Palaeoecological Data Research Articles

Necrophagy by insects in Oculudentavis and other lizard body fossils preserved in Cretaceous amber

When a vertebrate carcass begins its decay in terrestrial environments, a succession of different necrophagous arthropod species, mainly insects, are attracted. Trophic aspects of the Mesozoic environments are of great comparative interest, to understand similarities and differences with extant counterparts. Here, we comprehensively study several exceptional Cretaceous amber pieces, in order to determine the early necrophagy by insects (flies in our case) on lizard specimens, ca. 99 Ma old. To obtain well-supported palaeoecological data from our amber assemblages, special attention has been paid in the analysis of the taphonomy, succession (stratigraphy), and content of the different amber layers, originally resin flows. In this respect, we revisited the concept of syninclusion, establishing two categories to make the palaeoecological inferences more accurate: eusyninclusions and parasyninclusions. We observe that resin acted as a “necrophagous trap”. The lack of dipteran larvae and the presence of phorid flies indicates decay was in an early stage when the process was recorded. Similar patterns to those in our Cretaceous cases have been observed in Miocene ambers and actualistic experiments using sticky traps, which also act as “necrophagous traps”; for example, we observed that flies were indicative of the early necrophagous stage, but also ants. In contrast, the absence of ants in our Late Cretaceous cases confirms the rareness of ants during the Cretaceous and suggests that early ants lacked this trophic strategy, possibly related to their sociability and recruitment foraging strategies, which developed later in the dimensions we know them today. This situation potentially made necrophagy by insects less efficient in the Mesozoic.

A Historical Ecology of the Limpopo and Kruger National Parks and Lower Limpopo Valley

The paper uses new palaeo-ecological data and a selective review of archaeological and written sources to show how social and natural history over the last 1200 years have interacted to form the present day landscape of Limpopo National Park and Northern Kruger National Park. The long-term mosaic of different communities in this landscape, hunter and gatherers, pastoralists, farmers and traders has, over time, contributed to shape and reshape a heterogeneous landscape. While some features in this landscape, such as water scarcity, have remained stable over time, there have also been major transformations in both the physical landscape and social life. The natural mosaics have been utilised and enhanced over time and the combination of natural and cultural mosaics are reflected in the landscape through archaeological sites, the pollen record and in the present day landscape.

Modelling the location of interglacial microrefugia for cold-adapted species

Rhododendron tomentosum is a vascular plant common in northern regions of Europe, Asia and North America. In central Europe, this species is considered a glacial relict. In recent years, systematic floristic mapping of the distribution of this species in the Bohemian Switzerland National Park has revealed that this rare and highly sensitive species is strongly associated with a specific type of habitat. Rhododendron tomentosum adapted to environmental changes throughout the Holocene: soil acidification, repeated fires and negative effect of forestry on species composition. To identify the survival mechanisms of R. tomentosum, suitable sites in well-preserved parts of a sandstone landscape with assumed minimal human influence were selected. Based on data from intensive field sampling and a set of environmental variables for the whole area of the National Park, habitat suitability maps for R. tomentosum were developed. These maps were then combined with the palaeoecological data and used to identify sources of pollen with greater precision and extend the knowledge of the potential distribution of R. tomentosum outside the intensively sampled areas. Palaeoecological data was particularly useful for reconstructing the past population dynamics of R. tomentosum. The results indicate the long-term stability of sites suitable for this species, supported by locally diversified vegetation development in sandstone areas. This research could therefore provide the first direct evidence of the persistence of R. tomentosum micropopulations throughout a large part of the Holocene.

The natural environment in the vicinity of Lake Sporovskoye in the Late Glacial and Holocene

Due to the relatively small number of lakes in the southern part of Belarus, in the Polesie region, each lake and its bottom sediments are of great scientific interest for palaeoecological reconstructions. Lake Sporovskoye is one of the largest lakes in Belarusian Polesie, previously studied in the field of palaeoecology by a number of researchers. The discovery in 2018 of the Kakoryca-4 archaeological site near Lake Sporovskoye inspired the beginning of this study, during which new palaeoecological data were obtained. The purpose of the article was to build palaeoecological reconstructions based on spore-pollen, macroremains, sedimentological, and radiocarbon analyses of the Sporovo II core. As a result, it was proven that Lake Sporovskoye was formed in the Late Glacial. Water level changes in the lake, as well as a period of increased activity of the Yaselda River, were detected in the Holocene. The author proposed the reconstructions of regional and local vegetation and corrected some previous views on the development of the study area in the Late Glacial and Holocene.

