Atapuerca - Patrimonio de la humanidad

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The Physical and Geological Setting

La Sierra de Atapuerca se encuentra en el Corredor de la Bureba, un corredor estratégico entre las cuencas del Duero y el Ebro The Sierra de Atapuerca is located in the north of Spain, on the northern Meseta, and has a maximum altitude of 1,085 meters above sea level. Located some 15 km to the East of the city of Burgos, the Sierra is surrounded by the Pico, Vena and Arlanzón rivers and forms part of the last mountains which comprise the Iberian System. It is separated from the southern border of the Cantabrian Cordillera by a tectonic corridor (known as the 'Strait of Burgos') which links the tertiary depression of the Duero River Basin with that of the Ebro River, two of the largest rivers in Spain, and which forms a natural pass providing access to the interior of the Iberian Peninsula. At the same time, the Sierra is located at a biogeographic crossroads where Mediterranean, Atlantic and Continental climatic regimes converge. This climatic confluence has fostered the coexistence of a wide variety of species of fauna and flora and an ecosystem (with a great diversity of biotopes) which has been exploited by different groups of humans throughout time.

The Sierra de Atapuerca corresponds tectonically to a fallen anticline (sedimentary layers forming an inverted V-shape), with a NE inclination and an orientation similar to the nearby Sierra de la Demanda, i.e. NNW-SSE, which was formed in limestone of marine origin laid down during the Upper Cretaceous period (between 80-100 million years ago). On the borders of the Sierra, materials from the Tertiary period (from 25-5 million years ago) overlay the inclined layers of the anticline. These ar materials are of continental origin, formed by Oligocene lime conglomerates and red clays, and e in turn overlain by a more or less horizontal lithological sequence of marls, clays, gypsums, and marly and limey pockets, typical of the lacustrine environment of the Duero River Basin.

Toward the end of the Pliocene and the beginning of the Pleistocene, the Arlanzón River began to carve out a fluvial valley, showing a stairstep-like profile with 15 levels of terraces. The valley is strongly asymmetric as it passes by the Sierra, whose peak reaches barely 50 meters above the uppermost terrace.

Present Ecosystems

The altitude and geographic location of the Sierra de Atapuerca produce a continental climate which is attenuated by the amount of rainfall and Mediterranean influence. The combination of these climatic factors with the particular substrate characteristics of the region (with a 'c' calcareous horizon and very thin soil in the Sierra) gives rise to an optimal natural vegetation coverage which is defined as 'subsclerofilous forest'. The Sierra is covered by low, patchy woods that rarely reach above four meters in height and are comprised principally of two varieties of oak trees. Throughout the Sierra, 'Quercus ilex' is predominant, with groups of 'Quercus faginea' proliferating in zones with greater depths of soil. These woody areas primarily extend along the southwest slope, leaving some clearings visible where the presence of the limestone bedrock has reduced the soil cover to only superficial levels.

El páramo castellano (derecha) y la Sierra de ATapuerca (izquierda) se encuentran aproximadamente a la misma altura The species of shrubs that grow at the boundaries between the woody areas and clearings include hawthorn ('Ramnus sexatillis' and 'Crataegus monogyna'), honeysuckles ('Lonicera etrusca' and' Lonicera splendida'), wild rose ('Rosa canina') and less frequently blackthorn ('Prunus spinosa') and wild jasmine ('Jasminum fruticans'). Herbaceous plants also appear, including, among others, 'Thimus cinereum', 'Heliantemum hirtum', 'Teucrium polium', 'Lavandula latifolia', 'Knautia arvensis', and 'Onopordon acanthium'. In the highest parts of the Sierra, the oaks are concentrated in patches, allowing a very thin vegetation cover to take over, composed of thyme ('Thimus szigis'), lavender ('Lavandula pedunculata') and gorse.

