Early Holocene greening of the Sahara requires Mediterranean winter rainfall.

The greening of the Sahara, associated with the African Humid Period (AHP) between ca. 14,500 and 5,000 y ago, is arguably the largest climate-induced environmental change in the Holocene; it is usually explained by the strengthening and northward expansion of the African monsoon in response to orbital forcing. However, the strengthened monsoon in Early to Middle Holocene climate model simulations cannot sustain vegetation in the Sahara or account for the increased humidity in the Mediterranean region. Here, we present an 18,500-y pollen and leaf-wax δD record from Lake Tislit (32° N) in Morocco, which provides quantitative reconstruction of winter and summer precipitation in northern Africa. The record from Lake Tislit shows that the northern Sahara and the Mediterranean region were wetter in the AHP because of increased winter precipitation and were not influenced by the monsoon. The increased seasonal contrast of insolation led to an intensification and southward shift of the Mediterranean winter precipitation system in addition to the intensified summer monsoon. Therefore, a winter rainfall zone must have met and possibly overlapped the monsoonal zone in the Sahara. Using a mechanistic vegetation model in Early Holocene conditions, we show that this seasonal distribution of rainfall is more efficient than the increased monsoon alone in generating a green Sahara vegetation cover, in agreement with observed vegetation. This conceptual framework should be taken into consideration in Earth system paleoclimate simulations used to explore the mechanisms of African climatic and environmental sensitivity.

Fifty thousand years of Arctic vegetation and megafaunal diet.

Although it is generally agreed that the Arctic flora is among the youngest and least diverse on Earth, the processes that shaped it are poorly understood. Here we present 50 thousand years (kyr) of Arctic vegetation history, derived from the first large-scale ancient DNA metabarcoding study of circumpolar plant diversity. For this interval we also explore nematode diversity as a proxy for modelling vegetation cover and soil quality, and diets of herbivorous megafaunal mammals, many of which became extinct around 10 kyr bp (before present). For much of the period investigated, Arctic vegetation consisted of dry steppe-tundra dominated by forbs (non-graminoid herbaceous vascular plants). During the Last Glacial Maximum (25-15 kyr bp), diversity declined markedly, although forbs remained dominant. Much changed after 10 kyr bp, with the appearance of moist tundra dominated by woody plants and graminoids. Our analyses indicate that both graminoids and forbs would have featured in megafaunal diets. As such, our findings question the predominance of a Late Quaternary graminoid-dominated Arctic mammoth steppe.

Quantification of Asian monsoon variability from 68 ka BP through pollen-based climate reconstruction.

The glacial-interglacial variability of precipitation and its driving mechanism in monsoonal regions has long been a subject of debate. However, there are few records of quantitative climate reconstruction dating to the last glacial cycle in areas dominated by the Asian summer monsoon. Here, using a pollen-based quantitative climate reconstruction based on three sites in areas exposed to the Asian summer monsoon, we demonstrate that climate has undergone great variability over the past 68 ka. The differences between the last glacial and the Holocene optimum could have been as much as 35%-51% for precipitation, and 5-7 °C for mean annual temperature. Our findings also reveal regional heterogeneity during the abrupt climate events of Heinrich Event 1 and Younger Dryas, that drove drier conditions in southwestern China dominated by the Indian summer monsoon, and a wetter climate in central eastern China. The pattern of variation in reconstructed precipitation, exhibiting strong glacial-interglacial variability, is broadly consistent with the stalagmite δ18O records from Southwest China and South Asia. Our results of reconstruction quantify the sensitivity of the MIS3 precipitation to orbital insolation changes, and highlight the prominent influence of interhemispheric temperature gradients on Asian monsoon variability. Comparison with transient simulations and major climate forcings has shown that the mode of precipitation variability during the transition from the last glacial maximum to the Holocene has been significantly modulated by weak or collapsed Atlantic meridional overturning circulation events in addition to insolation forcing.

Climate change threatens olive oil production in the Levant.

The olive tree (Olea europaea L.) is one of the species best adapted to a Mediterranean-type climate1-8. Nonetheless, the Mediterranean Basin is deemed to be a climate change 'hotspot' by the Intergovernmental Panel on Climate Change9,10 because future model projections suggest considerable warming and drying11,12. Within this context, new environmental challenges will arise in the coming decades, which will both weaken and threaten olive-growing areas, leading to a loss of productivity and changes in fruit and oil quality13-15. Olive growing, a core of the Mediterranean economy, might soon be under stress. To probe the link between climate and olive trees, we here report 5,400 years of olive tree dynamics from the ancient city of Tyre, Lebanon. We show that optimal fruiting scales closely with temperature. Present-day and palaeo data define an optimal annual average temperature of 16.9 ± 0.3 °C for olive flowering that has existed at least since the Neolithic period. According to our projections, during the second half of the twenty-first century, temperature increases in Lebanon will have detrimental consequences on olive tree growth and olive oil production, especially in the country's southern regions, which will become too hot for optimal flowering and fruiting. These data provide a template to understand present and future thresholds of olive production under climate change.