Understanding Spatiotemporal Variation in Richness and Rate of Within-Site Turnover for Vegetation Communities in Western Eurasia over the Last 4000 Years

Vegetation communities are intricate networks of co-occurring species. Logistical challenges in collecting primary data means research often utilises short-term data from restricted geographical areas. In this study, we examine spatiotemporal change in richness and turnover of vascular plants and bryophytes over the last 4000 years at 23 sites in western Eurasia using high-resolution palaeoecological data. We find support for the Latitudinal Diversity Gradient and Altitudinal Diversity Gradient in both the overall vegetation community (arboreal and non-arboreal species) and the shrub and herb sub-community (non-arboreal species only), as well as a significant temporal increase in the gradient of both relationships. There was a temporal increase in (alpha) richness; the rate of turnover was high but temporally consistent for the overall vegetation community and high but decreasing over time for the shrub and herb sub-community. The rate of change in turnover was affected by latitude (steeper negative relationship at higher latitudes) and altitude (steeper negative relationship at lower altitudes). The Diversity-Stability Hypothesis was supported: vegetation communities changed from “lower richness, higher turnover” historically to “higher richness, lower turnover” more recently. Causal mechanisms for these complex interlinked biogeographical patterns remain ambiguous, but likely include climate change, non-native introductions, increasing homogenisation of generalist taxa, landscape simplification, and anthropogenic disturbance. Further research into drivers of the spatiotemporal patterns revealed here is a research priority, which is especially important in the context of biodiversity decline and climate change.

The avian remains from El Juyo, Lower Magdalenian Cantabrian Spain

El Juyo is one of the Cantabrian sites of Iberia known from long ago for its important Lower Magdalenian sequence. The present study reports the results of the zooarchaeological and taphonomic analysis of the avian remains recovered at the site, which complements the archaeological and palaeoecological data already available. The remains recovered are limited, but they seem to indicate that humans were the main accumulating agent in the site, with sporadic presence of owls’ activity. Additionally, humans could have used birds for other purposes than as a dietary resource, as suggested by cut marks observed on two wing bones.

Pollen, macrofossils and sedaDNA reveal climate and land use impacts on Holocene mountain vegetation of the Lepontine Alps, Italy

Both climate change and anthropogenic disturbance affect vegetation composition, but it is difficult to separate these drivers of vegetation change from one another. A better understanding of past vegetation dynamics is necessary to disentangle the influence of different forcing factors and assess future vegetation change. Here we present the first multi-proxy palaeoecological study combining sedimentary ancient DNA (sedaDNA), pollen, spores, stomata, charcoal and plant macrofossils from the Alps. We reconstructed the Holocene vegetation dynamics and fire history at Lago Inferiore del Sangiatto (1980 m asl), a small lake in the subalpine belt of the Ossola region, Italian Lepontine Alps. Afforestation in response to climate warming started at 10,700 cal yr BP with Larix decidua and tree Betula, which formed open forests together with Pinus cembra from 10,500 cal yr BP onwards. Human impact on the regional vegetation started at 5100 cal yr BP, resulting in expansions of Picea abies and Alnus viridis and the collapse of Abies alba. Species response models and ordination analysis show that livestock grazing and fire were major drivers of vegetation change at Lago Inferiore del Sangiatto during the late Holocene. Finally, increasing human impact during the Bronze (ca. 4200–2900 cal yr BP) and Iron Age (ca. 2900–2000 cal yr BP) led to the formation of species-rich larch meadows and alpine pastures that are still dominant today. The palaeoecological data suggest that under projected climate change and land abandonment, the treeline ecotone will likely shift to higher altitudes, leading to important changes in species composition and increasing the risk of biodiversity loss.