The Sierra is surrounded by grain fields that remain fallow during the winter and whose principal crops are wheat and barley. The soil is acidic and deep, having been laid down by the Arlanzón River. On the fields that remain fallow year round, opportunistic species of herbaceous plants such as Bromus and a variety of mustard plants proliferate during springtime.

La acción del Rio Arlazón ha dado forma a las cuevas y a los depósitos que en ellas se encuentran desde hace más de un millón de años Another important ecosystem in the Sierra de Atapuerca has developed along the banks of the Arlanzón River and consists of a mixed-gallery forest comprised of poplar and willow groves, including black poplar trees ('Populus nigra'), aspens ('Populus tremula') and various species of willow ('Salix neotricha', 'Salix alba' y 'Salix fragilis'). The river also forms various islands, whose size and shape changes depending on the annual water levels, and which are home to various types of rushes ('Thipha latifolia', 'Thipha dominguensis' and 'Tipha angustifolia') and reeds ('Phragmites australis', 'Scirpus lacustris', 'Scirpus maritimus', 'Juncus implexus', 'Juncus conglomeratus'). On the banks, the black poplars and willows are interspersed with pastures, fern groves and beds of reeds, where wild rose, mint ('Mentha longifolia') and clover ('Trifolium repens') grow. During the summer months, the clearings are utilized for grazing livestock, and the inhabitants of the nearby pueblo of Ibeas de Juarros take advantage of the presence of the Arlanzón River for recreational activities. Patches of elm trees are also present at various points along the river bank, although a severe attack of Dutch elm disease has reduced their numbers significantly.

A stroll from the Arlanzón River to the Sierra de Atapuerca

The Arlanzón is inhabited by numerous species typical of a river ecosystem. The trout ('Salmo trutta') swimming in its waters consumes a variety of aquatic invertebrates as well as flying insects which perch near the river’s surface. The water snake ('Natrix natrix') is another of the predators in this ecosystem.

Just beyond the riverbank, we might notice the jumps of some common frogs ('Rana perezi') as we pass by and we can see warblers ('Hippolais poliglota'), chiffchaffs (Phylloscopus collybita) and groups of sparrows ('Passer domesticus'). Some of the river birds are difficult to see, but their songs are easily heard. This is the case of the nightingale ('Cettia cetti'), which hides in the dense willows and thorny bushes, or the blue nuthatch, whose cries are heard from the branch of a black poplar along the banks of the Arlanzón. This is also the habitat of the wagtail ('Motacilla alba'), the moorhen ('Gallinula chloropus'), more easily seen in areas where the river current is calmer, and of the small white wagtail ('Actitis hypoleucos'), which runs along the riverbanks hiding among the reeds.

Not far from the Arlanzón river is the pueblo of Ibeas de Juarros. Upon passing through its streets to begin our climb to the Sierra de Atapuerca, we encounter flocks of sparrows throughout the year, and beginning in March the sky is filled with swallows ('Hirundo rustica'), swifts ('Apus apus') and common martins ('Delinchon urbica') which nest in the roofs of the houses.

After leaving the streets of Ibeas and continuing our journey to the Sierra, we enter a small open forest of oak trees ('Quercus pirenaica'). During the winter months, this small oak grove is filled with thrushes ('Turdus phylomenos'), groups of goldfinches ('Fringilla coelebs') and linnets ('Carduelis cannabina'), robins ('Erithacus rubecula') and common blackbirds ('Turdus merula'). The branches of the trees which form the border of the oak grove are favored by owls ('Tyto alba') that lie in wait for small rodents during the night and by buzzards ('Buteo buteo') during the day.

From here, we enter the fields which surround the Sierra. During the winter we find might see flocks of red partridges ('Alectoris rufa'), lapwings ('Vanellus vanellus'), choughs ('Pyrrhocorax pyrrhocorax'), rooks ('Corvus monedula'), crows ('Corvus corone') and magpies ('Pica pica'), all of which feed on small vertebrates and seeds which remain in the wheat fields. Pale hawks ('Circus cyaneus') make low-flying sweeps above the fields in search of small mammals and unwary chicks. Later, in springtime, they’ll travel to the north of Europe to breed, but in their place we see a sister species, the gray hawk ('Cyrcus pygargus'), which arrives from Africa to nest in the protection which the wheat fields provide.

