# Linking environmental changes with human occupations between 900 and 400 ka in Western Europe

Marie-Hélène Moncel, Amaelle Landais, Vincent Lebreton, Nathalie

Combourieu-Nebout, Sébastien L Nomade, Lucie Bazin

► To cite this version:

Marie-Hélène Moncel, Amaelle Landais, Vincent Lebreton, Nathalie Combourieu-Nebout, Sébastien L Nomade, et al.. Linking environmental changes with human occupations between 900 and 400 ka in Western Europe. Quaternary International, 2018, 480, pp.78-94. 10.1016/j.quaint.2016.09.065. hal-02099957

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# Linking environmental changes with human occupations between 900 and 400 ka in Western Europe

Marie-Hélène Moncel (1), Amaëlle Landais (2), Vincent Lebreton (1), Nathalie Combourieu-Nebout (1), Sébastien Nomade (2), Lucie Bazin (2,3)

(1) UMR 7194 CNRS, Department of Prehistory, National Museum of Natural History, Paris, France

(2) Laboratoire des Sciences du Climat et de l'Environnement, LSCE/IPSL, CEA-CNRS-Université Paris Saclay, Université Paris-Saclay, F-91198 Gif-sur-Yvette, France

(3) School of Earth and Ocean Sciences, Cardiff University, Cardiff CF10 3AT, UK

## Abstract

Human occupation in Europe strongly fluctuated over the Quaternary. Archaeological records suggest an intermittent human occupation in Western Europe between 900 and 500 ka, especially in the north of Europe at latitude highern than 45°N. On the opposite, southern Europe, more stable from a palaeoenvironmental point of view, was occupied continuously. This period is followed by a more widespread and dense occupation over the last 450 ka. In parallel, the last 900 ka are characterized by global climatic oscillations and display shifts between glacial/drier and interglacial/wetter periods that modulate the general repartition of fauna and flora. The pacing of these climatic periods is well recorded in numerous palaeoclimatic archives that provide global as well as regional information concerning past climatic and environmental changes. A transition is observed from 1.25 Ma until up to 450 ka (Mid-Pleistocene Revolution) with a change of the dominant periodicity of climate cycles from 41ka to 100 ka in the absence of substantial change in orbital forcing. After 450 ka, the amplitude between glacials and interglacials increases. The change in periodicity since 450 ka corresponds to a change in the density of human occupations as well as the Acheulean technoculture expansion in Europe. There is a general perception that these climatic and environmental oscillations have played a role in human occupation and his morphological evolution. For instance, temperate environments might have favoured permanent occupations or occupations over larger territories with
long periods dominated by dry meadows and steppes and followed by the expansion of broadleaf deciduous and confireous forests. Testing this hypothesis for the period encompassing 900 to 500 ka is a challenge because of the lack of a common chronological framework between climatic/environmental archives and sites of human occupation, but also because archaeological records are only snapshots of cultural and morphological changes of hominins.

**Key-Words** : Middle Pleistocene, Western Europe, hominin occupations, environment, climatic data

**Introduction**

The link between hominin occupations and environmental conditions from the earliest evidence of occupations until 400 ka is still under debate (Carbonell et al., 2010; Martinón-Torres et al., 2011a; Mosquera et al., 2013; Rodríguez et al., 2011, 2015; Voinchet et al., 2015). It has been suggested that climatic and environmental changes over the period encompassing 900 to 400 ka have an impact on the continuity of occupation by small hominin groups. Some regional studies put forward an important role of the climatic changes (Rodríguez et al., 2011; Preece and Parfitt, 2012), while others just observe the lack of hominin occupations in some areas without any climatic-based reasons (Despriée et al., 2011). Considering a geographical delimitation at around 45°N, a widespread hypothesis is that Northern Europe would have been occupied predominantly during favourable climatic periods (Carrión et al., 2011; MacDonald et al., 2012). In contrast, Southern Europe would have been occupied more continuously due to moderate climatic variations over time (Almogi-Labin, 2011; Cuenca-Bescós et al., 2011; Rodriguez et al., 2011; Abbate and Sagri, 2012; Muttoni et al., 2015; Orain et al., 2013b).

Over the last few decades, new data from both Northern and Southern Europe established the earliest appearances of assemblages with bifacial tools in Europe between 900 and 500 ka, providing evidence for the onset of new behaviours, including handaxe-making on a core-and-flake substrate. In
addition, recent discoveries in Spain, France and England enriched our vision of human colonization
in both the southern and the northern parts of the continent. They reveal that the onset of the
bifacial technology happened well before 500 ka, i.e. in parallel to occupations without bifaces and to
new behavioural strategies for raw materials supply and land-use patterns. The bifacial technology
would thus have been sporadic before 500 ka, before becoming widespread after the glacial episode
of Marine Isotope Stage (MIS) 12. While handaxes appeared in Africa as early as 1.8 Ma ago, this
tradition appeared later on the European continent limited to Western and Southern Europe (de la
Torre, 2011; Lepre et al., 2011; Beyene et al., 2013). Bifacial technology is observed in some well
dated open-air and cave sites along a geographical belt from Southern and North-Western Europe to
the South Caucasus. There is no trace of bifacial technology in Central Europe, Russia or Central Asia
until 250 ka.

The 1 Ma-500 ka period of time also raises questions concerning the relationship between various
groups of hominins and the onset of these new behaviours, including bifacial or Acheulean
technology (Moncel et al., 2013; Vallverdú et al., 2014). The few hominin fossils dated from this
period (Gran Dolina TD6, Mauer, Boxgrove, Notarchirico) are attributed to either *Homo antecessor* or
*Homo heidelbergensis*, and the diversity of anatomical features suggests possible hominin intra- or
inter-diversity in Europe due to longitudinal migrations of new hominin groups from Asia and/or
speciation events in Africa or Eurasia prior to 600 ka (Bermúdez de Castro et al., 2011; Dennell et al.,
2011; Martinón-Torres et al., 2011a, b; Stringer, 1996, 2012; Vincenzo et al., 2015).

This paper aims to place the main archaeological sites dated back to 1 Ma (early evidence of bifacial
technology), 700 ka (onset of innovation in core technology for instance) and around 500 ka (before
the glacial period MIS 12 considered to be a break in the hominin settlement) within the global
climatic and local environmental frameworks to present a critical review of the current state of
knowledge on the potential impact of climate on the distribution of earliest European hominin
occupations. In the first section, we concentrate on available archaeological data, dating constraints
and in-situ environmental proxies on several European sites that would help to establish the link
between local environment and hominin occupations. Focusing on Mediterranean and northern
temperate areas, the types of lithic series (with or without bifaces) are thus compared with the
available climatic and environmental data. An overview of the glacial-interglacial cycles over the last
million years is presented in the following two sections to introduce the main global climate changes
over this long period as well as the associated regional vegetation responses. We finally discuss the
limitations of a direct comparison between archeological and climatic records. These limitations
result from inherent biases due to archaeological deposits, dating methods uncertainties,
quantitative interpretation of analysed proxies and/or degree of preservation of records. Keeping
these limitations in mind, we present an attempt to discuss hominin occupations/cultures vs global
climatic and environmental records.

Early Hominin occupations in Europe: available archaeological data.

The archaeological data available are variable in quality and in quantity depending on the area. We
decided to select some sites from particular areas of Europe (Figure 1). These sites provide short or
long sequences with well-dated levels of occupations as well as multiple proxies for
paleoenvironmental reconstructions: plant remains, mammals and micro-mammals, types (i.e.
alluvial, lacustrine, volcanic) and dynamic of deposits. These sites are dated by various methods with
variable margins of error (biostratigraphy, radiometric methods, chronostratigraphy) presented in
table 1. Some of the sites show small series of artefacts and provide limited data for our purpose.
However, we decided to keep them since they have recorded useful paleoenvironmental information.
For the following discussion, we separate our sites within two large geographical areas with
delimitation at 45°N (limit of the current Mediterranean climate): (1) the great European northern plain and small basins and (2) the plains and valleys from the south of Europe.

## South Great-Britain

At Happisburgh Site 3, the human occupation is dated to the end of either MIS 21 (866–814 ka) or MIS 25 (970–936 ka). An assemblage of 78 flint cores, flakes and flake-tools has been excavated from fluvial gravels, laminated estuarine sands and silts of the ancestral River Thames and Bytham River (Cromer Forest-bed Formation CF-bF) (Parfitt et al., 2010).

The environment and climate during the period of human occupation can be reconstructed with combination of plant remains, foraminifera, marine molluscs, barnacles, beetles and vertebrates. Most of the artefacts are associated with *Pinus* and *Picea* pollen, conifer wood and pine-cones, indicating regional conifer-dominated forest. Local grassland is indicated by the range of grazers (*Equus suessenbornensis*, *Bovidae and Microtus* spp.) and indirectly by preserved pollen from hyaena coprolite. Mammals include *Mammuthus* cf. *meridionalis*, *Equus suessenbornensis* and at least two species of *Mimomys*. The mean summer temperatures are estimated to lie between 16°C and 18°C and mean winter temperatures between 0°C and -3°C based on the conifer-dominated woodland similar to what exists in southern Scandinavia today.

At Pakefield, a maximum age is indicated by palaeomagnetic data (normal polarity) as possibly the early part of the Brunhes Chron. The hominin presence can be dated to MIS 17 (about 680 ka) at the very youngest or to the later part of MIS 19 (about 750 ka). 32 flakes in local flint came from four different contexts within an interglacial infill of a channel incised into Early Pleistocene marine sediments (Parfitt et al., 2005).

Macro-remains (*Trapa natans*, *Salvinia natans* and *Corema album*), beetles (*Cybister lateralmarginalis*, *Oxytelus opacus* and *Valgus hemipterus*) and the presence of *Hippopotamus* indicate warmer summers (between 18°C and 23°C) and mild winters. *Mammuthus trogontherii*, *Stephanorhinus hundsheimensis*, *Megaloceros savini*, *M. dawkinsi*, *Bison* cf. *schoetensacki*, and
carnivores (*Homotherium* sp., *Panthera leo*, *Canis lupus* and *Crocuta crocuta*) compose the faunal assemblage. Pollen analysis indicates an interglacial with broadleaf woodland including coexisting *Carpinus*. *Mimomys savini* and *M. aff. pusillus*.

The sites of *Brandon Fields, Maidscross Hill and Warren Hill* yielded evidence of a bifacial technology.

Flint bifaces indicate both tools produced by some deep removals and thinner tools produced by a series of removals and final working of the cutting edges and the tip. Recent geological fieldwork has suggested that the sediments with archaeological findings lie on the 2nd terrace of the Bytham (Bridgland et al., 2006). The age of this terrace possibly dates back to MIS 15 (Ashton et al., 2011; Preece and Parfitt, 2012). New ESR-U series investigations confirm the age of MIS 15-14 for the earliest bifaces in Great-Britain (Voinchet et al., 2015) but the environment of occupations cannot be reconstructed. Microfaunal data place these sites in MIS 13 under a landscape of boreal species and open grassland (Candy et al., 2015).

*High Lodge* is 1 km north of Warren Hill, but in a different geological context (Ashton and Lewis, 2012; Ashton et al., 1992). A thick sequence of glacio-fluvial sands and gravels lies above the sediments with archaeological findings. The site contains a rich scraper flint assemblage, without bifaces, described in the past as Clactonian, associated with *Stephanorhinus hundsheimensis* dated back to MIS 13 or possibly earlier. The floodplain sediments contain pollen and beetle remains that indicate coniferous forest in a cool-temperate climate.

The site of *Boxgrove* is included into sediments of freshwater and lagoonal silts with overlying soil horizons, under periglacial sediments, attributed to a temperate climate at the end of MIS 13 (Stringer, 1996; Roberts and Parfitt, 1999; Smith, 2013). The site yielded a large quantity of similar thin and ovate flint bifaces found on *in situ* knapping floors. There is only occasional production of flake tools (simple scrapers or minimally retouched flakes). The fauna, microfauna (*Arvicola*) and plants suggest a mosaic of grassland and freshwater pools, with developing scrub and woodland in a relatively cool climate (Roberts and Parfitt, 1999).
## North France

Cagny-la-Garenne is located in a fluvial unit of the Middle Terrace of the Somme Valley. La Garenne 1 is composed of fluvial silts interstratified with chalky sediments derived from the nearby Chalk slope. The gravels have been attributed to the end of the MIS 13, beginning of MIS 12 based on the regional geological framework of the Somme (Antoine et al., 2007). ESR/U-series dates place this unit at 400 ± 101 ka. The sequence is composed of six levels of occupations (CXCA, CA, LJ, LG, CXB and CXV) and assemblages (bifaces, core and flake manufacture with notches and denticulates) made from the locally available flint. This site was probably a workshop. Assemblages of levels CXCA and CA are in primary context close to the Chalk slope. Series of levels LJ and LG come from fluvial silts, CXB from limestone gravels and the top of level CXV comes from coarse, periglacial gravels (Lamotte and Tuffreau, 2001a, 2001b, 2015).

At La Garenne 2, 100 m from La Garenne 1, the fluvial sequence is banked up against the Chalk slope. Towards the base, four archaeological levels (R1 to R3 and KR) were recovered, while at the top, five levels (I0 to I4 and J) came from gravels. All stages of core working (unifacial and Levallois cores) and biface manufacture are present, including crude and broken bifaces, tranchet flakes, notches, denticulates, scrapers, and a few choppers and chopping tools (Lamotte and Tuffreau, 2001a, b).

## Central France

The site of la Noira is located along a tributary of the Loire River. The substratum of Tertiary lacustrine limestone (Stratum a), is overlain by five Quaternary strata; a coarse slope deposit (Stratum b) is covered by two sequences of fluvial sediments (Strata c and d). The artefacts are associated with the basal coarse sand of stratum b, the lower part of which consists of an accumulation of local 'millstone', a rock formed as slabs by diagenetic silicification within Oligocene lacustrine limestone. The earliest archaeological level with cores, flakes and bifaces is located at the bottom of the sequence in a quarry. The position of the artefacts between deposition of slope materials on the limestone bedrock and later phases of gelifluction and cryoturbation, suggests that hominins were
present after the period of river incision at the beginning of a cold climate corresponding to the start
of MIS 16 (average age of 655 ± 55 ka by ESR on quartz, Despriée et al., 2011). Hominins left the site
and the area when cold conditions arrived. No faunal remains were preserved. The assemblage
recovered from stratum b consists of around 1000 artefacts, including diverse bifaces (bifaces and
bifacial tools), cleavers on slab or flake, cores (bifacial, unifacial, orthogonal), flakes and retouched
flakes (Moncel et al., 2013, 2015b).

## South France

At *Caune de l'Arago*, levels P and Q are dated back to 532 ± 106 ka (Falguères et al., 2015). These levels belong to the Unit I. Sedimentological analysis shows that the deposits are essentially composed of sands blown into the cave by strong northeast winds during a dry phase of MIS 14. The levels yielded assemblages with bifaces (Barsky and Lumley, 2010; Barsky, 2013; Falguères et al., 2015). Discoidal working was applied to quartz or siliceous stone cobbles from 15-30 km away. Retouched flakes are abundant and mainly pointed. Bifacial tools on a variety of raw materials include well-worked bifaces of various sizes with overall management of the volume by a series of invasive bifacial removals and with final retouching. There are also some cleavers on flakes.

The faunal spectrum includes *Ursus deningeri*, *Cuen priscus*, *Vulpes vulpes*, *Lynx spelaeus*, *Panthera cf. pardus*, *Equus mosbachensis*, *Stephanorhinus hemitoechus*, *Bison sp.*, *Ovis ammon antiqua*, *Hemitragus bonali*, *Rangifer tarandus* and *Cervus elaphus* (Moigne et al., 2006). Horses (28%), reindeers (20%) and bears (28%) are most frequent. The cave was alternately a bear den or a den for other carnivores, and habitats for short-term human occupations.