A condensed chalk–marl succession on an Early Cretaceous intrabasinal structural high, Danish Central Graben: Implications for the sequence stratigraphic interpretation of the Munk Marl Bed

The Deep Adda-1 well in the Danish Central Graben, North Sea, provides a record of mid-Cretaceous sedimentation on the eastern flank of the intrabasinal Adda–Tyra inversion high. An upper Hauterivian – lower Barremian core in the Tuxen Formation spans the lower boundary of the laminated organic-rich Munk Marl Bed (MMB), a key marker bed in North Sea Cretaceous stratigraphy. Multidisciplinary sedimentological–biostratigraphic–palaeoecological data document the abrupt environmental shift at this boundary. The upper Hauterivian – lowermost Barremian lower Tuxen Formation (nannozones BC10 – lowermost BC14), beneath the MMB, represents a well-ventilated, current-swept setting supporting a diverse benthic fauna and characterized by a condensed succession with hardgrounds, at one level defining a biostratigraphic hiatus, and stacked, thin shallowing-upward parasequences. The succeeding lower Barremian MMB (nannozone BC14) attests to poorly oxygenated bottom waters and a total lack of epi- and infauna; the calm, inhospitable sea floor was intermittently disturbed by muddy turbidity currents and debris flows. The base-MMB surface is a complex fractured hardground indicative of relative sea-level fall and protracted winnowing of the cemented sea floor. The Deep Adda-1 core thus records a sea-level excursion that accompanied the onset of early Barremian oxygen depletion in concert with additional potential forcing factors such as coeval volcanism and watermass warming.

Radiocarbon dates and late Quaternary vertebrates from cave deposits in the central Canadian Rocky Mountains

New radiocarbon dates on biological remains from cave deposits in the Canadian Rocky Mountains range in age from infinite to modern but are largely restricted to the Holocene (9560±40 years B.P. to modern). The consistency in radiocarbon ages is interpreted as resulting from geologic processes associated with deglaciation and may reflect timeframes when cave conditions in some areas of the Canadian Rocky Mountains became suitable for deposition and preservation of biological remains. When considered at taxonomic levels permitted by morphological identification, recovered vertebrate remains from cave deposits are consistent with the modern biota of the region, suggesting early post-Late Glacial Maximum establishment of the modern biota and relative stability of mammal assemblages through the Holocene. Preliminary evaluation of recovered plant remains highlights a potentially underevaluated source of palaeoecological data for higher elevations.

Scaling Behavior of Peat Properties during the Holocene: A Case Study from Central European Russia

A better understanding of past climate change is vital to our ability to predict possible future environmental dynamics. This study attempts to investigate the dynamic features of the temporal variability of peat humification, water table depth and air temperature by analyzing palaeoecological data from the Valdai Uplands region (Central European Russia). The regression analysis revealed the presence of a periodicity of about 6000 years in the reconstructed peat humification timeseries. Nonlinear analysis showed that humification time variability, water table depth and air temperature exhibit persistent long-range correlations of 1/f type. This indicates that a fluctuation in these variables in the past is very likely to be followed by a similar one in the future, but is magnified by 1/f power-law. In addition, it dictates that humification, water table depth and temperature are key parameters of a system that implies the existence of a special structure, such as self-organized criticality, operating close to a minimum stability configuration, and achieves it without any fine adjustment by external forcing. These conclusions point to new avenues for modeling future ecosystem disturbances and, in particular, for predicting relevant extreme events.

Learning from the past: opportunities for advancing ecological research and practice using palaeoecological data.

Palaeoecology involves analysis of fossil and sub-fossil evidence preserved within sediments to understand past species distributions, habitats and ecosystems. However, while palaeoecological research is sometimes made relevant to contemporary ecology, especially to advance understanding of biogeographical theory or inform habitat-based conservation at specific sites, most ecologists do not routinely incorporate palaeoecological evidence into their work. Thus most cross-discipline links are palaeoecology → ecology rather than ecology → palaeoecology. This is likely due to lack of awareness and/or the misnomer that palaeoecology invariably relates to the "distant past" (thousands of years) rather than being applicable to the "recent past" (last ~ 100-200years). Here, we highlight opportunities for greater integration of palaeoecology within contemporary ecological research, policy, and practice. We identify situations where palaeoecology has been, or could be, used to (1) quantify recent temporal change (e.g. population dynamics; predator-prey cycles); (2) "rewind" to a particular point in ecological time (e.g. setting restoration/rewilding targets; classifying cryptogenic species); (3) understand current ecological processes that are hard to study real-time (e.g. identifying keystone species; detecting ecological tipping points); (4) complement primary data and historical records to bridge knowledge gaps (e.g. informing reintroductions and bioindicator frameworks); (5) disentangle natural and anthropogenic processes (e.g. climate change); and (6) draw palaeoecological analogues (e.g. impacts of pests). We conclude that the possibilities for better uniting ecology and palaeoecology to form an emerging cross-boundary paradigm are as extensive as they are exciting: we urge ecologists to learn from the past and seek opportunities to extend, improve, and strengthen their work using palaeoecological data.

