of potential refugia for summer-green trees during the Last Glacial Maximum (LGM).
Locations  Southern Europe and south-western Asia.
Methods  Time-slice simulations of the atmospheric climate with the ECHAM3 model are used for the LGM. Limiting factors beyond which cool and warm groups of deciduous trees cannot grow (such as temperature in growing degree days, minimum monthly temperature and precipitation in summer) are chosen. A limited validation by fossil pollen and charcoal records from LGM sites was done.
Results  Two sets of maps extending from Europe to the Caspian region for cool and warm summer-green trees are presented. Three criteria are combined using contour lines to indicate confidence levels. Small areas within the three southern peninsulas of Europe (Spain, Italy and Greece) are highlighted as possible refugia for summer-green trees. Further, areas that have remained poorly known are now proposed as refugia, including the Sakarya–Kerempe region in northern Turkey, the east coast of the Black Sea and the area south of the Caspian Sea.
Main conclusions  The maps produced in this study could be used to facilitate better long-term management for the protection of European and south-western Asian biodiversity.     

The spread of<Emphasis Type="Italic"> Abies</Emphasis> throughout Europe since the last glacial period: combined macrofossil and pollen data

In this paper we reconstruct the location of the last glacial refugia and postglacial spread of Abies throughout Europe based on combined pollen and macrofossil data. More than 208 pollen sequences available in the European Pollen Database (EPD) and 38 macrofossil sites are used to produce distribution maps encompassing a time span between 38000 and 5500 years B.P. The investigation excludes more recent periods, because these could be strongly influenced by human impact. The pollen data presented here confirm long-lasting refugial areas such as southern Italy and Greece already described in previous studies. The combined pollen/macrofossil dataset identifies the Pyrenees as a further important refugium. In addition the pollen data indicate potential refugia in south-east France and north-west Italy. Possible migration tracks of Abies are discussed by comparing the palaeobotanical evidence with isozyme studies on gene markers of recent fir populations.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00334-004-0049-4.     

Lateglacial and Holocene vegetation history in the Insubrian Southern Alps—New indications from a small-scale site

Fundamental uncertainties exist in the study region about the former lowland vegetation at local scales. All existing palaeoecological results are derived from sediments of medium- to large-sized sites (8–5000 ha), which are thought to record mainly regional vegetation in their pollen content. Therefore the very small mire at Balladrum (0.05 ha) was analysed for pollen, plant-macrofossils, and charcoal and the results compared with those of previous studies in the same region. Common regional signals were detected, but also new insights for the tree species Pinus cembra (L.), Abies alba (Mill.) and Castanea sativa (Mill.). Our palaeobotanical data reveal the local dominance of the timberline species P. cembra during the Lateglacial (16500–14250 cal b.p.) at low-altitudes. For A. alba an early presence in the area is suggested by pollen data, corroborating previous high-altitudinal studies indicating the presence of glacial refugia in the region. Occasional findings of C. sativa pollen throughout the Holocene may indicate the local but very rare presence of this species in the Insubrian Southern Alps, in contrast to the conventional opinion that C. sativa was introduced during the Roman Period. Altogether the results confirm the need of multiproxy palaeobotanical records from basins of variable size to assess the past composition of vegetation more accurately. Communicated by F. Bittmann     