## Spain

At Atapuerca, the sequences of *Sima del Elefante* and *Grand Dolina* display several archaeological
levels.
Lower units TE7 to TE16 at Sima del Elefante show a reversed polarity magnetization that is attributed to the Matuyama Chron, between 1.3 and 1.0 Ma (Parès et al., 2006; Berger et al., 2008; de Lombera-Hermida et al., 2015; Bennàsar et al., 2016). A few cores and flakes on local raw materials were recovered. The occupations were characterized by a humid and warm climate before the cooling recorded from MIS22 (ca. 900 ka).

Upper units TE17 to TE21 exhibit normal polarity. Luminiscence ages for Units TE16 and TE17 confirm the palaeomagnetic data. Unit TE16 has thus been dated to 804 ± 47 ka and 864 ± 88 ka. The luminiscence ages obtained for TE17 are 724 ± 43 ka and 781 ± 63 ka. Pollen shows the presence of *Pinus sylvestris*. The industry is a core-and-flake type on various raw materials as well.

Layer TD6 at Gran Dolina is composed of 2–2.5 m of blocks and gravels within a poor clayey matrix. This layer contains hominin remains (Carbonell et al., 2005) and has been divided into three lithostratigraphical units. TD6 has a pre-Matuyama negative polarity (> 0.78 Ma) (Blain et al., 2013; Parès et al., 2013) confirmed by biostratigraphy. Radiometric dating by ESR on quartz and U-series methods provided an age between 800 and 880 ka (Moreno et al., 2015) associated with MIS 21 (Rodriguez et al., 2011). TD6 displays a core-and-flake industry made of various local raw materials (Carbonell et al., 1999; Ollé et al., 2013).

Sub-unit TD6-2 is characterized by warm and humid climatic conditions (i.e., a continental Mediterranean climate) (Cuenca-Bescós et al., 2016). The high abundance of *Celtis* seeds at sub-unit TD6-2 evidences Mediterranean conditions (Rodriguez et al., 2011). The landscape indicates open woodland and steppe. This may be correlated with the transitional phase of forest development during a cold to warm climatic transition (possibly correlated with the MIS 22/21 transition). A moderate increase in the presence of open dry taxa occurs in several samples from sub-unit TD6-3 to TD6-1, although this should not be interpreted as indicative of cold steppes but of steppe habitats in a mosaic environment.
The site of La Boella is located in a fluvio-deltaic area. Six lithostratigraphic units (Units I–VI, from bottom to top) are described (Vallverdú et al., 2014; Mosquera et al., 2015a, 2015b), one including two crude bifacial tools. Palaeomagnetic analyses indicate reverse polarity for Units I (archaeologically sterile), II and III, all ascribed to the Matuyama chron. The Brunhes/Matuyama boundary was recorded at the base of Unit IV in Pit 1. A robust geochronological age for the lithic assemblages found within Unit II indicates a position within the late Early Pleistocene or late Matuyama Chron (0.96–0.78Ma). The presence of *Mammuthus meridionalis* and the morphology of *Mimomys savini* molars from the top of Unit II at the three localities support the age of the archaeological deposits. The faunal assemblage of level 2 (Unit II) of the Pit 1 locality is dominated by *Mammuthus meridionalis* associated with a few *Dama* cf. *vallonetensis* and *Equus* sp. indicating a mixture of temperate environments.

## Italy

*Notarchirico* is a seven meters thick fluvial derived sedimentary sequence rich in volcanic materials coming from the Monte Vulture stratovolcano. The particularity of this site is the coexistence in the sediments of quartz and volcanic material (mainly sanidines and clinopyroxenes) allowing the use of several radio-isotopic methods (ESR and $^{40}$Ar/$^{39}$Ar). New studies of $^{40}$Ar/$^{39}$Ar and ESR on quartz ages allowed to constrain the time of occupation of this site between 670 ± 4 ka and 614 ± 4 ka, therefore spanning the entire cold stage MIS 16 (Pereira et al., 2015).

The site documents one of the earliest occurrences (levels B, D, F) of bifacial technology in Southern Europe (Lefèvre et al., 2010; Pereira et al., 2015; Piperno, 1999; Santagata, in press). Origin of raw material and core technology are homogenous along the sequence. Hominins produced small flint en-products.. The oldest handaxes found at Notarchirico (levels F) are now securely dated between 670 ± 4 ka and 661 ± 4 ka and prove that hominin populations lived here as early as the beginning of MIS16 (Pereira et al., 2015).
The faunal remains found in the lower levels (E/E1, F, and G) are *Elephas antiquus*, *Dama clactoniana*, *Bos primigenius* and *Bison shoetensacki*. In the higher levels (α, sub-α, A/A1, B and D) new species such as *Megaceros solilhacus* and *Cervus elaphus* were described. The faunal assemblage of Notarchirico belongs to the "Ponte Galeria" faunal unit. From the microfaunal point of view, several species were identified, principally in horizon E1 such as *Pliomys episcopalis*, *Chionomys nivalis*, *Microtus* sp. and *Arvicola cantianus*. Micro-mammals indicate a dry climate typical of Middle Pleistocene glacial period in the Italian peninsula. The vegetation was characteristic of an open and cold environment made of meadows Poaceae.

The stratigraphy of *Isernia* is composed of 5 units. Unit 3 contains the archaeological deposits and a human deciduous tooth and includes 3 sub-units (Gallotti and Peretto, 2015; Peretto et al. 2015a). The geological age of this site has been evaluated through different dating methods, such as ³⁹Ar/⁴⁰Ar, paleomagnetism and biostratigraphy.

Sanidines dating lead to ages of 610 ± 10 and 606 ± 2 ka for sub-unit 3E, which directly overlies archeological layer t.3a (Coltorti et al., 2005). In contrast, the ESR/U-series dating method on tooth enamel led to underestimated ages with an error weighted mean age of 435 ± 24 ka (Qingfeng, 2011). Recent and more robust ³⁹Ar/⁴⁰Ar dates of 583-561 ka (MIS15/MIS14) were proposed by Peretto et al. (2015a) for this site based on more than 90 single crystal sanidine dates. Unit 3a is the richest level, composed of a high concentration of flint and limestone artefacts (without bifaces). Gallotti and Peretto (2015) demonstrate that the lithic assemblage corresponds to well-established mental pictures despite the lack of bifaces. The knappers used mainly a discoid method, applying this cutting method regardless of the size and shape of the original matrix to maintain a high productivity. This method also permits to produce medium-sized flakes that could be turned into small tools. The large number of herbivores indicates that during the MIS15/MIS14 period the Isernia region was an area of open vegetation of woodland-steppe type, rich in pastures, which was home to bison herds and to numerous pachyderms. This kind of environment is a climate with two seasons, a long and arid
season coupled with a shorter one during which the annual rainfall was concentrated (L Thun Hohenstein et al., 2009; Lebreton, 2004; Manzi et al., 2011; Orain et al., 2013b).

Using the different archaeological sites depicted above, it is possible to make a spatial and temporal synthesis of the technological history of European hominins over the last million years.

Before 1 Ma, Western Europe is occupied by hominins using a technology based on cores and flakes despite the few artefacts found in some sites. Core technologies are simple and share common features between sites. Between 1 Ma and 500 ka, sites exhibit two groups of lithic series: core-and-flake industries and series with evidence of a bifacial technology (« Acheulean »?) with or without new behavioural strategies including a larger perimeter for raw material supply and/or more complex core technologies with less dependence to the raw material constraints (Moncel et al., 2015a). Although the lack of bifacial tools may be due in some cases to small excavations or functional reasons, the existence of bifacial tools (Large Cutting Tools-LCTs or bifaces) on these European sites reduces the chronological gap between Africa and Europe for the emergence of this technology (Scott and Gibert, 2009; Jiménez-Arenas et al., 2011; Antoine et al., 2015; Ashton and Scott, 2015; Moncel et al., 2015a; Voinchet et al., 2015). Then, a closer examination of lithic assemblages with bifaces on sites dated between 1 Ma and 500 ka should enable one to assess whether they result from episodic arrivals of new hominin groups, an influx of new ideas (moving faster than hominins), or from a local innovation (Roberts and Parfitt, 1999; Dennell et al., 2010; Hosfield, 2011; Stringer, 2011, 2012; Ashton and Lewis, 2012; Moncel et al., 2013; Ashton et al., 2014; García-Medrano et al., 2014; Meyer et al., 2014; Voinchet et al., 2015). Some sites are associated with an elaborated bifacial technology, related to more complex core technologies, suggesting external origins (Piperno, 1999; Barsky and de Lumley, 2010; Despriée et al., 2011; Lefèvre et al., 2010; Barsky, 2013; Falguères et al., 2015; Pereira et al., 2015; Moncel et al., 2013, 2015a; Ollé et al., 2016). In contrast, the recent discovery of the site of la Boella with bifacial tools dated back to 900-800 ka (Mosquera et al., 2015b; Vallverdú et al., 2014) has shed new light on the emergence of European bifacial technology which could be of local
origin. Finally, among the series without or with bifaces, dated between 700 and 600 ka, some technological innovations are recorded (for instance, at Isernia and la Noira). These features suggest that different traditions possibly cohabited and contemporaneous core-and-flake industries persisted. That also attests that the onset of the bifacial technology is composed of a mixture of behavioral changes.

The archaeological record for the Northwest part of Europe (e.g. Great Britain) is mainly linked with the changing geographical configuration and its connection with the continent. Indeed in the absence of ubiquitous control of fire, the human occupations in Europe were mainly located in the temperate and Mediterranean biotopes and no evidence of continuous occupation above latitude 53°N is known (Parfitt et al., 2010; Hosfield, 2011; Roebroeks and Villa, 2011; Cohen et al., 2012). Prior to MIS 12, Britain was a peninsula of mainland Europe with continuous land links. The earliest sites of Happisburgh Site 3 (either MIS 25 or 21) and Pakefield (probably MIS 17) both appear to lack handaxes, perhaps due to the small sampling size, in contrast to similar sites on the continent. During the glacial time of MIS 16, there is no evidence of occupation in Northwest and mid-West Europe (Ashton and Lewis, 2012; Ashton et al., 2014; Moncel et al., 2015a, 2015b; Parfitt et al., 2005, 2010; Voinchet et al., 2015). The British records might reflect the incursion of several different populations using the bifacial technology after MIS 15. Unfortunately due to the lack of well-dated sites that can be attributed to MIS 13 or possibly MIS 15 on mainland Europe, the source areas for these colonizing populations is not known. There is no evidence for human occupation of Britain during the extreme cold of MIS 12.

The low number of well-dated sites before 500 ka and the uncertain origin of the associated new bifacial technology may also explain the diversity of strategies and assemblage composition, since each site displays individual variations due to activities, raw materials constraints, specific traditions, and possibly taphnomical disturbances (Moncel et al., 2016a, b).

Current data suggest co-existence of diverse traditions: continuity of core-and-flake industries, core-and-flake industries with innovation, introduction of external traditions bringing the bifacial
technology, sometimes more complex core technologies but also new land-use strategies. The flexibility of these site-dependant behaviours, the raw material quality and the continuity or discontinuity of occupations according to areas (absence of cultural transmission) would explain the extreme variability of the assemblages. Clactonian and other so-called traditions without bifaces recall the long-lasting questions about the meaning of these series, considered sometimes as distinct traditions, due to the bad preservation of sites or to the size of the excavated area (Ashton et al., 1994; McNabb, 1996).

Sites are dispersed across the South and North of Europe. The scarcity of sites over such a long period of time suggests short-lived dispersal events and, possibly, a source-sink dynamic from the South with phases of depopulation and recolonization that could be impacted by the environmental and climatic conditions (Dennell et al., 2011; MacDonald et al., 2012; Cuenca-Bescós et al., 2013; Huguet et al., 2013; Rodríguez-Gómez et al., 2014; Leroy et al., 2011; Palombo, 2016). After MIS 12, the number of sites in Western Europe increases probably because of a demographic expansion and adaptability of populations to contrasted climates despite the lack of evidence for the use of fire (i.e. the case of Schöningen in Germany; Conard et al., 2015; Stahlschmidt et al., 2015).

## Climatic context of the last million years

The glacial – interglacial cycles punctuating the last million years have been recorded in many different archives in the marine and continental realms. These records should enable one to depict the variety of the successive glacial and interglacial periods useful to replace the Earliest hominin occupations both in Northern and Southern Europe (Tzedakis et al., 2009; Sadori et al., 2016) (Figure 2). The majority of records are local, rendering it difficult to draw the global picture necessary to assess the global Earth system response, such as the annual mean temperature, inferred from modelling experiments (e.g. Yin and Berger, 2012).
Probably one of the most global climatic parameters is the global sea level that can be inferred by
coral proxies (e.g. Clark et al., 2009) and marine foraminifera δ¹⁸O signal corrected from deep ocean
temperature. A synthetic effort was done by Lisiecki and Raymo (2005) gathering 57 benthic δ¹⁸O
records from different ocean regions leading to a stack covering the last 5.3 Ma often used as a
reference in a number of studies to discuss the evolution of glacial and interglacial amplitudes (e.g.
Masson-Delmotte et al., 2010; Yin and Berger, 2012) (Figure 2a). Alternative sea level curves have
been proposed in parallel based on planktonic δ¹⁸O records with correction for the hydrological signal
(i.e. sapropel events, Rohling et al., 2014, Elderfield et al., 2012, Shakun et al., 2015) (Figure 2b).
Because of the difficulty of (1) deciphering the sea level and deep ocean temperature signals or (2)
properly correcting for local hydrological effects, these different sea level curves are often associated
with large uncertainties (> ± 20 m, Spratt and Lisiecki, 2015). Still, it is possible to identify some strong
interglacials (MIS 19, MIS 15, MIS 11, MIS 9 and MIS 5) and strong glacial maxima (MIS 16, MIS 12,
LGM) in these records (Figure 2b).

Antarctic polar ice cores also provide some insight for documenting the regional to global climatic
and environmental history. The great advantage of polar ice cores is that they provide high resolution
climatic records (~ 20 years resolution, Pol et al., 2014) with a well constrained chronology based on
absolute, orbital and relative time constraints as well as glaciological models (Parrenin et al., 2007;
Lemieux-Dudon et al., 2010; Bazin et al., 2013; Veres et al., 2013). The longest ice core climatic record
has been retrieved at Dome C in East Antarctica (EPICA community Members, 2004). While the
isotopic composition of ice (δD) is a local temperature signal (Jouzel, 2003), it is striking to observe a
remarkable similitude between the sea-level curves and the Antarctic δD record, thus proving that
the Antarctic climate is not isolated and should be integrated in our understanding of the global Earth
climate system. The puzzling element is the response of Antarctic climate to the precession signal.
When a warming occurs during a maximum in Northern hemisphere summer insolation, the northern
warming, as inferred from Greenland ice cores, is delayed compared to Antarctic warming (Masson-
Delmotte et al., 2010).
Ice cores also record global parameters in the air trapped in ice such as the concentration of greenhouse gases (N₂O, CO₂ and CH₄). While CO₂ seems to be mainly driven by changes in the austral ocean over the past 800 000 years (Lüthi et al., 2008), CH₄ variations are linked to the hydrological cycle of the low latitudes during glacial periods (Chappellaz, 2007; Rhodes et al., 2015). Dust concentration recorded in polar ice cores is also an indicator of the lower latitudes and mainly reflects the aridity of the source regions and transportation. The 800 ka dust record from EDC (Lambert et al., 2012) mainly reflects changes in erosion in South America (Delmonte et al., 2008) that can be directly compared with the timing of temperature changes in high latitudes. In general, there is an excellent inverse correlation between temperature in Antarctica and dust concentration, apart from shifts of several thousands of years observed for some deglaciations (e.g. Termination III, 240 ka before present, Röthlisberger et al., 2008). Finally, one noticeable tool for linking high and low latitudes in polar ice cores is the isotopic composition of atmospheric oxygen, hereafter δ¹⁸O_atm (Bender et al., 1994; Landais et al., 2010). Indeed, despite the numerous processes that can be involved, Reutenauer et al. (2015) used a modelling approach to show that δ¹⁸O_atm is mainly dependent on the low latitude hydrological cycle with strong similarities with δ¹⁸O of calcite of low latitude speleothems over the last climatic cycles (Wang et al., 2008; Severinghaus et al., 2009; Landais et al., 2010).