Synthesis of palaeoecological data from the Polish Lowlands suggests heterogeneous patterns of old-growth forest loss after the Migration Period

Human impact on Central European forests dates back thousands of years. In this study we reanalyzed 36 published pollen data sets with robust chronologies from Polish Lowlands to determine the patterns of large-scale forest decline after the Migration Period (fourth to sixth century CE). The study revealed substantial heterogeneity in the old-growth forest decline patterns. Using new high-resolution studies, we could better understand the timing of this transition related to increasing economic development. After the Migration Period, forest expansion continued until the seventh to ninth centuries cal. CE, when the dawn of Slavic culture resulted in large-scale forest decline, especially in north-western and north-central Poland. Later, forest decline was recorded mainly in north-eastern Poland and was related to Prussian settlements, including activities associated with the Teutonic Order, as well as with new settlements from the fourteenth century. The composite picture shows a varied spatio-temporal forest loss and transition towards the present-day, human activity dominated landscapes. However, some sites, such as in north-eastern Poland, are characterized by a less abrupt critical transition. The pristine nature of the oak-hornbeam forest had already been destroyed in Early Medieval times (eighth to ninth centuries cal. CE) and the potential for recovery was largely lost. Our study has confirmed previous assumptions that the decline of hornbeam across the Polish Lowlands may be an early indicator of local settlement processes, preceding severe forest loss, and establishment of permanent agriculture.

Long‐term drivers and timing of accelerated vegetation changes in African biomes and their management implications

AbstractAimOwing to its diverse bioclimatic zones, long human history and intense anthropogenic impacts, Africa provides a model system for studying how global terrestrial ecosystems might respond to accelerated socio‐environmental stress. Africa is particularly vulnerable to climate change and human impact, and insufficient baseline data hamper current environmental management efforts. Using palaeoecological data, we seek to identify the timing, pace and drivers of change in African biomes on a long‐term scale to inform current ecosystem management frameworks on the continent.LocationAfrica.Time period0–12 ka.Major taxa studiedAfrican biomes.MethodsSixty‐four pollen records across Africa and nearby sites were retrieved from multiple databases/sources and grouped into biomes. Turnover (quantified using the squared chord distance dissimilarity metric) and rarefaction analyses were conducted on pollen records in each biome group to reconstruct regional temporal vegetation turnover and richness. Reconstructed vegetation turnover and richness were compared with independent records of climate, fire and human activity to identify possible drivers of change.ResultsWe found that the most stable biomes were those with the greatest floristic richness. Southern Africa's mediterranean‐type (SAM) ecosystems were the most stable and northern Africa's mediterranean‐type (NAM) ecosystems were the most unstable (mainly owing to fire). Tropical savannas (TS) and SAM ecosystems expressed the most sensitivity to climatic shifts from ≥6 ka, whereas tropical forests (TF) were relatively stable before human activities intensified from c. 2 ka. Floristic richness also declined across the tropics from c. 2 ka.Main conclusionsOur analysis pinpoints NAM ecosystems as undergoing the fastest acceleration in turnover on the continent in response to fire, whereas TF and TS have been undergoing unprecedented changes in biodiversity in the last 2,000 years. We expect further changes in biodiversity where climate becomes warmer and drier and where human impacts are novel and rapid in comparison to long‐term baselines.