Holocene persistence of wooded steppe in the Great Hungarian Plain

Aim We used a combination of new and previously published palaeoecological data to test three hypotheses: (1) that wooded steppe persisted in the Great Hungarian Plain throughout the Holocene; (2) that wooded steppe and steppe were most extensive between c. 9900 and 8300 cal. yr bp (the ‘Boreal steppe’ period); and (3) that Southern Continental, Pontic and Eastern Sub‐Mediterranean steppe species reached the region during the early Holocene via the ‘Lower Danube Corridor’. Location Sarló‐hát oxbow lake, Hungary and the Eastern European wooded steppe zone. Methods Holocene sediments deposited in the Sarló‐hát oxbow lake were subjected to pollen and microcharcoal analyses. Twelve radiocarbon age estimates were obtained to determine sediment chronology. In addition, previously published palaeoecological data from the Great Hungarian Plain were compiled, analysed and compared with previous studies in other regions of steppe and wooded steppe in eastern Europe. Results Palynological data from two sediment cores extending to c. 11,400 cal. yr bp indicate the persistent dominance of the landscape by temperate deciduous wooded steppe throughout the Holocene, although with varying canopy composition. Warm‐continental steppe grasslands and saline tall‐grass meadows developed on edaphically constrained areas, which remained steppe‐dominated throughout the Holocene. The extent of steppe grasslands did not increase between 9900 and 8300 cal. yr bp . After c. 3100 cal. yr bp , anthropogenic activities led to the development of cultural steppe. Thermophilous steppe species of the Southern Continental, Pontic and Sub‐Mediterranean floristic elements probably reached the Great Hungarian Plain principally via the Lower Danube Corridor during the late glacial interstadial and Holocene. Eurythermic members of these elements, however, probably survived the Last Glacial Maximum in favourable microsites, extending their ranges during the Holocene from these local sources. Main conclusions Our results confirm the Holocene persistence of wooded steppe in the Great Hungarian Plain, disprove the ‘Boreal steppe’ theory, and suggest an Early Holocene period of greater vegetation openness between 11,400 and 9900 cal. yr bp . Evidence for the post‐glacial immigration of south‐eastern steppe elements into the Carpathian Basin is equivocal: the last glacial/interglacial presence of several southern steppe species suggests that the Hungarian Plain hosted suitable habitats for them during warm and cold phases alike.     

Post-glacial changes in spatial patterns of vegetation across southern New England

Aim We analysed lake‐sediment pollen records from eight sites in southern New England to address: (1) regional variation in ecological responses to post‐glacial climatic changes, (2) landscape‐scale vegetational heterogeneity at different times in the past, and (3) environmental and ecological controls on spatial patterns of vegetation. Location The eight study sites are located in southern New England in the states of Massachusetts and Connecticut. The sites span a climatic and vegetational gradient from the lowland areas of eastern Massachusetts and Connecticut to the uplands of north‐central and western Massachusetts. Tsuga canadensis and Fagus grandifolia are abundant in the upland area, while Quercus, Carya and Pinus species have higher abundances in the lowlands. Methods We collected sediment cores from three lakes in eastern and north‐central Massachusetts (Berry East, Blood and Little Royalston Ponds). Pollen records from those sites were compared with previously published pollen data from five other sites. Multivariate data analysis (non‐metric multi‐dimensional scaling) was used to compare the pollen spectra of these sites through time. Results Our analyses revealed a sequence of vegetational responses to climate changes occurring across southern New England during the past 14,000 calibrated radiocarbon years before present (cal yr bp ). Pollen assemblages at all sites were dominated by Picea and Pinus banksiana between 14,000 and 11,500 cal yr bp ; by Pinus strobus from 11,500 to 10,500 cal yr bp ; and by P. strobus and Tsuga between 10,500 and 9500 cal yr bp . At 9500–8000 cal yr bp , however, vegetation composition began to differentiate between lowland and upland sites. Lowland sites had higher percentages of Quercus pollen, whereas Tsuga abundance was higher at the upland sites. This spatial heterogeneity strengthened between 8000 and 5500 cal yr bp , when Fagus became abundant in the uplands and Quercus pollen percentages increased further in the lowland records. The differentiation of upland and lowland vegetation zones remained strong during the mid‐Holocene Tsuga decline (5500–3500 cal yr bp ), but the pattern weakened during the late‐Holocene (3500–300 cal yr bp ) and European‐settlement intervals. Within‐group similarity declined in response to the uneven late‐Holocene expansion of Castanea, while between‐group similarity increased due to homogenization of the regional vegetation by forest clearance and ongoing disturbances. Main conclusions The regional gradient of vegetation composition across southern New England was first established between 9500 and 8000 cal yr bp . The spatial heterogeneity of the vegetation may have arisen at that time in response to the development or strengthening of the regional climatic gradient. Alternatively, the differentiation of upland and lowland vegetation types may have occurred as the climate ameliorated and an increasing number of species arrived in the region, arranging themselves in progressively more complex vegetation patterns across relatively stationary environmental gradients. The emergence of a regional vegetational gradient in southern New England may be a manifestation of the increasing number of species and more finely divided resource gradient.     