Figure 3 illustrates the major climatic variations recorded in the EPICA Dome C ice core over the past 800 ka. Even if the different ice core proxies presented here (CH₄, δ¹⁸O_atm, dust flux, δD of water) reflects different influences from different latitudes, we nevertheless observe some general features regarding the length and amplitude of the glacial and interglacial stages. The strongest glacial - interglacial transitions, as identified from ice core records, are thus Termination II (from MIS6 to MIS5), Termination IV (from MIS10 to MIS9) and Termination V (from MIS12 to MIS11). These transitions often correspond to the coldest glacials (MIS 2, 12 and 16) and warmest interglacials (MIS5, 9 and 11) already identified in ice cores and global climatic records (Wolff et al., 2015).
It has often been discussed how the succession of glacial-interglacial periods recorded in the different proxies of figure 2 are influenced by orbital parameters. A tilt increase leads to warmer polar temperatures and a precession minimum leads to an increase of Antarctic temperature and methane concentration as well as a decrease in both δ¹⁸O_atm and benthic δ¹⁸O. Gradient between high and low latitudes may thus be linked to the interactions between obliquity and precession as observed over the last glacial inception in the northern hemisphere (Sánchez-Goñi et al., 2008). Precession is indeed a key driver of the low latitude summer insolation changes, inducing oscillations of the Inter-Tropical Convergence Zone (ITCZ) position and hence variations of the monsoon (or low latitude water cycle) signal.

The changes in the timing and amplitudes of glacial – interglacial periods over the last million years result from changes in orbital parameters and greenhouse gases concentration forcing combined with internal feedbacks (albedo, vegetation repartition, strength and organization of the oceanic and atmospheric circulation). Yin and Berger (2012) have recently performed simulations using an intermediate complexity model to decipher the relative contributions of insolation and CO₂ to the amplitude of interglacial periods over the last 800 ka. Modelling outputs confirm data finding for warm MIS5, 9 and 11 and relatively cold MIS13 and 17. They also show that while orbital and greenhouse forcing act together for the warm MIS5 and 9, the orbital forcing over MIS11 is rather weak so that the high temperature level recorded during this period is mainly due to high greenhouse gases concentration. It follows from this study that the latitudinal temperature gradient and vegetation distribution vary under different greenhouse gases and orbital forcing. It is thus expected that the latitudinal temperature gradient should be less pronounced during MIS11 than during the other warm interglacials (MIS5 and MIS9). As noted in the introduction, the timing of glacial-interglacial cycles has clearly shifted from a ~41 ka to a ~100 ka periodicity depicted as the Mid-Pleistocene transition, with a clear 100 ka periodicity since MIS11. Such transition can be explained by simple conceptual models forced by the orbital parameters even if no evident modification of the orbital forcing is detected in the astronomic curves (e.g. Paillard, 1998). While the exact associated
mechanism is still unknown, the driver for the modification of the glacial-interglacial periodicity from
a dominant influence of obliquity to a dominant influence of precession is assumed to be a
progressive cooling of the Earth (Clark et al., 2006; Raymo et al., 2006), possibly induced by a
decrease in atmospheric CO2.

## Regional glacial-interglacial climatic and vegetation patterns during the last million years

While the aforementioned marine and ice cores records can provide a general climatic context for the
last million years, regional and local climatic and environmental records are further required to
examine whether migration and colonization of hominins in Europe can be related to specific
environmental conditions.

Continuous regional climatic records in the Mediterranean region can be obtained from sea surface
temperature proxies in marine cores. In figure 2, we present two continuous records. The first one is
a Mediterranean record of δ¹⁸O of planktonic foraminifera for the last 1 Ma (36°15.25′N, 17°44.34′E
Ionian sea, Lourens, 2004; Figure 2c) and the second is the sea surface temperature reconstruction
based on alkenone from Central North Atlantic (41°0.07′N; 32°57.44′W; U1313; Naafs et al., 2011; Fig
2d). These two records evidence clear differences between the local climate and the global sea level
curves presented above. Before 400 ka, notable differences are also observed between the
Mediterranean and the Atlantic records. First, while the cold MIS12 is very prominent in the Central-
North Atlantic, as observed in the more global records of previous sections, and along the Atlantic
coast (Sanchez-Goñi et al., 2016), it is not recorded as significantly cold in the Mediterranean record:
the same glacial level is actually reached for all displayed glacial stages older than MIS12 in the
Mediterranean region (Figure 2c). Second, while relatively strong amplitudes of glacial – interglacial
cycles are observed in the global records before 400 ka, the Mediterranean curve suggests
significantly lower amplitude of the sea surface temperature changes before 400 ka in Southern
Europe compared to the situation after MIS11.

Climate from the Mediterranean region can also be depicted from pollen records if the climatic
control on vegetation is well understood. Over the Mediterranean region, precession driven summer
insolation associated with ITCZ displacements and North Atlantic Oscillations (NAO) constrains
summer aridity. A Mediterranean bioclimate with seasonal contrasts gradually took place during the
Neogene (Pons et al., 1995; Sadori et al., 2013). However, summer hydric seasonal stress occurred
about 2.6 Ma ago (Pons et al., 1995; Klotz et al., 2006; Combourieu-Nebout et al., 2015). Rainfall amount and seasonality are the dominant factors explaining the distribution of vegetation cover in the Mediterranean region (e.g. Peinado Lorca and Martínez-Parras, 1987; Yin and Berger, 2012). The main characteristic of the Mediterranean vegetation is the ability to withstand long and drastic summer dryness (Quézel and Médail, 2003). Actually, the Mediterranean bioclimate specificity determines the current vegetation contrast in Europe between the middle and low latitudes respectively with the temperate and Mediterranean biomes (Ozenda, 1994; Quézel and Médail, 2003).

No continuous pollen records exist in Europe for our period of interest. Still, from discontinuous sequences, it is possible to depict the characteristic glacial vs interglacial vegetation dynamic in Southern and Northern Europe. Existing pollen records show that global climate changes have a clear impact on the regional vegetation structure and dynamic as attested by singular differences between North and South Europe (Zagwijn, 1996; Tzedakis, 2007; Bertini, 2010; Fletcher et al., 2010; Magri, 2010; Magri and Palombo, 2013; Combourieu-Nebout et al., 2015). The transition from a dominant periodicity of 41 ka to 100 ka of glacial – interglacial climate cycles results in an extended duration and intensification of glacial dry periods in the European region. However, the dynamic of climate changes remains similar with asynchronous oscillations of temperature and precipitation as reconstructed from the vegetation succession of pioneer forest at the beginning of an interglacial, mixed oak forest for the full interglacial, coniferous forest and finally open vegetation for the full glacial (Figure 4; de Beaulieu and Suc, 1985). However, when looking in more details at the composition and structure of the vegetation over the last million years, clear differences are observed that should be linked to latitudinal and/or altitudinal contrasts (Figure 5). The main changes in the temperate and Mediterranean regions are presented below in order to discuss the latitudinal environmental gradients over Europe.

In the temperate environment, some records from the North of Europe illustrate the vegetation dynamics over the last million years (Woillard, 1978; de Beaulieu and Reille, 1984, 1989, 1992a, 1992b; Zagwijn, 1992, 1996; Drescher-Schneider, 2000; Nitychoruk et al., 2000; Müller et al., 2003; Satkunas et al., 2003; de Beaulieu et al., 2006; Hahne et al., 2008; Knipping, 2008; Parfitt et al., 2010; Lindner and Marks, 2015; Pidek, 2015) (Figure 5). Since MIS25 and at least up to MIS17, warm periods are initiated with the spread of a pioneer forest (Pinus and Betula). A mixed-oak forest then developed during the warm optimum, with Alnus, Corylus, Quercus, Ulmus, Carpinus or Ostrya-type. Within this temperate forest, subtropical taxa still persisted such as Eucommia and Pterocarya. By the end of the warm period in the Netherlands and Poland, a cool humid forest extended, dominated by Tsuga and then Picea, associated with Abies in the Netherlands. However in the United Kingdom, heath (Ericaceae) and coniferous forest (Picea and Pinus) developed at the same time. After each
warm period, dry meadows expanded with the spread of herbs (Poaceae, Asteraceae and Cyperaceae) during the cold phases of the climate cycles. The intense glacial related to MIS16 may have acted as a threshold for the most recent glacials and interglacials. The mixed-oak forest became more and more impoverished in taxa such as *Carya*, *Pterocarya*, *Zelkova*, *Celtis*, *Tsuga* from this period and all of them disappeared after the strong MIS12 glacial at ca. 450 ka. Later, when the 100 ka climate cycles are definitively established, the enhanced duration of the cycles and concomitant associated glacial periods impacted the composition and dynamic of the vegetation. The pioneer and coniferous forests were still recorded respectively at the beginning and termination of the warm optimum characterized by the mixed-oak forest without any subtropical trees. Steppic plants increased among the dry meadow during the glacials.

In the Mediterranean environment, numerous pollen records document the vegetation dynamics over the last million years (Wijmstra, 1969; Allen et al., 1999, 2000; Bertini, 2000; Bruguès and Suc, 1980; rmolli and Cheddadi, 1997; Florschütz et al., 1971; Follieri et al., 1988, 1998; Tzedakis et al., 2002, 2006; Joannin et al., 2007, 2008, 2011; Leroy, 1990; Leroy et al., 1994, 1996; Magri, 1999; Magri and Sadori, 1999; Tzedakis, 1999; Moscariello et al., 2000; Okuda et al., 2001; Lebreton, 2002; BCorrado and Magri, 2011; Rodríguez et al., 2011; Eertini et al., 2015; Orain et al., 2015; Pross et al., 2015; Sánchez-Goñi et al., 2016) (Figure 5). The pioneer forest was mainly composed of *Pinus* and the spread of deciduous *Quercus* is recorded at the very beginning of the glacial – interglacial transitions. Then a thermophilous forest spread during the warm optimum with deciduous and evergreen *Quercus*, *Carpinus*, *Ulmus*, together with *Pinus* and few Mediterranean taxa. Mesothermic taxa requiring moisture such as *Carya*, *Pterocarya*, *Zelkova* and *Eucommia* persisted during the optimum although a decreasing trend is notable. The expansion of conifers (*Pinus*, *Abies*, *Picea*) marked the end of the warm periods. In Northern Italy, the forest was enriched in conifers (*Abies*, *Picea*, *Tsuga*, *Cedrus*). In the northern Mediterranean, the glacial landscapes were covered by dry meadows with steppic elements (mainly Poaceae, Asteraceae and *Artemisia*, *Chenopodiaceae*).

From MIS11, both the lengthening and the increasing amplitudes of glacial/interglacial cycles impacted vegetation. At the beginning and end of interglacials, the pioneer and the coniferous forests expanded as observed in interglacials older than MIS11. During the warm optimum, the mixed oak forest with rare subtropical trees dominated. However, several Mediterranean sites record mesothermic and relic taxa such as *Carya*, *Pterocarya*, *Zelkova*, and *Tsuga* in regular representation long after 450 ka, until ca. 320 ka, within MIS9 (Magri, 2010; Orain et al., 2013a, 2013b). At the same time, during interglacial periods, more Mediterranean taxa reached the composition of the Mediterranean mixed forest observed in the Holocene (Combourieu-Nebout et al., 2015). During the glacials, the lowering of temperature and precipitation favored the installation of a steppe dominated
by the presence of herbs. The development of these open environmentsis linked to the increasing
glacial dryness (Combourieu-Nebout et al., 2015).

Since the Middle Pleistocene, the Mediterranean vegetation is constrained by aridity rather than
temperature (Tzedakis, 2007). Thus the Mediterranean region has to be considered as a “warm spot”
even during glacial episodes. However, several refuge areas with permanent available moisture
enabling the persistence of the most requiring thermophilous and subtropical taxa were located over
the northern Mediterranean during the Quaternary (Bennett et al., 1991; Bennett and Provan, 2008;
Médail and Diadema, 2009; Stewart et al., 2010). Furthermore, the local persistence of trees should
be connected with faunal occurrence and human occupations in that time window to better
understand this key period for human evolution and cultural transition (Orain et al., 2013b;
Rodríguez-Rey et al., 2015).

Discussion

It is tempting to draw a parallel between hominin dispersal, cultural evolution and global/regional
climatic and environmental changes. However, we first would like to address the limitations inherent
to such an approach. Five major limiting factors should be considered:

1) The link between global climate variations and regional environment is difficult to establish for the time period between 1 Ma and 400 ka. One of the reasons is that, if global as well regional palaeoclimatic data (i.e. mainly marine records) are continuous, the palaeoenvironmental data (pollen records) from both Northern and Southern Europe remain partial, far from being continuous and do not even exist for some periods. Similarly, only a few archeological sites have been identified over this period. The scarcity of sites can be linked to the recurrence of ice sheet growth during Early and Middle Pleistocene glacials such as MIS16 and MIS12. Indeed, while ice sheet growth probably had an impact on the hominin occupations over Eurasia (Parfitt et al., 2010; Preece and Parfitt, 2012), the polar ice cap might also have erased all the northern evidence of human sites during previous interglacials. Finally, the paleoenvironmental proxies found in each archeological site (e.g. micro and macro fauna, pollens) remain often very low in
number and quality. Archaeological sites can thus only provide snapshots of local past
environmental changes.

2) Many sites are not dated precisely enough to be correlated with confidence to regional climatic and paleoenvironmental records. Indeed, only a handful of sites could be correlated with confidence to a single climatic stage based on available radio-isotopic ages. This could result from uncertainties in the different dating methods of natural and archeological deposits. For example ESR dating provides ages with large uncertainties whereas K/Ar or ³⁹Ar/⁴⁰Ar methods on tephra layers deliver ages with a narrow uncertainty. In the case of large uncertainties, the position of a human occupation cannot be strictly linked to glacial or interglacial periods solely based on the age. Furthermore, the duration of occupation (i.e. seasons, months, years?) at a specific site cannot be accessed directly. As a consequence, questions could be raised on the link between paleoenvironmental reconstructions that usually represent an average of several hundreds of years of environmental changes and archeological remains.

3) Archeological characteristics of the sites should be properly related to hominin occupation. For example the lithic as well as fauna assemblages, sometimes scarce, could be biased by the nature of the site itself (e.g. butchery site, seasonal settlement) or by the origin of the fauna (selected hunting, opportunistic butchery...) (García et al. 2009; Saladié et al., 2014).

4) Comparing local environmental data (pollen or microfauna) in an archaeological context within the global climate and vegetation frameworks require much caution in relation to methodology and taphonomy. Thus, the differential preservation of pollen in mineral sediments may strongly affect the fossil assemblages and bias the local palaeoenvironmental reconstructions (Lebreton et al., 2010). In that way, taphonomic processes (e.g. oxidation, biodegradation) have major effects on pollen preservation. Consequently, the most resistant pollen taxa can be over-
represented, which induces misinterpretation of vegetation records. This is particularly well illustrated through (i) artificial glacial pictures produced by the over-representation of Cichorioideae due to taphonomic processes or (ii) frozen images of the vegetation for long periods due to gaps in sedimentation combined with percolation of pollen grains in mineral sediments.