Springtime brings the arrival of several other bird species, which, together with the gray hawk, cross the Straits of Gibraltar. We might see the northern stonechat ('Saxicola rubetra'), the common shrike ('Lanius senator'), or hear the song of the quail ('Coturnix coturnix'). These species, together with corn buntings ('Miliaria calandra'), green finches ('Carduelis chloris'), calandra larks ('Melanocoripha calandra'), common larks ('Alauda arvensis') and crested larks ('Galerida cristata') are the most common inhabitants of cultivated areas.

At sundown, we surprise a group of roe deer ('Capreolus capreolus') emerging from the thick vegetation that covers the Sierra to feed in the wheat fields, or an owl perched on the rock piles accumulated by farmers upon tilling their fields.

In spring, summer and part of fall, we can see the booted eagle ('Hieratus pennatus') and the short-toed eagle ('Circaetus galicus'), but other predators inhabit the Sierra year round, such as the fox ('Vulpus vulpes') or the common falcon ('Falco tinnunculus').

Tunel en el trazado del ferrocarril minero que recorría la Sierra de Atapuerca The road leads us to the route of the old mining railroad. Following it northwest we approach the slopes of the Sierra until we reach a clearing where we enter the limestone bedrock of the mountain. We can hear the beautiful song of the nightingale ('Luscinia megarynchos'), one of the most difficult birds to see. To the right of us is the Cueva del Silo where an eagle owl has occasionally been sighted, most likely a habitual occupant of one of the limestone walls that remain from past quarrying activities in the Sierra. The road now leads us into the interior of the railway trench. Falcons raise their young within the confines of the trench, but they aren’t alone, rooks and choughs make their nests in the numerous cracks and cavities in the limestone walls.

Inside the trench we walk along a path lined with honeysuckles, hawthorns, green hellebore ('Helleborus viridis'), asparagus plants and wild roses. Among them we spot the robin and the redstart ('Phoenicurus ochruros'). Both species inhabit the Sierra year round, and move between the bushes which grown along the roads and in the clearings.

En el interior de la Trinchera del Ferrocarril de la Sierra de Atapuerca (Burgos) A road leads out from the railway trench, through the woods, and ascends to the highest part of the Sierra. Among the dense oak trees, we can also find the coal tit ('Parus major'), the common tit and the common blackbird. On occasion, we might be surprised by wood pigeons ('Columba palumbus') when they emerge from the bushes or come across the common jay (Garrulus glandarius). A few species, such as the wild boar ('Sus scrofa'), the fox or the marten ('Martes foina') take refuge in the undergrowth that covers the Sierra. Their nocturnal habits make them even more difficult to spot, although their footprints and other traces of their presence are easily seen. Some of the clearings in the woods, formed by stony ground which derives from the old limestone quarries, are home to various lizard species, such as the jewelled lizard ('Lacerta lepida') or the Iberian lizard ('Podarcis hispanica'). We can also find the asp ('Vipera aspis'), which tends not to stray too far from these stony areas with low shrubs. Other snakes found in the Sierra include the viper ('Natrix maura') and the southern gray snake ('Coronela girondica').

The road leads us between oaks until we reach the summit of the Sierra, where the vegetation changes. The oak forest becomes patchier, joined by ever-larger clearings with thyme, lavender and gorse.

IMG_IZQ]135[/IMG_IZQ] Some species also arrive from the regions surrounding the Sierra. Groups of Griffon vultures ('Gips fulvus') come from the limestone bluffs of the Arlanza River to the south, or the ravines of the Pancorbo to the north. We might also see the fleeting presence of peregrine falcons ('Falco peregrinus') from these same regions.