Pinus cembra L. (arolla pine), a common tree in the inner French Alps since the early Holocene and above the present tree line: a synthesis based on charcoal data from soils and travertines

Aim In this study, charcoal‐based data for Pinus cembra L. (arolla pine) were gathered from soil and travertine sequences in order to reconstruct its historical biogeography at the landscape level in the inner western Alps during the Holocene. Location The study sites are located between 1700 and 2990 m a.s.l., in the southern (Queyras Massif and Ubaye Valley) and the northern (Maurienne Valley) parts of the inner French Alps. Methods Charcoal fragments were extracted from sediments by water sieving, using meshes of 5, 2, 0.8 and 0.4 mm. The charcoal mass of P. cembra was determined in each charcoal assemblage. Accelerator mass spectrometry and conventional ¹⁴C measurements were used to date the fragments. Results Supported by 40 ¹⁴C datings, the fragments show that, over 2000 m a.s.l., P. cembra accounts for around 40% (mean value) of identified fragments. Data reveal that arolla pine once extended between 260 and 375 m above the present‐day local tree lines. It was established in the southern and the northern French Alps from at least c. 9000 and 6000 cal yr bp , respectively. Main conclusions While present‐day populations of P. cembra are very fragmented in the inner French Alps, charcoal records indicate large past occurrences of this tree since the early Holocene. Human disturbance since the Neolithic seems to be the main reason for the regression of the arolla pine woodlands. On the south‐facing slopes of the study sites, currently deforested, this species extended up to 2800 m a.s.l. In the northern areas, charcoal records of the P. cembra expansion are consistent with the regional pollen archives, but in the southern massifs charcoal records indicate its presence c. 2600 years earlier than other palaeobotanical studies suggest. This discrepancy highlights the necessity to crosscheck data using several different proxies in order to assess the validity of conclusions regarding tree development in space and time.     

Revisiting tree-migration rates: Abies alba (Mill.), a case study

At northern temperate latitudes trees have adjusted their ranges substantially in response to changing climates during the Holocene. Results from dispersal model simulations suggest that postglacial migration rates may have been over-estimated from fossil pollen data. As a contribution to this debate, we infer the migration rates of Abies alba (Mill.), silver fir, as a case-study species, by using a spatially explicit approach based on fossil pollen but taking into account its modern genetic diversity pattern. Maximum estimates of migration rates from fossil pollen data alone are higher than 700 m yr^?1 during the Holocene. Considering the potential refugia as suggested from all the fossil data but restricting the area over which silver fir expanded from each glacial refugium using data on the current haplotype distribution, the estimated maximum migration rates of silver fir are less than 250 m yr^?1. Genetic information may allow for (1) the exclusion of those refugial areas where the species may have survived during the last glacial period but from which it did not spread or spread only very locally and (2) the delineation of the areas over which the species spread from each glacial refugium. The estimated rates in the present study are generally consistent with rates suggested from modelling approaches. This study shows that integrating fossil pollen records can improve simulations of dispersal processes and, thus, allow for better predictions of future changes in tree species’ ranges.     