5) We also have to keep in mind that hominin fossils are scarce and diverse. Some hominin remains at Atapuerca (Spain) indicate a first phase which occurred at level TE9 at Sima del Elefante and levels TD3-TD4 to TD6 at Gran Dolina with *Homo antecessor* without bifaces (Rodriguez et al., 2011; van der Made et al., 2015). At around 700 ka, *Homo antecessor* is supposed to be replaced by *H. heidelbergensis* (possibly a diverse group of hominins) who is considered to be the craftsman of the bifacial technology in Europe (Wagner et al., 2011; Stringer, 2012).

Despite the numerous limitations we have listed above, several general conclusions as well as new research tracks could be highlighted using the general review of available archeological, chronological, paleoclimatic and environmental records now available.

On Northern sites, records indicate that hominin occupations took place during either dry steppic environments or woodlands (table 2). Some sites display more confident environmental data based on multiple proxies such as the British sites (Happisburg 3 and Pakefield), while others are less reliable due to the lack of well-preserved remains or alternative interpretation (Bytham River sites) (Gibbard et al., 2009; Candy et al., 2015). Some sites, such as Cagny-la-Garenne (Somme Valley), are only related to pleniglacial phases by the deposits. In this latter case, occupations could be placed either at the end of MIS 13 or at the beginning of MIS 12, but records clearly showed that hominins left the area when climatic conditions became too hard (i.e. full glacial conditions). Similarly, in la Noira (Centre France), hominins as well as fauna disappeared when very cold and dry conditions
occurred at the beginning of MIS 16, considered as one of the strongest glacial phases since 1 Ma.

The MIS 16 intensity would explain the chronological gap for hominin occupations in the Centre of France between 700 and 500 ka (« bottleneck »). The specific time period in question here is centered on the Middle Pleistocene Transition (MPT) after the Brunhes-Matuyama shift at 780 ka, which is considered to have led to an extension of the grassland habitat into higher latitudes, thereby opening and/or closing corridors for migration in these regions.

In southern Europe, the data obtained on Spanish sites suggests recurrent humid and warm environments during occupations but steppic environments are also found (Rodríguez et al., 2011). In Italy and South France, dry environments of MIS 16 or 14 are associated with occupations (Moigne et al., 2006; Barsky and Lumley, 2010; Pereira et al., 2015), suggesting a lower intensity of the glacial phases in the South compared to the North as evidenced by some regional records, such as the reconstructed Mediterranean sea surface temperatures during these periods (Lourens et al., 2004).

As explained in section 2, the shift to a 100 ka climatic cycle during the Early Pleistocene is associated with a change in the vegetation cover from almost permanent widespread forests to a well-marked alternation of forests and more open environments. This change in the vegetation cover has probably favored the expansion of hominin, according to its tolerance to climate variability, and faunal turnovers (Guthrie, 1984; Turner, 1992; Abbazzi et al., 2000; Stiner, 2002; Almogi-Labin, 2011; Bar-Yosef and Belmaker, 2011; Chapais, 2011; Cuenca-Bescos et al., 2011; Leroy et al., 2011; Rodríguez et al., 2011; Abbate and Sagri, 2012; Martínez et al., 2014). As indicated by sites older than 1 Ma, hominin occupations would have been possible only at transitions from glacial to interglacial periods during the Early Middle Pleistocene (Leroy et al., 2011). However, we have to keep in mind that the lack of records for cold periods could be also only due to taphonomical constraints and not due to the absence of occupations.

In the absence of the use of fire (Roebroeks and Villa, 2011; Gowlet, 2016), cyclical climate changes, such as the major glaciations of MIS 16 and MIS 12, would have led to successive depopulations or extinctions in the North followed by recolonization events aided by new techniques and new social
organizations (Dennell et al., 2011) or the diffusion of traditions from group to group (Manzi, 2004) during transitions to milder conditions. Current environmental and climatic data as well as chronological constraints available at archeological sites both limit models that explain the distribution of hominin occupations in Western Europe. The Mid-Brunhes Event (MBE) that occurred during MIS 11 is associated with a widespread diffusion of elaborate bifacial manufacture techniques.

The diffusion of this technology is attributed by some authors to the sporadic control of fire (Gowlett, 2006, 2016; Roebroeks and Villa, 2011). Looking at the palaeoclimatic and palaeoenvironmental records, it can also be proposed that the particular climatic features of this period (a long stable MIS 11 interglacial stage followed by the relatively short MIS 10 glacial interval) helped the expansion of occupations over the whole Western Europe (Conard et al., 2015). Indeed, after 450 ka, from MIS 11 to 9, the diversity of technological behaviours characterizes again the series, with or without bifacial tools. The subsistence strategies indicate more complex hunting scheme and new land-use patterns.

These observations are based on more series and artefacts allowing statistical comparisons and trends. Changes in the stability, amplitude and duration of climate cycles could be among the factors that explain these behavioural changes, favouring the emergence of regional traditions and technological novelties (for instance Levallois), demographic expansion and genetic exchanges (growth and regular adaptation without demographic discontinuities) and new skills (see examples of more or less rapid processes of population dynamics and cultural changes) (Lamotte and Tuffreau, 2001a, b; Hölzchen et al., 2015; Peretto et al., 2015b; Sánchez Yustos and Diez Martín, 2015; Vaesen et al., 2016). However, we do not exclude the possibility of hominin expansion before 500 ka in northern Europe in favour of centennial or yearly scale climate variations that we are not able to perceive through existing records. That would explain why fire mastery was not necessarily useful to hominin groups to occupy the Northern territories.

**Conclusion**
To conclude, despite the wealth of climatic and environmental information that are now available at the local or global scales showing the evolution of environmental and climatic gradients over the last million years, it is still hazardous to make the link between climate, environment and hominin occupations. Information is still missing in order (1) to determine the palaeoenvironmental and climatic factors that may have influenced or regulated the hominin occupations, associated pathways and timing of dispersals, and (2) to understand how hominin activities and behaviours could have been influenced by the climatic and environmental changes (McPherron, 2000; Goren-Inbar and Sharon, 2006).

Sporadic hominin sites between 900 and 500 ka associated with large dating uncertainties and lack of continuous climatic/environmental proxies records in Europe over this period do not currently allow one to relate hominin occupations to clear climatic constraints. After MIS 12, it has been observed that demographic expansion over Western Europe is related to new skills despite the lack of evidence of fire before 400 ka. However, the influence of climate and environment is not clearly identified, expansion being either related to micro-climatic phases or diffusion of the bifacial technology by new populations adapted to these areas. Progress in dating methods, further acquisitions of continuous records of the climatic gradients for the last 1 Ma in Europe; improvement of climate and biosphere modelling are necessary to demonstrate the eventual role of climate on hominin occupations and phases of expansion.

## Acknowledgements

This study was supported by the French-British project ANR (PremAcheuSep) devoted to the earliest evidence of occupations with bifacial technology in North-West Europe. It was also supported by the Department of Prehistory, National Museum of Natural History, Laboratoire des Sciences du Climat et de l'Environnement, LSCE/IPSL, CEA-CNRS-UVSQ, the University Paris-Saclay, France.

We would like to thank the Editor and the five reviewers who helped us to enrich and improve the paper.
References

Abbate, E., Sagri, M., 2012. Early to Middle Pleistocene Homo dispersals from Africa to Eurasia: Geological, climatic and environmental constraints. Quaternary International 267, 3–19.

Abbazzi, L., Fanfani, F., Ferretti, M.P., Rook, L., Cattani, L., Masini, F., Mallegni, F., Negrino, F., Tozzi, C., 2000. New Human Remains of Archaic Homo sapiens and Lower Palaeolithic Industries from Visogilano (Duino Aurisina, Trieste, Italy). Journal of Archaeological Sciences 27, 1173–1186.

Allen, J.R.M., Brandt, U., Brauer, A., Hubberten, H.-W., Huntley, B., Keller, J., Kraml, M., Mackensen, A., Mingram, J., Negendank, J.F.W., 1999. Rapid environmental changes in southern Europe during the last glacial period. Nature 400, 740–743.

Allen, J.R.M., Watts, W.A., Huntley, B., 2000. Weichselian palynostratigraphy, palaeovegetation and palaeoenvironment; the record from Lago Grande di Monticchio, southern Italy. Quaternary International 73, 91–110.

Almogi-Labin, A., 2011. The paleoclimate of the Eastern Mediterranean during the transition from early to mid Pleistocene (900 to 700 ka) based on marine and non-marine records: An integrated overview. Journal of Human Evolution 60, 428–436.

Antoine, P., Limondin-Lozouet, N., Chaussé, C., Lautridou, J.P., Pastre, J.F., Auguste, P., Bahain, J.J., Falguères, C., Galehb, B., 2007. Pleistocene fluvial terraces from northern France (Seine, Yonne, Somme) : synthesis and new results. Quaternary Science Reviews 26, 2701–2723.

Antoine, P., Moncel, M.-H., Locht, J.-L., Limondin-Lozouet, N., Auguste, P., Stoetzel, E., Dabkowski, J., Voinchet, P., Bahain, J.-J., Falgueres, C., 2015. Dating the earliest human occupation of Western Europe: New evidence from the fluvial terrace system of the Somme basin (Northern France). Quaternary International 370, 77–99.
Ashton, N.M., Cook, J., Lewis, S.G., Rose, J., 1992. High Lodge, Excavations 1962-8 by G. de G. Sieveking and J. Cook, 1988. British Museum Press.

Ashton, N., McNabb, J., Irving, B., Lewis, S., Parfitt, S., 1994. Comtemporaneity of Clactonian and Acheulian flint industries at Barnham, Suffolk. Antiquity 68, 585-589.

Ashton, N., Lewis, J.E., Hosfield, R., 2011. Mapping the Human Record: Population Change in Britain during the Early Palaeolithic, in: Ashton N., Lewis J.E., Stringer C. (Eds.), The Ancient Human Occupation of Britain. Quaternary Science.

Ashton, N., Lewis, S.G., 2012. The environmental contexts of early human occupation of northwest Europe: The British Lower Palaeolithic record. Quaternary International 271, 50-64.

Ashton, N., Lewis, S.G., Groote, I. De, Duffy, S.M., Bates, M., Bates, R., Hoare, P., Lewis, M., Parfitt, S.A., Peglar, S., Williams, C., Chris Stringer, 2014. Hominin Footprints from Early Pleistocene Deposits at Happisburgh, UK. PloS One 9, e88329.

Ashton, N., Scott, B., 2015. The British Middle Palaeolithic. Quaternary International doi:http://dx.doi.org/10.1016/j.quaint.2015.06.011

Barsky, D., 2013. The Caune de l'Arago stone industries in their stratigraphical context. Comptes Rendus Palevol 12, 305-325. Barsky, D., de Lumley, H., n.d. Early European Mode 2 and the stone industry from the Caune de l'Arago's archeostratigraphical levels "P" Quaternary International 223-224, 71-86.

Barsky, D., Lumley, de H., 2010. Early European Mode 2 and the Stone Industry from the Caune de l'Arago's Archeostratigraphical Levels "P." Quaternary International 223, 71-86.

Bar-Yosef, O., Belmaker, M., 2011. Early and Middle Pleistocene Faunal and hominins dispersals through Southwestern Asia. Quaternary Science Reviews 30, 1318-1337.

Bazin, L., Landais, A., Lemieux-Dudon, B., Toyé Mahamadou Kele, H., Veres, D., Parrenin, F.,
Martinerie, P., Ritz, C., Capron, E., Lipenkov, V., Loutre, M.-F., Raynaud, D., Vinther, B., Svensson, A., Rasmussen, S., Severi, M., Blunier, T., Leuenberger, M., Fischer, H., Masson-Delmotte, V., Chappellaz, J., Wolff, E., 2013. An optimized multi-proxies, multi-site Antarctic ice and gas orbital chronology (AICC2012): 120-800 ka. Clim. Past 9, 1715-1731.

Beaulieu de, J.-L., Reille, M., 1984. A long upper Pleistocene pollen record from Les Echets, near Lyon, France. Boreas 13, 111-132.

Beaulieu, J.-L. de, Suc, J.-P., 1985. Les pollens et l'histoire de la végétation. Histoire archéologie 93, 67-73.

Beaulieu, J.-L. de, Reille, M., 1989. The transition from temperate phases to stadials in the long upper Pleistocene sequence from Les Echets (France). Paleogeography, Paleoclimatology, Paleoecology. 72, 147-159.

Beaulieu, J.-L., de Reille, M., 1992a. The last climatic cycle at La Grande Pile (Vosges, France) a new pollen profile. Quaternary Science. Reviews 11, 431-438.

Beaulieu, J.-L. de, Reille, M., 1992b. Long Pleistocene pollen sequences from the Velay Plateau (Massif Central, France). Vegétation Histoire Archaeobotanique 1, 233-242.

Beaulieu, J.-L. de, Andrieu-Ponel, V., Cheddadi, R., Guiter, F., Ravazzi, C., Reille, M., Rossi, S., 2006. Apport des longues séquences lacustres à la connaissance des variations des climats et des paysages pléistocènes. Comptes Rendus Palevol 5, 65-72.

Bender, M., Sowers, T., Labeyrie, L., 1994. The Dole Effect and its variations during the last 130,000 years as measured in the Vostok Ice Core. Global Biogeochemistry Cycles 8(3), 363-376.

Bennàsar, M., Cáceres, I. and Cuenca-Bescós, G., 2016. Paleoecological and microenvironmental aspects of the first European hominids inferred from the taphonomy of small mammals (Sima del Elefante, Sierra de Atapuerca, Spain). Comptes Rendus Palevol, in press.
Bennett, K.D., Tzedakis, P.C., Willis, K.J., 1991. Quaternary refugia of north European trees. Journal of Biogeography 103–115.

Bennett, K.D., Provan, J., 2008. What do we mean by “refugia”? Quaternary Science Reviews 27, 2449–2455.

Berger, GW, Pérez-González, A, Carbonell, E, Arsuaga, JL, Bermúdez de Castro, JM, Ku T-L, 2008. Luminescence chronology of cave sediments at the Atapuerca paleoanthropological site, Spain. Journal of Human Evolution 55(2), 300-311

Bermúdez de Castro, J.M., Martinón-Torres, M., Gómez-Robles, A., Prado-Simón, L., Martín-Francés, L., Lapresa, M., Olejniczak, A., Carbonell, E., 2011. Early Pleistocene human mandible from Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain): A comparative morphological study. Journal of Human Evolution 61, 12–25.

Bertini, A., 2000. Pollen record from Colle Curti and Cesi: early and middle Pleistocene mammal sites in the Umbro–Marchean Apennine Mountains (central Italy). Journal of Quaternary Sciences 15, 825–840.

Bertini, A., 2010. Pliocene to Pleistocene palynoflora and vegetation in Italy: state of the art. Quaternary International 225, 5–24.

Bertini, A., Toti, F., Marino, M., Ciaranfi, N., 2015. Vegetation and climate across the Early–Middle Pleistocene transition at Montalbano Jonico, southern Italy. Quaternary International doi:http://dx.doi.org/10.1016/j.quaint.2015.01.003

Beyene, Y., Katoh S., WoldeGabriel G., Hart W.K., Sudo M., Kondo M., Hyodo M., Renne P.R., Suwa G., Asfaw B., 2013. The characteristics and chronology of the earliest Acheulean at Konso, Ethiopia. PNAS 110, 1584–1591.

Blain, H. -A., Cuenca-Bescos, G., Burjachs, F., Lopez-Garcia, J., Lozano-Fernandez, I., Rosell, J., 2013.
Early Pleistocene palaeoenvironments at the time of the Homo antecessor settlement in the
Gran Dolina cave (Atapuerca, Spain). Journal of Quaternary Sciences 28, 311–319.