Brazilian montane rainforest expansion induced by Heinrich Stadial 1 event.

The origin of modern disjunct plant distributions in the Brazilian Highlands with strong floristic affinities to distant montane rainforests of isolated mountaintops in the northeast and northern Amazonia and the Guyana Shield remains unknown. We tested the hypothesis that these unexplained biogeographical patterns reflect former ecosystem rearrangements sustained by widespread plant migrations possibly due to climatic patterns that are very dissimilar from present-day conditions. To address this issue, we mapped the presence of the montane arboreal taxa Araucaria, Podocarpus, Drimys, Hedyosmum, Ilex, Myrsine, Symplocos, and Weinmannia, and cool-adapted plants in the families Myrtaceae, Ericaceae, and Arecaceae (palms) in 29 palynological records during Heinrich Stadial 1 Event, encompassing a latitudinal range of 30°S to 0°S. In addition, Principal Component Analysis and Species Distribution Modelling were used to represent past and modern habitat suitability for Podocarpus and Araucaria. The data reveals two long-distance patterns of plant migration connecting south/southeast to northeastern Brazil and Amazonia with a third short route extending from one of them. Their paleofloristic compositions suggest a climatic scenario of abundant rainfall and relative lower continental surface temperatures, possibly intensified by the effects of polar air incursions forming cold fronts into the Brazilian Highlands. Although these taxa are sensitive to changes in temperature, the combined pollen and speleothems proxy data indicate that this montane rainforest expansion during Heinrich Stadial 1 Event was triggered mainly by a less seasonal rainfall regime from the subtropics to the equatorial region.

Past and future global transformation of terrestrial ecosystems under climate change.

Impacts of global climate change on terrestrial ecosystems are imperfectly constrained by ecosystem models and direct observations. Pervasive ecosystem transformations occurred in response to warming and associated climatic changes during the last glacial-to-interglacial transition, which was comparable in magnitude to warming projected for the next century under high-emission scenarios. We reviewed 594 published paleoecological records to examine compositional and structural changes in terrestrial vegetation since the last glacial period and to project the magnitudes of ecosystem transformations under alternative future emission scenarios. Our results indicate that terrestrial ecosystems are highly sensitive to temperature change and suggest that, without major reductions in greenhouse gas emissions to the atmosphere, terrestrial ecosystems worldwide are at risk of major transformation, with accompanying disruption of ecosystem services and impacts on biodiversity.

Temperature Range Shifts for Three European Tree Species over the Last 10,000 Years.

We quantified the degree to which the relationship between the geographic distribution of three major European tree species, Abies alba, Fagus sylvatica and Picea abies and January temperature (Tjan) has remained stable over the past 10,000 years. We used an extended data-set of fossil pollen records over Europe to reconstruct spatial variation in Tjan values for each 1000-year time slice between 10,000 and 3000 years BP (before present). We evaluated the relationships between the occurrences of the three species at each time slice and the spatially interpolated Tjan values, and compared these to their modern temperature ranges. Our results reveal that F. sylvatica and P. abies experienced Tjan ranges during the Holocene that differ from those of the present, while A. alba occurred over a Tjan range that is comparable to its modern one. Our data suggest the need for re-evaluation of the assumption of stable climate tolerances at a scale of several thousand years. The temperature range instability in our observed data independently validates similar results based exclusively on modeled Holocene temperatures. Our study complements previous studies that used modeled data by identifying variation in frequencies of occurrence of populations within the limits of suitable climate. However, substantial changes that were observed in the realized thermal niches over the Holocene tend to suggest that predicting future species distributions should not solely be based on modern realized niches, and needs to account for the past variation in the climate variables that drive species ranges.

Genetic structure and diversity of the endangered fir tree of Lebanon (Abies cilicica Carr.): implications for conservation.

The threatened conifer Abies cilicica currently persists in Lebanon in geographically isolated forest patches. The impact of demographic and evolutionary processes on population genetic diversity and structure were assessed using 10 nuclear microsatellite loci. All remnant 15 local populations revealed a low genetic variation but a high recent effective population size. FST -based measures of population genetic differentiation revealed a low spatial genetic structure, but Bayesian analysis of population structure identified a significant Northeast-Southwest population structure. Populations showed significant but weak isolation-by-distance, indicating non-equilibrium conditions between dispersal and genetic drift. Bayesian assignment tests detected an asymmetric Northeast-Southwest migration involving some long-distance dispersal events. We suggest that the persistence and Northeast-Southwest geographic structure of Abies cilicica in Lebanon is the result of at least two demographic processes during its recent evolutionary history: (1) recent migration to currently marginal populations and (2) local persistence through altitudinal shifts along a mountainous topography. These results might help us better understand the mechanisms involved in the species response to expected climate change.