Changes in environmental controls on the growth of Abies alba Mill. in the Vosges Mountains, north-eastern France, during the 20th century

Aim  To analyse the radial growth of silver fir ( Abies alba Mill.) in north-eastern France in relation to soil, climate and interspecific competition factors and to check the temporal stability of their effects during the 20th century.
Location  The Vosges Mountains, located in north-eastern France.
Methods  Data were collected from 143 plots regularly distributed over three altitude ranges (from 330 to 1100 m), three humus forms (moder to mull) and seven types of stands, including a pure stand of silver fir and six mixed stands containing this species. For each plot, an index of mean radial growth of silver fir was calculated, independent of tree age and calendar year of ring formation. Relationships between radial growth and ecological factors were analysed using linear mixed-effects models.
Results  Radial growth of silver fir is low in the presence of Picea abies or of a high density of other dominant trees . Low aluminium concentration and good nitrogen supply, measured by the soil C:N ratio, are correlated with high radial growth. Water reserves, related to available water storage capacity and to lateral run-off, are also correlated with high radial growth. Analysis of environmental effects during the 20th century showed that growth was strongly correlated with nitrogen supply at the start of the century, and with aluminium toxicity and climate at the end of the century. The radial growth of trees located on sites with low nitrogen supply was lower before 1970 than that of trees located on nitrogen-rich soils. At the end of the century, radial growth of silver fir was the same for all levels of nitrogen nutrition.
Main conclusions  Our results indicate that nitrogen supply was a limiting factor for the mean radial growth of silver fir before the 1970s. Eutrophication during the 20th century may then have eliminated limitation by nitrogen.     

The mid-Holocene extinction of silver fir (Abies alba) in the Southern Alps: a consequence of forest fires? Palaeobotanical records and forest simulations

Pollen records suggest that Abies alba played a dominating role in both the montane and lowland forests at the border of the Southern Alps between ca. 8500 and 5700 years ago. Two major declines in fir, at about 7300–7000 cal b.p. and at ca. 6000 cal b.p., followed by the local extinction of the species are characteristic of the area below ca. 1000 m a.s.l. In order to test the impact of fire on the population dynamics of silver fir, a dynamic model (DisCForm) with a fire module was applied to simulate the early- and mid-Holocene forest development. Simulation outputs based on different fire scenarios were compared with the pollen record from Lago di Annone (226 m a.s.l.). The marked Abies decreases shown in the pollen record can be simulated with very intensive fire scenarios, but they do not result in an extinction of silver fir in the model. Low charcoal influx values related to the Abies declines in the palaeobotanical record suggest that fire was not the only reason for the extinction of silver fir. Human impact, as well as Holocene climatic changes leading to temporary moisture deficits and reduced adaptability due to low genetic variation may have had a significant impact on the Abies forests.     

From ice age to modern: a record of landscape change in an Andean cloud forest

Aim To investigate the palaeoecological changes associated with the last ice age, subsequent deglaciation and human occupation of the central Andes. Location Lake Pacucha, Peruvian Andes (13°36′26″ S, 73°19′42″ W; 3095 m elevation). Methods Vegetation assemblages were reconstructed for the last 24 cal. kyr bp (thousand calibrated ¹⁴C years before present), based on pollen analysis of sediments from Lake Pacucha. An age model was established using ¹⁴C accelerator mass spectrometry dates on bulk sediment. Fossil pollen and sedimentological analyses followed standard methodologies. Results Puna brava replaced the Andean forest at the elevation of Lake Pacucha at the Last Glacial Maximum (LGM). Deglaciation proceeded rapidly after 16 cal. kyr bp , and near‐modern vegetation was established by c. 14 cal. kyr bp . The deglacial was marked by the range expansion of forest taxa as grassland taxa receded in importance. The mid‐Holocene was marked by a lowered lake level but relatively unchanged vegetation. Quinoa and maize pollen were found in the latter half of the Holocene. Main conclusions Temperatures were about 7–8 °C colder than present at this site during the LGM. The pattern of vegetation change was suggestive of microrefugial expansion rather than simple upslope migration. The mid‐Holocene droughts were interrupted by rainfall events sufficiently frequent to allow vegetation to survive largely unchanged, despite lowering of the lake level. Human activity at the lake included a 5500‐year history of quinoa cultivation and 3000 years of maize cultivation.