Bridgland, D.R., Antoine, P., Limondin-Lozouet, N., Santisteban, J.I., Westaway, R., White, M.J., 2006.
The Palaeolithic occupation of Europe as revealed by evidence from the rivers: data from ICGP
449. Journal of Quaternary Sciences 21, 437–456.

Bruguès, R.J., Suc, J.-P., 1980. Analyse pollinique des dépôts lacustres du Pléistocène inférieur de Banyoles (Bañolas, site de la Bòbila Ordis-Espagne): un élément nouveau dans la reconstitution de l'histoire paléoclimatique des régions méditerranéennes d'Europe occidentale. Geobios 13, 5–19.

Candy, I., Schreve, D., White, T.S., 2015. MIS 13–12 in Britain and the North Atlantic: understanding the palaeoclimatic context of the earliest Acheulean. Journal of Quaternary Sciences 30(7), 593–610.

Carbonell, E., Garcí, M., Mallol, C., Mosquera, M., Ollé, A., Sahnouni, M., Sala, R. and Vergs, J.M.,
1999. The TD6 level lithic industry from Gran Dolina, Atapuerca (Burgos, Spain): production and
use. Journal of Human Evolution 37(3), 653-693.

Carbonell, E., De Castro, J.B., Arsuaga, J.L., Allue, E., Bastir, M., Benito, A., Cacères, I., Canals, T., Díez, J.C., van der Made, J. and Mosquera, M., 2005. An early Pleistocene hominin mandible from Atapuerca-TD6, Spain. Proceedings of the National Academy of Sciences of the United States of America, 102(16), 5674–5678.

Carbonell, E., Sala Ramos, R., Rodríguez, X.P., Mosquera, M., Ollé, A., Vergès, J.M., Martínez-Navarro, B., Bermúdez de Castro, J.M., 2010. Early hominid dispersals: A technological hypothesis for "out of Africa." Quaternary International 223-224, 36–44.

Carrión, J.S., Rose, J., Stringer, C., 2011. Early Human Evolution in the Western Palaearctic: Ecological
Scenarios. Quaternary Science Reviews 30, 1281–1295.

Channell, J.E.T., Hodell, D. a., Romero, O., Hillaire-Marcel, C., de Vernal, a., Stoner, J.S., Mazaud, a., Röhl, U., 2012. A 750-kyr detrital-layer stratigraphy for the North Atlantic (IODP Sites U1302-U1303, Orphan Knoll, Labrador Sea). Earth Planaery Sciences Letter. 317-318, 218–230.

Chapais B., 2011. The Deep Social Structure of Humankind. Science (80). 331, 1276–1277.

Chappellaz, J., 2007. Encyclopedia of Quaternary Science, Encyclopedia of Quaternary Science.
Elsevier.

Clark, P.U., Archer, D., Pollard, D., Blum, J.D., Rial, J.A., Brovkin, V., Mix, A.C., Pisias, N.G., Roy, M., 2006. The middle Pleistocene transition: characteristics, mechanisms, and implications for long-term changes in atmospheric pCO2. Quaternary Science Reviews 25, 3150–3184.

Clark, P.U., Dyke, A.S., Shakun, J.D., Carlson, A.E., Clark, J., Wohlfarth, B., Mitrovica, J.X., Hostetler, S.W., McCabe, A.M., 2009. The Last Glacial Maximum. Science 80-325, 710–714.

Cohen, K.M., MacDonald, K., Joordens, J.C., Roebroeks, W., Gibbard, P.L., 2012. The earliest occupation of north-west Europe: a coastal perspective. Quaternary International 271, 70–83.

Coltorti, M., Feraud, G., Marzoli, A., Peretto, C., Ton-Thate, T., Voinchet, P., Bahain, J.J., Minelli, A., Thun Hohenstain, U., 2005. New 40Ar/39Ar, stratigraphic ans paleoclimatic data on the Isernia La Pineta Lower Palaeolithic site, Molise, Italy. Quaternary International 131, 11–22.

Combourieu-Nebout, N., Bertini, A., Russo-Ermolli, E., Peyron, O., Klotz, S., Montade, V., Fauquette, S., Allen, J.R.M., Fusco, F., Goring, S., 2015. Climate changes in the central Mediterranean and Italian vegetation dynamics since the Pliocene. Revue de Palaeobotanique Palynolologie.

Conard, N.J., Serangeli, J., Böhner, U., Starkovich, B.M., Miller, C.E., Urban, B., Van Kolfschoten, T., 2015. Excavations at Schöningen and paradigm shifts in human evolution. Journal of human evolution 89, 1–17.
Corrado, P., Magri, D., 2011. A late Early Pleistocene pollen record from Fontana Ranuccio (central Italy). Journal of Quaternary Sciences 26, 335–344.

Cuenca-Bescós, G., Melero-Rubio, M., Rofes, J., Martínez, I., Arsuaga, J.L., Blain, H.A., López-García, J.M., Carbonell, E., Bermudez de Castro, J.M., 2011. The Early-Middle Pleistocene environmental and climatic change and the human expansion in Western Europe: A case study with small vertebrates (Gran Dolina, Atapuerca, Spain). Journal of Human Evolution 60 (4), 481–491.

Cuenca-Bescós, G., Rofes, J., López-García, J.M., Blain, H.A., Rabal-Garcés, R., Sauqué, V., Arsuaga, J.L., de Castro, J.M.B., Carbonell, E., 2013. The small mammals of Sima del Elefante (Atapuerca, Spain) and the first entrance of Homo in Western Europe. Quaternary International 295, 28-35.

Cuenca-Bescós, G., Blain, H.A., Rofes, J., López-García, J.M., Lozano-Fernández, I., Galán, J., Núñez-Lahuerta, C., 2016. Updated Atapuerca biostratigraphy: Small-mammal distribution and its implications for the biochronology of the Quaternary in Spain. Comptes Rendus Palevol.

de la Torre I., 2011. The Early Stone Age lithic assemblages of Gadeb (Ethiopia) and the Developed Oldowan/early Acheulean in East Africa. Journal of Human Evolution 60, 768–812.

de Lombera-Hermida, A., Bargalló, A., Terradillos-Bernal, M., Huguet, R., Vallverdú, J., García-Antón, M.-D., Mosquera, M., Ollé, A., Sala, R., Carbonell, E., Rodríguez-Álvarez, X.-P., 2015. The lithic industry of Sima del Elefante (Atapuerca, Burgos, Spain) in the context of Early and Middle Pleistocene technology in Europe. Journal of Human Evolution 82, 95-106

Delmonte, B., Andersson, P.S., Hansson, M., Schöberg, H., Petit, J.R., Basile-Doelsch, I., Maggi, V., 2008. Aeolian dust in East Antarctica (EPICA-Dome C and Vostok): Provenance during glacial ages over the last 800 kyr. Geophysic Research Letter 35, 2–7.

Dennell, R.W., Martinón-Torres, M., Bermúdez de Castro, J.M., 2010. Out of Asia: The initial colonisation of Europe in the Early and Middle Pleistocene. Quaternary International 223-224,
Dennell, R.W., Martinón-Torres, M., Bermúdez de Castro, J.M., 2011. Hominin variability, climatic instability and population demography in Middle Pleistocene Europe. Quaternary Science Reviews 30, 1511–1524.

Dreyfus, G. B., Parrenin, F., Lemieux-Dudon, B., Durand, G., Masson-Delmotte, V., Jouzel, J., Barnola, J.-M., Panno, L., Spahni, R., Tisserand, A., Siegenthaler, U., Leuenberger, M., 2007. Anomalous flow below 2700 m in the EPICA Dome C ice core detected using delta O-18 of atmospheric oxygen measurements. Climate Past 3, 341–353.

Dreyfus, G. B., Raisbeck, G. M., Parrenin, F., Jouzel, J., Guyodo, Y., Nomade, S., Mazaud, A., 2008. An ice core perspective on the age of the Matuyama-Brunhes boundary. Earth Planetary Sciences Letter 274, 151–156.

Despriée Jackie, Pierre Voinchet, Hélène Tissoux, Jean-Jacques Bahain, Christophe Falguères, Gilles Courcimault, Jean Dépont, Marie-Hélène Moncel, Sophie Robin, Marta Arzarello, Robert Sala, Laurent Marquer, Erwan Messager, Simon Puaud, Salah Abdessadok, 2011. Lower and Middle Pleistocene human settlements recorded in fluvial deposits of the middle Loire River Basin, Centre Region, France. Quaternary Science Reviews 1474–1485.

Drescher-Schneider, R., 2000. Die Vegetations-und Klimaentwicklung im Riss/Würm-Inter-Glazial und im Früh-und Mittelwürm in der Umgebung von Mondsee. Ergebnisse der pollenanalytischen Untersuchungen.

Elderfield, H., Ferretti, P., Greaves, M., Crowhurst, S., McCave, I.N., Hodell, D., Piotrowski, a. M., 2012. Evolution of Ocean Temperature and Ice Volume through the Mid-Pleistocene Climate Transition. Science 308, 704–709.

EPICA community Members, 2004. Eight glacial cycles from an Antarctic ice core. Nature 429, 623–
Ermolli, E.R., Cheddadi, R., 1997. Climatic reconstruction during the Middle Pleistocene: a pollen record from Vallo di Diano (Southern Italy). Geobios 30, 735-744.

Falguères, C., Shao, Q., Han, F., Bahain, J.J., Richard, M., Perrenoud, C., Moigne, A.M., De, H.L., 2015. New ESR and U-series dating at Caune de l'Arago, France: A key-site for European Middle Pleistocene. Quaternary Geochronology 30, 547-553.

Fletcher, W.J., Sánchez-Gòni, M.F., Allen, J.R.M., Cheddadi, R., Combourieu-Nebout, N., Huntley, B., Lawson, I., Londeix, L., Magri, D., Margan, V., Muller, U.C., Naughton, F., Novenko, E., Roucoux, K., Tzedakis, P.C., 2010. Millenial-scale variability during the last glacial in vegetation records from Europe. Quaternary Science Reviews 29, 2839-2864.
Florschütz, F., Amor, J.M., Wijmstra, T.A., 1971. Palynology of a thick Quaternary succession in southern Spain. Palaeogeography Palaeoclimatology Palaeoecology 10, 233-264.

Follieri, M., Magri, D., Sadori, L., 1988. 250000-year pollen record from Vallee di Castiglione (Roma). Pollen et Spores 30, 329-356.

Follieri, M., Giardini, M., Magri, D., Sadori, L., 1998. Palynostratigraphy of the last glacial period in the volcanic region of central Italy. Quaternary International 47, 3-20.

Gallotti, R., Peretto, C., 2015. The Lower/early Middle Pleistocene small débitage productions in Western Europe: New data from Isernia La Pineta t. 3c (Upper Volturno Basin, Italy). Quaternary International 357, 264-281.

García, N.G., Feranec, R.S., Arsuaga, J.L., de Castro, J.B., Carbonell, E., 2009. Isotopic analysis of the ecology of herbivores and carnivores from the Middle Pleistocene deposits of the Sierra De Atapuerca, northern Spain. Journal of Archaeological Science 36(5), 1142-1151.

García-Medrano, P., Ollé, A., Mosquera, M., Cáceres, I., Díez, C., Carbonell, E., 2014. The earliest
Acheulean technology at Atapuerca (Burgos, Spain): Oldest levels of the Galería site (GII Unit).
Quaternary International 353, 170–194.

Gibbard, P.L., Pasanen, A.H., West, R.G., Lunkka, J.P., Boreham, S., Cohen, K.M., Rolfe, C., 2009. Late Middle Pleistocene glaciation in East Anglia, England. Boreas 38(3), 504-528.

Goren-Inbar N., Sharon G., 2006. Axe Age: Acheulian Tool-making from Quarry to Discard. Equinox Publishing Ltd, London.

Gowlett, J.A.J., 2006. The early settlement of northern Europe: Fire history in the context of climate change and the social brain, In: Colloque « Climats-Cultures-Sociétés Aux Temps Préhistoriques. De L'apparition Des Hominidés Jusqu'au Néolithique ». Comptes Rendus Palevol, pp. 299-311.

Gowlett, J.A.J., 2016. The discovery of fire by humans: a long and convoluted process. Phil. Trans. R. Soc. B, 371(1696), 20150164.

Guthrie, R.D., 1984. Mosaics, allelochemics and nutrients: an ecological theory of Late Pleistocene megafaunal extinctions, in: Martin, P.S., Klein, R.G. (Eds.), Quaternary Extinctions: A Prehistoric Revolution. University of Arizona Press, pp. 259–298.

Hahne, J., Ellwanger, D., Stritzke, R., 2008. Evidence for a Waalian thermomer pollen record from the research borehole Heidelberg UniNord, Upper Rhine Graben, Baden-Württemberg. Eiszeit. Gegenw. Journal of Quaternary Sciences 57, 403–410.

Hodell, D.A., Channeil, J.E.T., Curtis, J.H., Romero, O.E., Röhl, U., Channell, J.E.T., Curtis, J.H., Romero, O.E., Röhl, U., 2008. Onset of “Hudson Strait” Heinrich events in the eastern North Atlantic at the end of the middle Pleistocene transition (~640 ka)? Paleoceanography 23, 1–16.

Hölzchen, E., Hertler, C., Timm, I., Lorig, F., 2015. Evaluation of Out of Africa hypotheses by means of agent-based modeling. Quaternary International
doi:http://dx.doi.org/10.1016/j.quaint.2015.11.022
Hosfield, R. 2011. The British Lower Palaeolithic of the early Middle Pleistocene. Quaternary Science Reviews 30, 1486–1510.

Huguet, R., Saladié, P., Cáceres, I., Díez, C., Rosell, J., Bennàsar, M., Blasco, R., Esteban-Nadal, M., Gabucio, M.J., Rodríguez-Hidalgo, A., Carbonell, E., 2013. Successful subsistence strategies of the first humans in south-western Europe. Quaternary International 295, 168-182.

Jiménez-Arenas, J.M., Santonja, M., Botella, M., Palmqvist, P., 2011. The oldest handaxes in Europe: fact or artefact? Journal of Archaeological Sciences 38, 3340–3349.

Joannin, S., Cornée, J.-J., Moissette, P., Suc, J.-P., Koskeridou, E., Lécuyer, C., Buisine, C., Kouli, K., Ferry, S., 2007. Changes in vegetation and marine environments in the eastern Mediterranean (Rhodes, Greece) during the Early and Middle Pleistocene. Journal of Geolological Society London. 164, 1119–1131.

Joannin, S., Ciaranfi, N., Stefanelli, S., 2008. Vegetation changes during the late Early Pleistocene at Montalbano Jonico (Province of Matera, southern Italy) based on pollen analysis. Palaeogeography Palaeoclimatology Palaeoecology 270, 92–101.

Joannin, S., Bassinot, F., Combourien-Nebout, N., Peyron, O., Beaudouin, C., 2011. Vegetation response to obliquity and precession forcing during the Mid-Pleistocene Transition in Western Mediterranean region (ODP site 976). Quaternary Science Reviews 30, 280–297.

Jouzel, J., 2003. Magnitude of isotope/temperature scaling for interpretation of central Antarctic ice cores. Journal of Geophysic Research 108, 4361.

Jouzel, J., Masson-Delmotte, V., Cattani, O., Dreyfus, G., Falourd, S., Hoffmann, G., Minster, B., Nouet, J., Barnola, J.M., Chappellaz, J., Fischer, H., Gallet, J.C., Johnsen, S., Leuenberger, M., Louergue, L., Luethi, D., Oerter, H., Parrenin, F., Raisbeck, G., Raynaud, D., Schilt, A., Schwander, J., Selmo, E., Souchez, R., Spahni, R., Stauffer, B., Steffensen, J.P., Stenni, B., Stocker, T.F., Tison, J.L.,
Werner, M., Wolff, E.W., 2007. Orbital and Millennial Antarctic Climate Variability over the Past 800,000 Years. Sciences 317, 793–796.