Cedrus libani (A. Rich) distribution in Lebanon: past, present and future.

Long-term vegetation studies are needed to better predict the impact of future climate change on vegetation structure and distribution. According to the IPCC scenario, the Mediterranean region is expected to undergo significant climatic variability over the course of this century. Cedrus libani (A. Rich), in particular, is currently distributed in limited areas in the Eastern Mediterranean region, which are expected to be affected by such climate change. In order to predict the impact of future global warming, we have used fossil pollen data and model simulations. Palaeobotanical data show that C. libani has been affected by both climate change and human activities. Populations of C. libani survived in refugial zones when climatic conditions were less favourable and its range extended during periods of more suitable climate conditions. Simulations of its future geographical distribution for the year 2100 using a dynamic vegetation model show that only three areas from Mount Lebanon may allow its survival. These results extrapolated for cedar forests for the entire Eastern Mediterranean region show that forests in Syria are also threatened by future global warming. In southern Turkey, cedar forests seem to be less threatened. These results are expected to help in the long-term conservation of cedar forests in the Near East.

Genetic consequences of glacial survival and postglacial colonization in Norway spruce: combined analysis of mitochondrial DNA and fossil pollen.

Norway spruce (Picea abies [L.] Karst.) is a broadly distributed European conifer tree whose history has been intensively studied by means of fossil records to infer the location of full-glacial refugia and the main routes of postglacial colonization. Here we use recently compiled fossil pollen data as a template to examine how past demographic events have influenced the species' modern genetic diversity. Variation was assessed in the mitochondrial nad1 gene containing two minisatellite regions. Among the 369 populations (4876 trees) assayed, 28 mitochondrial variants were identified. The patterns of population subdivision superimposed on interpolated fossil pollen distributions indicate that survival in separate refugia and postglacial colonization has led to significant structuring of genetic variation in the southern range of the species. The populations in the northern range, on the other hand, showed a shallow genetic structure consistent with the fossil pollen data, suggesting that the vast northern range was colonized from a single refugium. Although the genetic diversity decreased away from the putative refugia, there were large differences between different colonization routes. In the Alps, the diversity decreased over short distances, probably as a result of population bottlenecks caused by the presence of competing tree species. In northern Europe, the diversity was maintained across large areas, corroborating fossil pollen data in suggesting that colonization took place at high population densities. The genetic diversity increased north of the Carpathians, probably as a result of admixture of expanding populations from two separate refugia.

Similarity of vegetation dynamics during interglacial periods.

The Velay sequence (France) provides a unique, continuous, palynological record spanning the last four climatic cycles. A pollen-based reconstruction of temperature and precipitation displays marked climatic cycles. An analysis of the climate and vegetation changes during the interglacial periods reveals comparable features and identical major vegetation successions. Although Marine Isotope Stage (MIS) 11.3 and the Holocene had similar earth precessional variations, their correspondence in terms of vegetation dynamics is low. MIS 9.5, 7.5, and especially 5.5 display closer correlation to the Holocene than MIS 11.3. Ecological factors, such as the distribution and composition of glacial refugia or postglacial migration patterns, may explain these discrepancies. Comparison of ecosystem dynamics during the past five interglacials suggests that vegetation development in the current interglacial has no analogue from the past 500,000 years.

Impact of Ice Ages on the genetic structure of trees and shrubs.

Data on the genetic structure of tree and shrub populations on the continental scale have accumulated dramatically over the past decade. However, our ability to make inferences on the impact of the last ice age still depends crucially on the availability of informative palaeoecological data. This is well illustrated by the results from a recent project, during which new pollen fossil maps were established and the variation in chloroplast DNA was studied in 22 European species of trees and shrubs. Species exhibit very different levels of genetic variation between and within populations, and obviously went through very different histories after Ice Ages. However, when palaeoecological data are non-informative, inferences on past history are difficult to draw from entirely genetic data. On the other hand, as illustrated by a study in ponderosa pine, when we can infer the species' history with some certainty, coalescent simulations can be used and new hypotheses can be tested.

Glacial refugia: hotspots but not melting pots of genetic diversity.

Glacial refuge areas are expected to harbor a large fraction of the intraspecific biodiversity of the temperate biota. To test this hypothesis, we studied chloroplast DNA variation in 22 widespread European trees and shrubs sampled in the same forests. Most species had genetically divergent populations in Mediterranean regions, especially those with low seed dispersal abilities. However, the genetically most diverse populations were not located in the south but at intermediate latitudes, a likely consequence of the admixture of divergent lineages colonizing the continent from separate refugia.