THE PLEISTOCENE ECOSYSTEMS OF THE SIERRA DE ATAPUERCA

The Pleistocene: an Unstable World

The Pleistocene is the period of the Earth’s history from 1,800,000 to 10,000 years ago. It is mainly characterized by a series of cyclical climatic changes which affected the entire planet, although the effects were much more severe in the more northern latitudes.

These cycles correspond to the alternating of two climatic stages: glacial periods and interglacial periods. During the interglacial periods, the climate was very similar to the present day. However, during the glacial periods, the average temperature of the planet was lowered several degrees and the polar ice caps expanded significantly, particularly in the northern hemisphere, producing important drops in the sea level and affecting the rainfall distribution. In North America, for example, ice sheets covered the entire north of the continent reaching just south of the Great Lakes, and the desert Southwest was significantly more humid than present day.

El cuaternario: sus periodos temporales y culturales, las variaciones de temperatura y las glaciaciones In Europe, the glacial cycles left a profound mark on the continent. During a glacial cycle, Great Britain, Scandinavia, the Netherlands and northern Germany were covered year-round by ice sheets. In these arctic desert zones, life was almost impossible. To the south, in Central Europe, the ice disappeared during the spring and summer months, but the cold was so intense and the climate so dry that trees could not survive, leaving the landscape covered by immense steppes. Still further south, on the Iberian, Italian and Balkan peninsulas, the climate was milder, although different from today. Here there were areas where forests could still survive and took refuge from the cold, and the southern mountain slopes, canyons and ravines protected them from the strong, icy winds. In more open areas, the most resistant trees, like species of pine and cypress, could form open woods mixed with steppe-like vegetation.

Vegetation

The climatic oscillations which occurred during the Pleistocene caused changes in the natural environment, giving rise to distinct ecosystems. The reconstruction of these ecosystems through the fossil record can give us a good idea of the different landscapes which developed over the course of time in the Sierra.

The record of fossilized pollen is used to study the vegetational history of the Sierra de Atapuerca. To date, pollen analysis has been carried out at the sites of Gran Dolina and Galería, in the Trinchera. Between them an ample time period is covered, ranging from the end of the Lower Pleistocene, approximately 900,000 years ago, to the end of the Middle Pleistocene, some 200,000 years ago. It should be clarified that in the Gran Dolina, the archaeological levels are numbered from the base upwards. This numbering system is the opposite of that used at most archaeological sites. However, since the site has been cut by the railway trench, the entire stratigraphy is visible, and it isn’t necessary to excavate the site to know how many levels there are.

In the lower part of the sequence, in level TD4, some 900,000 years ago, there is a high percentage of tree pollen with a predominance of oak species (the precise species cannot currently be identified), together with wild olive trees. This brings to mind a wooded environment with Mediterranean vegetation, although with distinct characteristics, since other species typical of more humid climates are also present, such as beech, walnut and birch trees. Pollen from pine trees is also present. However, this data is more difficult to interpret, since these trees spread their pollen through the wind. Thus, even if the trees are located several kilometers away, there is always a high concentration of their pollen in the air.

In the next level, TD5, the percentage of tree pollen is somewhat reduced. Further, the species typical of humid climates disappear, increasing the importance of the oak species, and indicating that the climate was drier than in the previous level.

Level TD6 dates to around 800,000 years ago and corresponds to a transition from a cold glacial period to a milder interglacial period. Thus, the lower part of this level shows a low percentage of oak pollen, while cypress species abound, such as savines and junipers, which are typical of cold dry climates. In the upper part of TD6, in the Aurora stratum where the remains of 'Homo antecessor' were found, cypress species are barely present, and species typical of a Mediterranean climate, such as wild olive trees or the lentisk ('Pistacia'), appear.

In level TD7, the percentage of tree pollen is moderately high and, again, oak species predominate. These appear together with Mediterranean species, such as wild olive trees and vines, as well as more humid climate species such as beech trees.

In TD8, some 600,000 years ago, a different landscape is encountered, marked by the importance of warm and humid climate species, such as chestnut, beech and birch trees.