Using palaeolimnology to guide rehabilitation of a culturally significant lake in New Zealand

Abstract Lakes are becoming degraded at an accelerating rate owing to human activity, and understanding their past ecology is necessary for lake management and rehabilitation. Palaeolimnology provides numerous methods that enable the historical state of lakes to be determined. New Zealand provides an ideal setting in which to do this as human modification of the landscape occurred later here than in most regions of the world (approx. 1300 CE). Lake Oporoa is a shallow lake that is highly significant to the local indigenous Māori community. This study used multiple proxy palaeolimnology to explore how lake ecology shifted following Māori and European settlement in the catchment, and how palaeolimnological data can be used to inform lake rehabilitation and conservation measures, alongside the desires of the indigenous community. Sedimentary pollen, diatoms, bacterial communities, and elemental and hyperspectral imaging scanning were used to infer ecological changes in the lake and catchment from pre‐human times to present. Following Māori settlement (approx. 1620 CE) there was gradual vegetation change and a rapid shift in diatom and bacterial assemblages, but not in phytoplankton pigments or sediment geochemistry. An increasing abundance of diatom taxa Discostella stelligera and Staurosirella cf. ovata indicates early nutrient enrichment. European pastoralism from approximately 1840 CE resulted in further deforestation, and all proxies show evidence for enhanced primary productivity driven by a combination of nutrient enrichment and changing lake levels, particularly since the 1960s. This has caused degradation in water quality and is likely to have contributed to the decline in populations of tuna (eel, Anguilla spp.). Conversations with local Māori, together with the palaeolimnological results, indicate that a culturally acceptable and realistic rehabilitation target for Lake Oporoa aligns with ecological conditions in the 1950s. The palaeoecological data provide information to guide catchment and lake revegetation and other methods of nutrient abatement, with the eventual aim of restoring culturally important tuna and native fish populations.

Newly initiated carbon stock, organic soil accumulation patterns and main driving factors in the High Arctic Svalbard, Norway

High latitude organic soils form a significant carbon storage and deposition of these soils is largely driven by climate. Svalbard, Norway, has experienced millennial-scale climate variations and in general organic soil processes have benefitted from warm and humid climate phases while cool late Holocene has been unfavourable. In addition to direct effect of cool climate, the advancing glaciers have restricted the vegetation growth, thus soil accumulation. Since the early 1900’s climate has been warming at unprecedented rate, assumingly promoting organic soil establishment. Here we present results of multiple organic soil profiles collected from Svalbard. The profiles have robust chronologies accompanied by soil property analyses, carbon stock estimations and testate amoeba data as a proxy for soil moisture. Our results reveal relatively recent initiation of organic soils across the Isfjorden area. The initiation processes could be linked to glacier retreat, and improvement of growing conditions and soil stabilization. Carbon stock analyses suggested that our sites are hot spots for organic matter accumulation. Testate amoebae data suggested drying of soil surfaces, but the reason remained unresolved. If continued, such a process may lead to carbon release. Our data suggest that detailed palaeoecological data from the Arctic is needed to depict the on-going processes and to estimate future trajectories.

The influences of volcanic eruptions on the development of peatland: A case study from the Changbai Mountains, Northeast Asia

Peatlands often develop around active volcanoes. However, little research has been conducted on the impacts of volcanic eruptions on peatland development. Here, we investigated the impacts of Holocene volcanic eruptions of the Changbaishan Volcanic field on the development of Dongfanghong peatland by using basal peat ages, geochemical and palaeoecological data. Analytical results suggest that lateral expansion process of the Dongfanghong peatland was due to paludification. This process was mainly controlled by complex volcanic lava terrain rather than climatic changes. Volcanic rocks provided an impervious horizon for the establishment of peat plants. Thick tephra layers resulted in the retrogressive succession of vegetation community that changed from mesomoor community dominated by the mixture of herbs and Sphagnum into the lowmoor community dominated by herbs. The subsequent high carbon accumulation rates were induced through the phosphorus release from tephra layers and formation of a impervious horizon. These results suggest that volcanic eruptions can significantly influence peatland development.

Legacies of Indigenous land use and cultural burning in the Bolivian Amazon rainforest ecotone.