Klotz, S., Fauquette, S., Combourieu-Nebout, N., Uhl, D., Suc, J.-P., Mosbrugger, V., 2006. Seasonality intensification and long-term winter cooling as a part of the Late Pliocene climate development. Earth Planetary Sciences Letter 241, 174–187.

Knipping, M., 2008. Early and Middle Pleistocene pollen assemblages of deep core drillings in the northern Upper Rhine Graben, Germany. Netherlands Journal Geosciences en Mijnb 87, 51.

Lambert, F., Delmonte, B., Petit, J. R., Bigler, M., Kaufmann, P. R., Hutterli, M. A., Stocker, T. F., Ruth, U., Steffensen, J. P., Maggi, V., 2008. Dust-climate couplings over the past 800,000 years from the EPICA Dome C ice core. Nature 452, 616–619.

Lambert, F., Bigler, M., Steffensen, J.P., Hutterli, M., Fischer, H., 2012. Centennial mineral dust variability in high-resolution ice core data from Dome C, Antarctica. Climate Past 8, 609–623.

Lamotte, A., Tuffreau, A., 2001a. Les industries lithiques de Cagny-la-Garennes II (Somme, France), in: (ed.), T.A. (Ed.), L'Acheuléen Dans La Vallée de La Somme et Paléolithique Moyen Dans Le Nord de La France : Données Récentes. CERP, pp. 59–91.

Lamotte A., Tuffreau A., 2001b. Les industries acheuléennes de Cagny (Somme) dans le contexte de l'Europe du Nord-ouest, in: Tuffreau, A. (Ed.), L'Acheuléen Dans La Vallée de La Somme et Paléolithique Moyen Dans Le Nord de La France : Données Récentes. CERP, pp. 149–153.

Landais, a., Dreyfus, G., Capron, E., Masson-Delmotte, V., Sanchez-Goñi, M.F., Desprat, S., Hoffmann, G., Jouzel, J., Leuenberger, M., Johnsen, S., 2010. What drives the millennial and orbital variations of δ18O atm? Quaternary Science Reviews 29, 235–246.

Lamotte, A., Tuffreau, A., 2015. Acheulean of the Somme basin (France): Assessment of lithic changes during MIS 12 to 9. Quaternary International
Lebreton, V., 2002. Végétation et climat au Pléistocène inférieur et moyen à La Pineta (Isernia, Italie).
Comptes Rendus Palevol 1, 11–17.

Lebreton, V., 2004. Paysages et climats des premiers hominidés en Italie. Bar, International Series, 1204.

Lebreton, V., Messager, E., Marquer, L., Renault-Miskovsky, J., 2010. A neotaphonomic experiment in pollen oxidation and its implication for archaeopalynology. Review of Palaeobotany and Palynology 162, 29-38.

Lefèvre, D., Raynal, J.-P., Vernet, G., Kieffer, G., Piperno, M., 2010. Tephro-stratigraphy and the age of ancient Southern Italian Acheulean settlements: The sites of Loreto and Notarchirico (Venosa, Basilicata, Italy). Quaternary International 223-224, 360–368.

Lemieux-Dudon, B., Blayo, E., Petit, J.R., Waelbroeck, C., Svensson, A., Ritz, C., Barnola, J.M., Narcisi, B.M., Parrenin, F., 2010. Consistent dating for Antarctic and Greenland ice cores. Quaternary Science Reviews 29, 8–20.

Lepre C. L., Roche H., Kent D. V., Harmand S., Quinn R.L., Brugal J-P., Texier P-J., Lenoble A., Feibel C., 2011. An earlier origin for the Acheulian. Nature 477, 82–85.

Leroy, S., 1990. Paléoclimats plio-pléistocènes en Catalogne et Languedoc d'après la palynologie de formations lacustres. Thèse de l'Université Catholique, Louvain 1, 319.

Leroy, S., Ambert, P., Suc, J.-P., 1994. Pollen record of the Saint-Macaire maar (Hérault, southern France): a Lower Pleistocene glacial phase in the Languedoc coastal plain. Revue de Palaeobotanique Palynologie 80, 149–157.

Leroy, S.A.G., Giralt, S., Francus, P., Seret, G., 1996. The high sensitivity of the palynological record in the Vico maar lacustrine sequence (Latium, Italy) highlights the climatic gradient through
Europe for the last 90 ka. Quaternary Science Reviews 15, 189–201.

Leroy, S.A.G., Arpe, K., Mikolajewicz, U., 2011. Vegetation context and climatic limits of the Early Pleistocene hominin dispersal in Europe. Quaternary Science Reviews 30(11), 1448-1463.

Lindner, L., Marks, L., 2015. Early and Middle Pleistocene fluvial series in northern foreland of the Carpathians (Poland and Ukraine) and their relation to Dnistr River terraces. Quaternary International 357, 22–32.

Lisiecki, L.E., Raymo, M.E., 2005. A Pliocene-Pleistocene stack of 57 globally distributed benthic ?? 18O records. Paleoceanography 20, 1–17.

Loulergue, L., Schilt, A., Spahni, R., Masson-Delmotte, V., Blunier, T., Lemieux, B., Barnola, J.-M., Raynaud, D., Stocker, T.F., Chappellaz, J., 2008. Orbital and millennial-scale features of atmospheric CH4 over the past 800,000 years. Nature 453, 383–386.

Lourens, L.J., 2004. Revised tuning of Ocean Drilling Program Site 964 and KC01B (Mediterranean) and implications for the δ 18 O, tephra, calcareous nannofossil, and geomagnetic reversal chronologies of the past 1.1 Myr. Paleoceanography 19, PA3010, p. 1-20

Lüthi, D., Le Floch, M., Bereiter, B., Blunier, T., Barnola, J.-M., Siegenthaler, U., Raynaud, D., Jouzel, J., Fischer, H., Kawamura, K., Stocker, T.F., 2008. High-resolution carbon dioxide concentration record 650,000-800,000 years before present. Nature 453, 379–382.

MacDonald, K., Martinón-Torres, M., Dennell, R.W., Bermúdez de Castro, J.M., 2012. Discontinuity in the record for hominin occupation in south-western Europe: Implications for occupation of the middle latitudes of Europe. Quaternary International 271, 84–97.

Magri, D., 1999. Late Quaternary vegetation history at Lagaccione near Lago di Bolsena (central Italy). Revue de Palaeobotanique Palynolologie 106, 171–208.

Magri, D., 2010. Persistence of tree taxa in Europe and Quaternary climate changes. Quaternary
International 219, 145–151.

Magri, D., Palombo, M.R., 2013. Early to Middle Pleistocene dynamics of plant and mammal communities in South West Europe. Quaternary International 288, 63–72.

Magri, D., Sadori, L., 1999. Late Pleistocene and Holocene pollen stratigraphy at Lago di Vico, central Italy. *Végétation Histoire Archaeobotanique* 8, 247–260.

Manzi, G., 2004. Human Evolution at the Matuyama-Brunhes boundary. *Evolutionary Anthropology* 13, 11–24.

Manzi, G., Magri, D., Palombo, M.R., 2011. Early-Middle Pleistocene environmental changes and human evolution in the Italian peninsula. *Quaternary Science Reviews* 30, 1420–1438.

Martínez, K., Garcia, J., Burjachs, F., Yll, R., Carbonell, E., 2014. Early human occupation of Iberia: the chronological and palaeoclimatic inferences from Vallparadís (Barcelona, Spain). *Quat. Sci. Rev.* 85, 136–146. Martinón-Torres, M., Dennell, R., Bermúdez de Castro, J.M., 2011a. The Denisova hominin need not be an out of Africa story. *Journal of Human Evolution* 60, 251–255.

Martinón-Torres, M., Dennell, R. and Bermúdez de Castro, J.M., 2011a. The Denisova hominin need not be an out of Africa story. *Journal of Human Evolution*, 60(2), 251-255.

Martinón-Torres, M., Martín-Francés, L., Gracia, A., Olejniczak, A., Prado-Simón, L., Gómez-Robles, A., Lapresa, M., Carbonell, E., Arsuaga, J.L., Bermúdez de Castro, J.M., 2011b. Early Pleistocene human mandible from Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain): A palaeopathological study. *Journal of Human Evolution* 61, 1–11.

Masson-Delmotte, V., Stenni, B., Pol, K., Braconnot, P., Cattani, O., Falourd, S., Kageyama, M., Jouzel, J., Landais, a., Minster, B., Barnola, J.M., Chappellaz, J., Krinner, G., Johnsen, S., Röthlisberger, R., Hansen, J., Mikolajewicz, U., Otto-Bliesner, B., 2010. EPICA Dome C record of glacial and interglacial intensities. *Quaternary Science Reviews* 29, 113–128.
McNabb, J., 1996. More from the cutting edge: further discoveries of Clactonian bifaces. Antiquity 70, 428-436.

McPherron, S.P., 2000. Handaxes as a measure of the mental capabilities of early hominids. Journal of Archaeolological Sciences 27, 655-663.

Médail, F., Diadema, K., 2009. Glacial refugia influence plant diversity patterns in the Mediterranean Basin. Journal of Biogeography 36, 1333-1345.

Meyer, M., Fu, Q., Aximu-Petri, A., Glocke, I., Nickel, B., Arsuaga, J.-L., Martínez, I., Gracia, A., de Castro, J.M.B., Carbonell, E., 2014. A mitochondrial genome sequence of a hominin from Sima de los Huesos. Nature 505, 403-406.

Moigne, A.-M., Palombo, M.R., Belda, V., Heriech-Briki, D., Kacimi, S., Lacombat, F., Lumley, de M.-A., Moutoussamy, J., Rivals, F., Quiles, J., Testu, A., 2006. Les faunes de grands mamifères de la Caune de l'Arago (Tautavel) dans le cadre biochronologique des faunes du Pléistocène moyen italien. L'Anthropologie. 110, 788-831.

Moncel, M.-H., Despriée, J., Voinchet, P., Tissoux, H., Moreno, D., Bahain, J.-J., Courcimault, G., Falguères, C., 2013. Early Evidence of Acheulean Settlement in Northwestern Europe-La Noira Site, a 700 000 Year-Old Occupation in the Center of France. PloS One 8(11), e75529.

Moncel, M.-H., Ashton, N., Lamotte, A., Tuffreau, A., Cliquet, D., Despriée, J., 2015a. The Early Acheulian of north-western Europe. J. Anthropol. Archaeol. 40, 302-331.

Moncel, M.-H., Despriée, J., Voinchet, P., Courcimault, G., Hardy, B., Bahain, J.-J., Puaud, S., Gallet, X., Falguères, C., 2015b. The Acheulean workshop of la Noira (France, 700 ka) in the European technological context. Quaternary International 393, 112-136.

Moncel M-H., Arzarello M., Boëda E., Bonilauri S., Chevrier B., Gaillard C., Forestier H., Li Yinghua, Sémah F., Zeitoun V. 2016a. The assemblages with bifacial tools in Eurasia (first part). What is
going on in the West? Data on Western and Southern Europe and the Levant, Comptes Rendu Palevol, special issue Eurasian Pleistocene, in press.

Moncel M-H., Arzarello M., Boëda E., Bonilauri S., Chevrier B., Gaillard C., Forestier H., Li Yinghua, Sémah F., Zeitoun V. 2016b. Assemblages with bifacial tools in Eurasia (third part). Considerations on the bifacial phenomenon throughout Eurasia, Comptes Rendu Palevol, special issue Eurasian Pleistocene, in press.

Moreno, D., Falguères, C., Pérez-González, A., Voinchet, P., Ghaleb, B., Despriée, J., Bahain, J.-J., Sala, R., Carbonell, E., de Castro, J.M.B., Arsuaga, J.L., 2015. New radiometric dates on the lowest stratigraphical section (TD1 to TD6) of Gran Dolina site (Atapuerca, Spain). Quaternary Geochronolology 30, Part B, 535-540.

Moscariello, A., Ravazzi, C., Brauer, A., Mangili, C., Chiesa, S., Rossi, S., de Beaulieu, J.-L., Reille, M., 2000. A long lacustrine record from the Pianico-Sellere basin (middle-late Pleistocene, Northern Italy). Quaternary International 73, 47-68.

Mosquera, M., Ollé, A., Rodríguez, X.P., 2013. From Atapuerca to Europe: Tracing the earliest peopling of Europe. Quaternary International 295, 130-137.

Mosquera, M., Ollé, A., Saladié, P., Cáceres, I., Huguet, R., Rosas, A., Villalaín, J., Carrancho, A., Bourlès, D., Braucher, R., Pineda, A., Vallverdú, J., 2015a. The Early Acheulean technology of Barranc de la Boella (Catalonia, Spain). Quaternary International doi:http://dx.doi.org/10.1016/j.quaint.2015.05.005

Mosquera, M., Saladié, P., Ollé, A., Cáceres, I., Huguet, R., Villalaín, J.J., Carrancho, A., Bourlès, D., Braucher, R., Vallverdú, J., 2015b. Barranc de la Boella (Catalonia, Spain): an Acheulean elephant butchering site from the European late Early Pleistocene. Journal of Quaternary Sciences 30(7), 651-667.
Müller, U.C., Őrg Pross, J., Bibus, E., 2003. Vegetation response to rapid climate change in Central Europe during the past 140,000 yr based on evidence from the Füramoos pollen record. Quaternary Research 59, 235–245.

Muttoni, G., Kent, D. V, Scardia, G., Martin, R.A., 2015. Bottleneck at Jaramillo for human migration to Iberia and the rest of Europe? Journal of Human Evolution 80, 187–190.

Naafs, B.D.A., Hefter, J., Ferretti, P., Stein, R. and Haug, G.H., 2011. Sea surface temperatures did not control the first occurrence of Hudson Strait Heinrich Events during MIS 16. Paleoceanography, 26(4).

Nitychoruk, J., Ber, A., Hoefs, J., Krzywicki, T., Schneider, J., Winter, H., 2000. Interglaziale Klimaschwankungen in Nordost-Polen–palynologische und isotopengeochemische Untersuchungen an organischen Seesedimenten. Eiszeitalter Gegenwart 50, 86–94.

Okuda, M., Yasuda, Y., Setoguchi, T., 2001. Middle to Late Pleistocene vegetation history and climatic changes at Lake Kopais, Southeast Greece. Boreas 30, 73–82.

Ollé, A., Mosquera, M., Rodríguez, X.P., de Lombera-Hermida, A., García-Antón, M.D., García-Medrano, P., Peña, L., Menéndez, L., Navazo, M., Terradillos, M., Bargalló, A., 2013. The Early and Middle Pleistocene technological record from Sierra de Atapuerca (Burgos, Spain). Quaternary International 295, 138-167.

Ollé, A., Mosquera, M., Rodríguez-Álvarez, X.P., García-Medrano, P., Barsky, D., de Lombera-Hermida, A., Carbonell, E., 2016. The Acheulean from Atapuerca: Three steps forward, one step back. Quaternary International, in press.

Orain, R., Ermolli, E.R., Lebreton, V., Di Donato, V., Bahain, J.-J., Sémah, A.-M., 2015. Vegetation sensitivity to local environmental factors and global climate changes during the Middle Pleistocene in southern Italy—A case study from the Molise Apennines. Revue Palaeobotanique
Palynologie 220, 69–77.