Finally, in TD10, about 350,000 years ago, there is a high proportion of tree pollen dominated by pine species, which leads us to believe that the climate was colder and drier than in the lower levels.

To continue our journey through time we must change sites to consider the stratigraphic sequence at the Galería site. The lower units, GI-II don’t contain sufficient pollen to draw any conclusions. In unit GIII, which dates to around 340,000 years ago, the landscape is once again dominated by oak species together with other trees such as beech, hazel, chestnut, ash and willows. The final period for which we have pollen information is Galería unit IV, some 150,000-200,000 years ago where the warm and humid forest species disappear, giving way to Mediterranean species, such as the wild olive, the lentisk or honeysuckle.

From what has been laid out above, we can conclude that the landscape surrounding Atapuerca suffered climatic changes throughout the Lower and Middle Pleistocene correlated with the glacial-interglacial cycles. The nature of these changes is well represented in TD6, where we can see that the changes from a glacial period to an interglacial period, although marked, were not as drastic as in more northerly latitudes. In fact, at no point does the vegetation disappear, and trees are always present. Rather, we see a replacement of the Mediterranean species by those which are more resistant to cold and aridity.

In general, the landscape has been dominated throughout this epoch by oak species and has varied primarily in the proportion and variety of species that accompany them. In some cases, these were warm and humid species, while at other times Mediterranean types, less tolerant of cold but better adapted to aridity, were more abundant. During moments of more extreme climatic conditions, oak species were relegated to a few refuge zones and the landscape was dominated by wooded steppes with cypress species. We can infer from this that during this time period the successive vegetational regimes in the Sierra de Atapuerca, both during warmer and colder times, were probably quite similar to those which are currently distributed throughout the Iberian Peninsula.

Pollen

Study of the pollen found in soil can help to reconstruct past landscapes. In the case of Atapuerca, pollen has been found in sediments inside the cave sites, having been swept in by the wind or brought in by animals in their hair, or even in their stomachs. Its hard outer covering makes pollen very resistant, and it often survives to be deposited in the sediments. Part of this pollen derives from egagropilas, balls of hair, feathers or bone which birds of prey form in their stomachs from the indigestible parts of their victims, and which they later expel through their mouths. These birds tend to perch in the same sites, and if this happens to be a tree branch near the mouth of a cave, a significant number of egagropilas can accumulate inside the cave in a short time.

Unfortunately, caves are not the best sites to preserve pollen because the oxygen-rich environment accelerates their degradation. The best stratigraphic sequences with pollen are found in lakebed sediments or peat bogs where there is no oxygen, but there aren’t any sites like this in Atapuerca. Nevertheless, we can obtain sufficient information to reconstruct the basic trends in the evolution of the landscape by studying the vegetational changes throughout the Pleistocene.

In addition, in the case of certain plants, the presence of only a few grains of pollen can provide significant information about the climate at the time. For example, in the case of the wild olive tree (Olea europaea), we can be certain that the climate could not have been very cold, since this typical Mediterranean species doesn’t tolerate frost. Nevertheless, interpreting pollen data doesn’t only consist of determining the presence or absence of certain species, but also in comparing the different frequencies with which each one occurs and their variation throughout the stratigraphic sequence. Of particular significance is the production of tree pollen compared with other species, or the presence of herbaceous plants typical of steppe-like environments.

One additional problem is that in many cases, pollen grains cannot be identified to the species level, and we have to be satisfied knowing only the genus. This is why we often speak of the presence of oak trees in the Sierra without being able to be more precise.

Fauna

As we have seen, if we were walking around in the Sierra de Atapuerca during the Pleistocene, the landscape would be familiar to us. Anyone who has recently traveled throughout the Iberian Peninsula would recognize the vegetation and would remember having seen similar landscapes. However, the same can’t be said of the animals we would find. Lions, jaguars, sabre-tooth cats or hyenas were competing with the hominids in hunting or scavenging the remains of rhinoceroses, bison, giant deer, elephants and even hippopotamuses.