The southwestern Amazon Rainforest Ecotone (ARE) is the transitional landscape between the tropical forest and seasonally flooded savannahs of the Bolivian Llanos de Moxos. These heterogeneous landscapes harbour high levels of biodiversity and some of the earliest records of human occupation and plant domestication in Amazonia. While persistent Indigenous legacies have been demonstrated elsewhere in the Amazon, it is unclear how past human-environment interactions may have shaped vegetation composition and structure in the ARE. Here, we examine 6000 years of archaeological and palaeoecological data from Laguna Versalles (LV), Bolivia. LV was dominated by stable rainforest vegetation throughout the Holocene. Maize cultivation and cultural burning are present after ca 5700 cal yr BP. Polyculture cultivation of maize, manioc and leren after ca 3400 cal yr BP predates the formation of Amazonian Dark/Brown Earth (ADE/ABE) soils (approx. 2400 cal yr BP). ADE/ABE formation is associated with agroforestry indicated by increased edible palms, including Mauritia flexuosa and Attalea sp., and record levels of burning, suggesting that fire played an important role in agroforestry practices. The frequent use of fire altered ADE/ABD forest composition and structure by controlling ignitions, decreasing fuel loads and increasing the abundance of plants preferred by humans. Cultural burning and polyculture agroforestry provided a stable subsistence strategy that persisted despite pronounced climate change and cultural transformations and has an enduring legacy in ADE/ABE forests in the ARE. This article is part of the theme issue 'Tropical forests in the deep human past'.

Speleological and environmental history of Lida Ajer cave, western Sumatra.

Some of the earliest evidence for the presence of modern humans in rainforests has come from the fossil deposits of Lida Ajer in Sumatra. Two human teeth from this cave were estimated to be 73–63 thousand years old, which is significantly older than some estimates of modern human migration out of Africa based on genetic data. The deposits were interpreted as being associated with a rainforest environment based largely on the presence of abundant orangutan fossils. As well as the main fossil-bearing chamber, fossil-bearing passages are present below a sinkhole, although the relationship between the different fossil deposits has only been tenuously established. Here, we provide significant new sedimentological, geochronological and palaeoecological data aimed at reconstructing the speleological and environmental history of the cave and the clastic and fossil deposits therein. Our data suggest that the Lida Ajer fossils were deposited during Marine Isotope Stage 4, with fossils from the lower passages older than the main fossil chamber. Our use of stable carbon and oxygen isotope analyses of mammalian tooth enamel demonstrates that early humans probably occupied a closed-canopy forest very similar to those present in the region today, although the fossil orangutans may have occupied a slightly different niche.This article is part of the theme issue ‘Tropical forests in the deep human past’.

Palaeoecological data indicates land-use changes across Europe linked to spatial heterogeneity in mortality during the Black Death pandemic

The Black Death (1347–1352 ce) is the most renowned pandemic in human history, believed by many to have killed half of Europe’s population. However, despite advances in ancient DNA research that conclusively identified the pandemic’s causative agent (bacterium Yersinia pestis), our knowledge of the Black Death remains limited, based primarily on qualitative remarks in medieval written sources available for some areas of Western Europe. Here, we remedy this situation by applying a pioneering new approach, ‘big data palaeoecology’, which, starting from palynological data, evaluates the scale of the Black Death’s mortality on a regional scale across Europe. We collected pollen data on landscape change from 261 radiocarbon-dated coring sites (lakes and wetlands) located across 19 modern-day European countries. We used two independent methods of analysis to evaluate whether the changes we see in the landscape at the time of the Black Death agree with the hypothesis that a large portion of the population, upwards of half, died within a few years in the 21 historical regions we studied. While we can confirm that the Black Death had a devastating impact in some regions, we found that it had negligible or no impact in others. These inter-regional differences in the Black Death’s mortality across Europe demonstrate the significance of cultural, ecological, economic, societal and climatic factors that mediated the dissemination and impact of the disease. The complex interplay of these factors, along with the historical ecology of plague, should be a focus of future research on historical pandemics.

Coprolite palynology from Abrigo do Lagar Velho (Portugal) and a revision of Gravettian vegetation in the Iberian Peninsula

Palynological investigations on Gravettian coprolites of Abrigo do Lagar Velho (Vale do Lapedo, Leiria, Portugal) dated between ca. 30 to ca. 27.3 ka cal BP are presented. The palaeoecological data shows a semi-forested landscape with Pinus, Poaceae, Erica, Artemisia, Juniperus and Quercus as the dominant taxa. A diversity of trees, shrubs, and herbs is also noticeable, including broad-leaf trees, Mediterranean woody shrubs and trees, conifers, xerothermophytes, indicators of saline substrates and heliophytes such as Erica, Asteroideae, Cistaceae, Ephedra fragilis and E. distachya. The overall reconstruction is a mosaic of different landscapes or types of vegetation around the rock-shelter. Indeed, we provided approach on the Gravettian vegetation of the Iberian Peninsula, connecting the palaeoecosystem with the biogeography of past human populations during the late Pleistocene.