Orain, R., Lebreton, V., Ermolli, E.R., Combourieu-Nebout, N., Sémah, A.-M., 2013a. Carya as marker for tree refuges in southern Italy (Boiano basin) at the Middle Pleistocene. Palaeogeography Palaeoclimatololgy Palaeoecology 369, 295–302.

Orain, R., Lebreton, V., Russo Ermolli, E., Sémah, A.-M., Nomade, S., Shao, Q., Bahain, J.-J., Thun Hohenstein, U., Peretto, C., 2013b. Hominin responses to environmental changes during the Middle Pleistocene in central and southern Italy. Climate Past 9, 687–697.

Ozenda, P., 1994. Végétation du continent européen. Delachaux et Niestlé Lausanne.

Paillard, D., 1998. The timing of Pleistocene glaciations from a simple multiple-state climate model. Nature 391, 378–381.

Paillard, D., Labeyrie, L., Yiou, P., 1996. Macintosh Program performs time-series analysis. EOS Trans. 77, 379.

Palombo, M.R., 2016. Discrete dispersal bioevents of large mammals in Southern Europe in the post-Olduvai Early Pleistocene: A critical overview. Quaternary International, in press.

Parrenin, F., Barnola, J.-M., Beer, J., Blunier, T., Castellano, E., Chappellaz, J., Dreyfus, G., Fischer, H., Fujita, S., Jouzel, J., Kawamura, K., Lemieux-Dudon, B., Loulergue, L., Masson-Delmotte, V., Narcisi, B., Petit, J.-R., Raisbeck, G., Raynaud, D., Ruth, U., Schwander, J., Severi, M., Spahni, R., Steffensen, J.P., Svensson, A., Udisti, R., Waelbroeck, C., Wolff, E., 2007. The EDC3 chronology for the EPICA Dome C ice core. Climate Past 3, 485–497.

Parés, J.M., Pérez-González, A., Rosas, A., Benito, A., de Castro, J.B., Carbonell, E. and Huguet, R., 2006. Matuyama-age lithic tools from the Sima del Elefante site, Atapuerca (northern Spain). Journal of Human Evolution, 50(2), 163-169.

Parés, J.M., Arnold, L., Duval, M., Demuro, M., Pérez-González, A., de Castro, J.B., Carbonell, E.,
Arsuaga, J.L., 2013. Reassessing the age of Atapuerca-TD6 (Spain): new paleomagnetic results.
*Journal of archaeological science* 40(12), 4586-4595.

Parfitt, S.A., Barendregt, R.W., Breda, M., Candy, I., Collins, M.J., Coope, G.R., Durbridge, P., Field, M.H., Lee, J.R., Lister, A.M. and Mutch, R., 2005. The earliest humans in Northern Europe: artefacts from the Cromer Forest Formation at Pakefield. *Nature* 438, 1008-101

Parfitt S. A., Ashton N.M., Lewis S.G., Richard A. L., Coope G. R., Field M.H., Gale R., Hoare P. G., Larkin N. R., Lewis M.D., Karloukovski V., Maher B.A., Peglar S.M., Preece R.C., Whittaker J.E., Stringer C. B., 2010. Early Pleistocene human occupation at the edge of the boreal zone in northwest Europe. *Nature* 466, 229-233.

Peinado Lorca, M., Martínez-Parras, J.M., 1987. Castilla-La Mancha, In: Peinado Lorca, M., and Rivas Martínez, S., (Eds.) *La vegetación de España. Alcala de Henares, Universidad de Alcalá de Henares*, 163-196.

Pereira, A., Nomade, S., Voinchet, P., Bahain, J.J., Falguères, C., Garon, H., Lefèvre, D., Raynal, J.P., Scao, V., Piperno, M., 2015. The earliest securely dated hominin fossil in Italy and evidence of Acheulian occupation during glacial MIS 16 at Notarchirico (Venosa, Basilicata, Italy). *Journal of Quaternary Sciences* 30(7), 639-651.

Peretto, C., Arnaud, J., Moggi-Cecchi, J., Manzi, G., Nomade, S., Pereira, A., Falguères, C., Bahain, J.-J., Grimaud-Hervé, D., Berto, C., 2015a. A Human Deciduous Tooth and New 40Ar/39Ar Dating Results from the Middle Pleistocene Archaeological Site of Isernia La Pineta, Southern Italy. *PLoS One* 10, e0140091.

Peretto, C., Arzarello, M., Bahain, J.-J., Boulbes, N., Dolo, J.-M., Douville, E., Falguères, C., Frank, N., Garcia, T., Lembo, G., Moigne, A.-M., Muttillo, B., Nomade, S., Pereira, A., Rufo, M.A., Sala, B., Shao, Q., Hohenstein, U.T., Tessari, U., Turrini, M.C., Vaccaro, C., 2015b. The Middle Pleistocene site of Guado San Nicola (Monteroduni, Central Italy) on the Lower/Middle Palaeolithic
transition. Quaternary International doi:http://dx.doi.org/10.1016/j.quaint.2015.11.056

Pidek, I.A., 2015. Palynostratigraphy and vegetation changes during the early Middle Pleistocene, based on new studies of deposits from Ferdynandów (central eastern Poland). Acta Palaeobotanica 55, 54–67.

Piperno, M., 1999. Notarchirico. Un sito del Pleistocene medio iniziale nel bacino di Venosa. Edizioni Osanna.

Pol, K., Masson-Delmotte, V., Cattani, O., Debret, M., Falourd, S., Jouzel, J., Landais, A., Minster, B., Mudelsee, M., Schulz, M., Stenni, B., 2014. Climate variability features of the last interglacial in the East Antarctic EPICA Dome C ice core. Geophysic Research Letter 41, 4004–4012.

Pons, A., Suc, J.P., Reille, M., Combourieu-Nebout, N., 1995. The history of dryness in regions with a Mediterranean climate. Time scales Biol. responses to water constraints. SPB Acad. Publ. Bv., Amsterdam 169–188.

Preece, R.C., Parfitt, S.A., 2012. The Early and early Middle Pleistocene context of human occupation and lowland glaciation in Britain and northern Europe. Quaternary International 271, 6–28.

Pross, J., Koutsodendris, A., Christianis, K., Fischer, T., Fletcher, W.J., Hardiman, M., Kalaitzidis, S., Knipping, M., Kotthoff, U., Milner, A.M., 2015. The 1.35-Ma-long terrestrial climate archive of Tenaghi Philippon, northeastern Greece: Evolution, exploration, and perspectives for future research. Newsletters Stratigraphy 48, 253–276.

Qingfeng, S., 2011. Combined ESR/U-series Dating of Fossil Teeth from Middle Pleistocene Sites in Northern Europe and Mediterranean Area: Contributing to the Chronology of the Acheulian Settlement of Europe. Département de Préhistoire. Muséum national d'Histoire Naturelle, Paris.

Quézel, P., Médail, F., 2003. Ecologie et biogéographie des forêts du bassin méditerranéen.

Roberts, M.B., Parfitt, S.A., 1999. Boxgrove: A Middle Pleistocene Hominid Site at Eartham Quarry,
Boxgrove, West Sussex. Archaeological Report 17.

Raymo, M.E., Lisiecki, L.E., Nisancioglu, K.H., 2006. Plio-Pleistocene Ice Volume, Antarctic Climate, and the Global δ18O Record. Science 80-313, 492–495.

Roebroeks, W., Villa, P., 2011. On the earliest evidence for habitual use of fire in Europe. PNAS 108(13), 5209-5214.

Reutenauer, C., Landais, A., Blunier, T., Bréant, C., Kageyama, M., Woillez, M.-N., Risi, C., Mariotti, V., Braconnot, P., 2015. Quantifying molecular oxygen isotope variations during a Heinrich stadial. Climate Past 11, 1527–1551.

Rhodes, R.H., Brook, E.J., Chiang, J.C.H., Blunier, T., Maselli, O.J., McConnell, J.R., Romanini, D., Severinghaus, J.P., 2015. Enhanced tropical methane production in response to iceberg discharge in the North Atlantic. Science 80-348, 1016–1019.

Rodríguez, J., Burjachs, F., Cuenca-Bescós, G., García, N., Van der Made, J., Pérez González, A., Blain, H.A., Expósito, I., López-García, J.M., García Antón, M., Allué, E., Cáceres, I., Huguet, R., Mosquera, M., Ollé, A., Rosell, J., Parés, J.M., Rodríguez, X.P., Díez, C., Rofes, J., Sala, R., Saladié, P., Vallverdú, J., Bennasar, M.L., Blasco, R., Bermúdez de Castro, J.M., Carbonell, E., 2011. One million years of cultural evolution in a stable environment at Atapuerca (Burgos, Spain). Quaternary Science Reviews 30, 1396–1412.

Rodríguez, J., Mateos, A., Hertler, C., Palombo, M.R., 2015. Modelling human presence and environmental dynamics during the Mid-Pleistocene Revolution: New approaches and tools. Quaternary International doi:http://dx.doi.org/10.1016/j.quaint.2015.06.003

Rodríguez-Gómez, G., Mateos, A., Martín-González, J.A., Blasco, R., Rosell, J. and Rodríguez, J., 2014. Discontinuity of human presence at Atapuerca during the early Middle Pleistocene: a matter of ecological competition? PloS one 9(7), e101938.
Rodríguez-Rey, M., Herrando-Pérez, S., Gillespie, R., Jacobs, Z., Saltré, F., Brook, B.W., Prideaux, G.J., Roberts, R.G., Cooper, A., Alroy, J., Miller, G.H., Bird, M.I., Johnson, C.N., Beeton, N., Turney, C.S.M., Bradshaw, C.J.A., 2015. Criteria for assessing the quality of Middle Pleistocene to Holocene vertebrate fossil ages. Quaternary Geochronology 30, Part A, 69–79.

Roebroeks W., Villa P., 2011. On the earliest evidence for habitual use of fire. PNAS 106(13), 5209–5214.

Rohling, E.J., Foster, G.L., Grant, K.M., Marino, G., Roberts, A.P., Tamisiea, M.E., Williams, F., 2014. Sea-level and deep-sea-temperature variability over the past 5.3 million years. Nature 508, 477–482.

Röthlisberger, R., Mudelsee, M., Bigler, M., de Angelis, M., Fischer, H., Hansson, M., Lambert, F., Masson-Delmotte, V., Sime, L., Udisti, R., Wolff, E.W., 2008. The Southern Hemisphere at glacial terminations: insights from the Dome C ice core. Climate Past 4, 345–356.

Sadori, L., Bertini, A., Combourieu-Nebout, N., Kouli, K., Mariotti Lippi, M., Roberts, N., Mercuri, A.M., 2013. Palynology and Mediterranean vegetation history. Flora Mediterranea 23, 141–156.

Sadori, L., Koutsodendris, A., Panagiotopoulos, K., Masi, A., Bertini, A., Combourieu-Nebout, N., Francke, A., Kouli, K., Joannin S., Mercuri, A.M., Peyron, O., Torri, P., Wagner, B., Zanchetta, G., Sinopoli, G., Donders, T.H., 2016. Pollen-based paleoenvironmental and paleoclimatic change at Lake Ohrid (south-eastern Europe) during the past 500 ka. Biogeosciences 13, 1423–1437.

Santagata, C., 2016. Operating systems in units B and E of the Notarchirico (Basilicata, Italy) ancient Acheulean open-air site and the role of raw materials. Quaternary International, in press.

Saladié, P., Rodríguez-Hidalgo, A., Huguet, R., Cáceres, I., Díez, C., Vallverdú, J., Canals, A., Soto, M., Santander, B., de Castro, J.M.B., Arsuaga, J.L., 2014. The role of carnivores and their relationship to hominin settlements in the TD6-2 level from Gran Dolina (Sierra de Atapuerca, Spain).
Quaternary Science Reviews 93, 47-66.

Sánchez-Goñi, M.F., Landais, A., Fletcher, W.J., Naughton, F., Desprat, S., Duprat, J., 2008. Contrasting impacts of Dansgaard-Oeschger events over a western European latitudinal transect modulated by orbital parameters. Quaternary Science Reviews 27, 1136-1151.

Sánchez-Goñi, M. F., Sánchez, Llave, E., Oliveira, D., Naughton, F., Desprat, S., Ducassou, E., Hodell, D. A., Hernández-Molina, F. J. 2016. Climate changes western Iberia and Mediterranean Outflow variations during two contrasting cycles of the last 1Myrs: MIS 31–MIS 30 and MIS 12–MIS 11. Global and Planetary Change 136, 18-29.

Sánchez Yustos, P., Diez Martín, F., 2015. Dancing to the rhythms of the Pleistocene? Early Middle Paleolithic population dynamics in NW Iberia (Duero Basin and Cantabrian Region). Quaternary Science Reviews 121, 75-88.

Satkunas, J., Grigiene, A., Velichkevich, F., Robertsson, A.N.N., Sandgren, P., 2003. Upper Pleistocene stratigraphy at the Medininkai site, eastern Lithuania: a continuous record of the Eemian-Weichselian sequence. Boreas 32, 627-641.

Scott, G.R., Gibert, L., 2009. The oldest hand-axes in Europe. Nature 461, 82-85.

Severinghaus, J.P., Beaudette, R., Headly, M.A., Taylor, K., Brook, E.J., 2009. Oxygen-18 of O2 Records the Impact of Abrupt Climate Change on the Terrestrial Biosphere. Science 324, 1431-1434.

Shakun, J.D., Lea, D.W., Lisiecki, L.E., Raymo, M.E., 2015. An 800-kyr record of global surface ocean δ18O and implications for ice volume-temperature coupling. Earth Planetary Sciences Letters 426, 58-68.

Smith, G.M., 2013. Taphonomic resolution and hominin subsistence behaviour in the Lower Palaeolithic: differing data scales and interpretive frameworks at Boxgrove and Swanscombe (UK). J. Archaeol. Sci. 40, 3754-3767. Stewart, J.R., Lister, A.M., Barnes, I., Dalén, L., 2010.
Refugia revisited: individualistic responses of species in space and time. Proceedings Royal Society London B Biological Sciences 277, 661–671.

Spratt, R.M., Lisiecki, L.E., 2015. A Late Pleistocene sea level stack. Climate Past Discussion 11, 3699–3728.

Stahlschmidt, M.C., Miller, C.E., Ligouis, B., Hambach, U., Goldberg, P., Berna, F., Richter, D., Urban, B., Serangeli, J., Conard, N.J., 2015. On the evidence for human use and control of fire at Schöningen. Journal of human evolution 89, 181-201.

Stewart, J.R., Lister, A.M., Barnes, I., Dalén, L., 2010. Refugia revisited: individualistic responses of species in space and time. Proceedings of the Royal Society B 277, 661-671.

Stiner M. C., 2002. Carnivory, Coevolution, and the Geographic Spread of the Genus Homo. Journal of Anthropological Research 10, 1–63.

Stringer, C., 1996. The Boxgrove Tibia: Britain's Oldest Hominid and its place in the Middle Pleistocene Record, in: Gamble, C., Lawson, A.J. (Eds.), The English Heritage Reviewed. Wessex Archaeology, pp. 53–56.

Stringer C., 2011. The Changing Landscapes of the Earliest Human Occupation of Britain and Europe, in: Ashton N., Lewis S.G., Stringer C. (Eds.), The Ancient Human Occupation of Britain. Quaternary Science 14, pp. 1–11.

Stringer C., 2012. The Status of Homo heidelbergensis (Schoetensack 1908). Evolutionary Anthropology 21, 101–107.

Suwa, M., Bender, M. L., 2008. Chronology of the Vostok ice core constrained by O$_{2}$/$N_{2}$ ratios of occluded air, and its implication for the Vostok climate records. Quaternary Science Reviews 27, 1093–1106.

Thun Hohenstein, U., Di Nucci, A., Moigne, A.-M., 2009. Mode de vie à Isernia La Pineta (Molies,
Italie). Stratégies d'exploitation de Bison schoetensacki par les groupes humains au
Paléolithique inférieur. L'Anthropologie 113(1), 96-110.

Turner, A., 1992. Large carnivores and earliest European hominids: changing determinants of resource availability during the Lower and Middle Pleistocene. Journal of Human Evolution 22, 109–126.

Tzedakis, P.C., 1999. The last climatic cycle at Kopais, central Greece. Journal of Geological Society London. 156, 425–434.

Tzedakis, P.C., 2007. Seven ambiguities in the Mediterranean palaeoenvironmental narrative. Quaternary Science Reviews 26, 2042–2066.

Tzedakis, P.C., Frogley, M.R., Heaton, T.H.E., 2002. Duration of last interglacial conditions in northwestern Greece. Quaternary Research 58, 53–55.

Tzedakis, P.C., Hooghiemstra, H., Pälike, H., 2006. The last 1.35 million years at Tenaghi Philippon: revised chronostratigraphy and long-term vegetation trends. Quaternary Science Reviews 25, 3416–3430.

Tzedakis, P. C., Raynaud, D. R, McManus, J. F., Berger, A., Brovkin, V., and Kiefer, T., 2009. Interglacial diversity, Nature Geoscience 2, 751–755.

Vaesen, K., Collard, M., Cosgrove, R., Roebroeks, W., 2016. Population size does not explain past changes in cultural complexity. PNAS 113(16), e2241-E2247.

Vallverdú, J., Saladié, P., Rosas, A., Huguet, R., Cáceres, I., Mosquera, M., Garcia-Tabernero, A., Estalrrich, A., Lozano-Fernández, I., Pineda-Alcalá, A., 2014. Age and date for early arrival of the Acheulian in Europe (Barranc de la Boella, la Canonja, Spain). PloS One 9(7), e103634.

van der Made, J., Rosell, J., Blasco, R., 2015. Faunas from Atapuerca at the Early–Middle Pleistocene limit: The ungulates from level TD8 in the context of climatic change. Quaternary International
Veres, D., Bazin, L., Landais, a., Toyé Mahamadou Kele, H., Lemieux-Dudon, B., Parrenin, F., Martinerie, P., Blayo, E., Blunier, T., Capron, E., Chappellaz, J., Rasmussen, S.O., Severi, M., Svensson, a., Vinther, B., Wolff, E.W., 2013. The Antarctic ice core chronology (AICC2012): An optimized multi-parameter and multi-site dating approach for the last 120 thousand years. Climate Past 9, 1733–1748.

Vincenzo, F. Di, Rodriguez, L., Carretero, J.M., Collina, C., Geraads, D., Piperno, M., Manzi, G., 2015. The massive fossil humerus from the Oldowan horizon of Gombore I, Melka Kunture (Ethiopia, 1.39 Ma). Quaternary Science Reviews 122, 207–221.

Voinchet, P., Moreno, D., Bahain, J.-J., Tissoux, H., Tombret, O., Falguères, C., Moncel, M.-H., Schreve, D., Candy, I., Antoine, P., Ashton, N., Beamish, M., Cliquet, D., Despriée, J., Lewis, S., Limondin-Lozouet, N., Locht, J.-L., Parfitt, S., Pope, M., 2015. New chronological data (ESR and ESR/U-series) for the earliest Acheulian sites of north-western Europe. Journal of Quaternary Sciences. 30, 610–623.

Wagner, G.A., Maul, L.C., Löscher, M., Schreiber, H.D., 2011. Mauer - the type site of Homo heidelbergensis: palaeoenvironment and age. Quaternary Science Reviews 30, 1464–1473.

Wang, Y., Cheng, H., Edwards, R.L., Kong, X., Shao, X., Chen, S., Wu, J., Jiang, X., Wang, X., An, Z., 2008. Millennial- and orbital-scale changes in the East Asian monsoon over the past 224,000[thinsp]years. Nature 451, 1090–1093.

Wijmstra, T.A., 1969. Palynology of the first 30 metres of a 120 m deep section in northern Greece. Acta Botanica Neerland 18, 511–527.

Wolff, E.W., Berger, A., Crucifix, M., Hodell, D.A., Mangili, C., McManus, J.F., Otto-Bliesner, B., Pol, K., Raynaud, D., Skinner, L.C., Tzedakis, P.C., Yin, Q.Z., Abe-Ouchi, A., Barbante, C., Brovkin, V.,
Cacho, I., Capron, E., Ferretti, P., Ganopolski, A., Grimalt, J.O., Hönisch, B., Kawamura, K., Landais, A., Margari, V., Martrat, B., Masson-Delmotte, V., Mokeddem, Z., Parrenin, F., Prokopenko, A.A., Rashid, H., Schulz, M., Vazquez Riveiros, N., 2015. "Interglacials of the last 800,000 years." *Revue de Geophysique* n/a-n/a. doi:10.1002/2015RG000482

Woillard, G., 1978. Grande Pile peat bog: A continuous pollen record for the Last 140,000 years. *Quaternary Research* 9, 1-21.

Yin, Q.Z., Berger, A., 2012. Individual contribution of insolation and CO2 to the interglacial climates of the past 800,000 years. *Climate Dynamics* 38, 709-734.

Zagwijn, W.H., 1992. The beginning of the ice age in Europe and its major subdivisions. *Quaternary Science Reviews* 11, 583-591.

Zagwijn, W.H., 1996. The Cromerian Complex Stage of the Netherlands and correlation with other areas in Europe. *Early Middle Pleistocene Europe, Ed. by Turner, C.*, Balkema, Rotterdam 145-172.
## Figure captions

<caption>Figure 1: Map of the Western Europe archaeological sites selected for this study (white rounds).</caption>

<caption>Figure 2: Age Estimates (including margins of error for the dates) of Archaeological sites with indication of lithic tradition (Cagny La Garenne 1, 2; Madcross Hill; Pakefield; Happisburgh 3; High Lodge; la Noira; Happisburgh 1; Boxgrove; Gran Dolina TD6; La Pineta; Sima del Elefante; La Boella; La Caune de l'Arago; Notarchirico. a) black line: benthic stack (Lisiecki and Raymo,2005); blue line: *Cibicides wuellerstorfi* curve from ODP U1308 (Hodell et al., 2008). b) Global sea level reconstructions from Rohling et al., 2014 (green) and Elderfield et al., 2012 (light blue). c) Mediterranean planktonic curve from ODP 975 (Lourens et al., 2004); d) See Surface Temperature (SST) based on Alkenone (Central Atlantic ODP site U1313) (Naafs et al., 2011).</caption>

<caption>Figure 3. Ice core records from EPICA Dome C covering the last 800 ka from Antarctica. a) CH$_4$ concentration (Louergue et al., 2008), b) $\delta^{18}$O$_{atm}$ composition (composite of EDC and Vostok data, Bazin et al., 2013; Dreyfus et al., 2008, 2007; Suwa and Bender, 2008), c) dust flux (Lambert et al., 2008), d) $\delta$D(Jouzel et al., 2007), e) orbital parameters (obliquity in green, precession in blue) obtained with Analyseries (Paillard et al., 1996), f) records of Ca/Sr of marine cores U1302/03 and Si/Sr of marine core U1308 located respectively on the western and eastern border of the North Atlantic (IRD proxies, Channell et al., 2012; Hodell et al., 2008).</caption>
Figure 4. Schematic representation of vegetation dynamics, temperature and precipitation changes corresponding to glacial and interglacial periods as inferred from pollen records. A glacial period is roughly characterized by open vegetation, cold temperature and low precipitation. An interglacial period is composed of a changing vegetation (from a coniferous to a pionner forest), warm temperature and high precipitation.

Figure 5. Correlation of the vegetation dynamic in Europe with δ¹⁸O marine records and the chronology of the [900-500] ka archaeological sites discussed in this study. a) and e) Vegetation dynamic in respectively the temperate and Mediterranean area (ST, subtropical trees; TF, Temperate forest; MTS, Mediterranean trees and shrubs; PT, pioneer trees; C, Conifers; DM, dry meadow; St, steppe), b) δ¹⁸O from benthic foraminifera in the North Atlantic (in blue, after Channell et al., 2008) and global stack (in black, after Lisiecki and Raymo, 2005), c) Age Estimates of Archaeological sites with indication of lithic tradition, d) δ¹⁸O from planktonic foraminifera in the Mediterranean sea (green, after Lourens, 2004).
<figure><img src="image_2.png" /></figure>

NASA Earth Observatory
<figure><img src="image_3.png" /></figure>

Listeckii and Raymo, 2005, Hoddell et al., 2008

Elderfield et al., 2012

Robling et al., 2014

Lourens, 2004

Naafs et al., 2011
<figure><img src="image_4.png" /></figure>
Climate cycle

<figure><img src="image_5.png" /></figure>

I, intergacial; G, glacial

Vegetation

<figure><img src="image_6.png" /></figure>

P, pionner forest; MO, mixed oak forest, C, coniferous forest; OV, open vegetation

Temperature

<figure><img src="image_7.png" /></figure>

Opt, climate optimum; Min, climate minimum (from vegetation)

Precipitation

<figure><img src="image_8.png" /></figure>

Opt, climate optimum; Min, climate minimum (from vegetation)
<figure><img src="image_9.png" /></figure>

<figure><img src="image_10.png" /></figure>

<figure><img src="image_11.png" /></figure>

<figure><img src="image_12.png" /></figure>

<figure><img src="image_13.png" /></figure>
<table><caption>Table 1. 900-500 ka European sites selected for the paper</caption><thead><tr><td>Country</td><td>Sites</td><td>Lithic composition</td><td>Radiometric Ages</td><td>MIS</td><td>Dating Methods</td><td>Some references</td></tr></thead><tbody><tr><td rowspan="2">Great-Britain</td><td>Happisburg 3</td><td>Flakes and cores series</td><td>970-936 ka<br>870-814 ka</td><td>MIS 25<br>MIS 21</td><td>Biostratigraphy<br>(Paleomagnetism)</td><td>Parfitt et al., 2010</td></tr><tr><td>Pakefield</td><td>Flakes and cores series</td><td>700 ka</td><td></td><td>Amino-acid<br>Biostratigraphy<br>(Paleomagnetism)</td><td>Parfitt et al., 2005</td></tr><tr><td rowspan="2">Great-Britain</td><td>Maidcross Hill</td><td>Bifacial technology</td><td>529 +- 55 ka<br>631 +-6 ka</td><td>MIS 15-14</td><td>ESR/U-Th quartz<br>Stratigraphy</td><td>Preece and Parfitt, 2012<br>Voinchet et al., 2015</td></tr><tr><td>Happisburg 1</td><td>Flakes and cores series with 1 biface</td><td>-</td><td>MIS 13</td><td>Amino-acid<br>Biostratigraphy</td><td>Parfitt et al., 2010</td></tr><tr><td rowspan="2">Great-Britain</td><td>Boxgrove</td><td>Bifacial technology</td><td>-</td><td>MIS 13</td><td>Biostratigraphy<br>Stratigraphy</td><td>Robert and Parfitt, 1999</td></tr><tr><td>High Lodge</td><td>Flakes and cores series</td><td>-</td><td>MIS 13</td><td>Biostratigraphy<br>Stratigraphy</td><td>Ashton et al., 1992<br>Ashton and Lewis, 2012<br>Preece and Parfitt, 2012</td></tr><tr><td rowspan="2">France (North)</td><td>Cagny-la-Garenne I-II</td><td>Bifacial technology</td><td>400 ± 101 ka</td><td>MIS 13/12</td><td>ESR teeth<br>Biostratigraphy<br>Stratigraphy</td><td>Antoine et al., 2007<br>Lamotte and Tuffreau, 2015</td></tr><tr><td>Brinay, la Noira</td><td>Bifacial technology</td><td>655 ± 55 ka</td><td>MIS 16</td><td>ESR quartz<br>Stratigraphy</td><td>Voinchet et al., 2010<br>Despriee et al., 2011<br>Moncel et al., 2013</td></tr><tr><td>France (Centre)</td><td>Tautavel-Arago P-Q</td><td>Bifacial technology</td><td>532 ± 106</td><td>MIS 14</td><td>Biostratigraphy<br>ESR/U-Th teeth</td><td>Barsky and Lumley, 2010<br>Falguères et al., 2015</td></tr><tr><td>Spain</td><td>La Boella</td><td>Flakes and cores series and Bifacial technology</td><td>990 to 780 ka</td><td>MIS 25</td><td>Paleomagnetism<br>Biostratigraphy<br>Cosmogenic</td><td>Vallverdu et al., 2014<br>Mosquera et al., 2015</td></tr></tbody></table>
<table><tr><td>Spain</td><td>Atapuerca Sima del Elefante</td><td>Flakes and cores series</td><td>TE 16 = 804 ± 47 ka and 864 ± 88 ka TE17 = 724 ± 43 ka and 781 ± 63</td><td>MIS 21</td><td>ESR/U-Th teeth - quartz Paleolagnetism Biostratigraphy</td><td>De Lombrera-Hermida et al., 2015</td></tr><tr><td>Spain</td><td>Atapuerca, Gran Dolina-TD6</td><td>Flakes and cores series</td><td>857 - 780 ka 731 ± 63 ka</td><td>MIS 25 MIS 21 MIS 19</td><td>Paleomagnetism ESR/U-Th Biostratigraphy TL anf IRSL</td><td>Falguères et al., 1999 Cuenca Bescos et al., 2010 Berger et al., 2008</td></tr><tr><td>Italy</td><td>Nortarchirico-di-Venosa</td><td>Flakes and cores series and Bifacial technology</td><td>39Ar/40Ar = 670 ± 4 ka to 614 ± 4 ka ESR = 637 ± 31 ka</td><td>MIS 16</td><td>ESR Quartz TL 40Ar/39Ar</td><td>Piperno ed., 1999 Lefèvre et al., 2010 Pereira et al., 2015</td></tr><tr><td>Italy</td><td>Isernia-la-Pineta</td><td>Flakes and cores series</td><td>583-561 ka</td><td>MIS 15/ MIS 14</td><td>39Ar/40Ar</td><td>Peretto et al., 2015</td></tr></table>
<table><caption>Table 2. Summary of the environmental and climatic context for the hominin occupations in Western Europe between 900 and 500 ka beyond and under the 45th parallel.</caption><thead><tr><th></th><th colspan="2">Series without bifaces</th><th colspan="2">Series with bifaces</th></tr><tr><th></th><th>1 Ma-700 ka</th><th>700-500/450 ka</th><th>1 Ma-700 ka</th><th>700-500/450 ka</th></tr></thead><tbody><tr><td>Beyond the 45th parallel</td><td>Happisburg 3 = conifer-dominated woodland</td><td>High-Lodge = coniferous forest (cool-temperate climate ?)</td><td></td><td>Bytham River sites = open grassland</td></tr><tr><td>South Great-Britain, North and Centrer France</td><td>Pakefield = broad-leaf woodland (climatic optimum)</td><td></td><td></td><td>Boxgrove = scrub and woodland<br/>Cagny-la-Garenne I-II = pleniglacial environment<br/>La Noira = beginning of a cold phase, pleniglacial</td></tr><tr><td>Under the 45th parallel</td><td>Sima del Elefante TE7-TE16 = Mediterranean optimum climatic</td><td>Isernia = open vegetation of woodland evolving to steppic conditions</td><td>La Boella = ?</td><td>Notarchirico = open steppic vegetation</td></tr><tr><td>Italy, Spain, South France</td><td>Gran Dolina TD6 = alternating Midterranean optimum climatic and open steppe vegetation (mosaic environments)</td><td></td><td></td><td>Caune de l'Arago P-Q = open steppic vegetation</td></tr></tbody></table>