Extensión de los hielos en Europa durante la última glaciación Just as the landscape changed through the course of time, so did the animals, particularly the mammals, which inhabited the Sierra and its immediate surroundings, although not in a cyclical manner. This is clearly different from the rest of Europe, where we can speak of a “glacial fauna” and an 'interglacial fauna' composed of different species and which parallel the climatic and vegetational changes. During the glacial periods, species which couldn’t survive in the cold central European steppes, such as deer, hippopotamuses or wild boar, disappeared from these areas and were replaced by more cold-adapted species which during interglacial times lived further north, such as arctic foxes, reindeer, wooly rhinos and mammoths. At Atapuerca, we can only speak of faunal complexes which occur through time. However, rather than representing an adaptation to two climatic extremes, changes in species composition are due to evolutionary factors or migration. An important type fossil at Atapuerca is the water vole, 'Mimomys savini', which went extinct a little more than 500,000 years ago and was replaced by another species, 'Arvicola cantianus', which was very similar to living voles. Thus, the fossils that are associated with Mimomys savini are all older than 500,000 years, while those which are found with 'Arvicola cantianus' are more recent in time.

The three sites where most work has been done to date are the Gran Dolina, Galería and the Sima de los Huesos. We know there are both older and younger fossils in other caves in the Sierra which are currently being prospected. The Sima del Elefante in the Trinchera contains the oldest levels in Atapuerca, while el Mirador and the site of Portalón, in the entrance to Cueva Mayor, have the youngest. In the Gran Dolina, 'Mimomys savini' is found from levels TD3 through the lower part of TD8, some eight and a half meters of sediments which span the time period from nearly one million years ago to just older than 500,000 years ago.

En colores claros, la distribución máxima del mamut lanudo en Eurasia durante la última glaciación The large mammals that inhabited the Sierra de Atapuerca in ancient times were quite varied and would have been a spectacular sight to see. In terms of the herbivores, there were large double-horned rhinoceroses of the species 'Stephanorhinus etruscus', wild boar, horses, long-legged bison and deer. There were also primitive giant deer and mammoths. In TD7, the hind legs of a musk ox were recovered, a relative of the species currently alive today. During this time period, the musk ox was still not adapted to periglacial environments, preferring pastures in close proximity to wooded areas. Further, hippos swam in the Arlanzón River and its tributaries, where beavers built their dams.

In addition to beavers, two other large sized rodents worth mentioning have been recovered from the oldest levels in the Gran Dolina: the porcupine ('Hystrix refossa') and the marmot. The living species of porcupine which are most similar to the form found at Gran Dolina inhabit warm climates in Africa and Asia, although this was a common taxon during the European Pleistocene as well, particularly during milder times. The marmot, on the other hand, lives in the cold climates of the alpine fields above the tree line and hibernates in its underground burrows. It is possible they inhabited the Sierra during colder times, between 600,000 and 900,000 years ago.

'Homotherium latidens', un tigre de dientes de sable,  fue uno de los grandes depredadores del Pleistoceno Inferior y Medio de Atapuerca Let’s take a look now at the hunters in these ancient ecosystems. Leaving aside for the moment the role of hominids, the largest predator was the saber-tooth cat, 'Homotherium latidens'. This large felid was about the size of a lion and had enormous curved upper canines with serrated edges. Although it disappeared from Western Europe a little less than 500,000 years ago, its cousin species 'Homotherium serum' survived in America until the end of the Ice Age. These powerful animals were capable of killing prey much larger than themselves, including young mammoths.

Another large felid found in the oldest levels in the Sierra de Atapuerca is the European jaguar, 'Panthera gombaszoegensis,' which went extinct some 400,000 years ago. It was smaller than the sabre-tooth cat, but larger than a leopard, meaning it was about the size of the modern American jaguar. A somewhat smaller felid, the lynx, was also present in the lower levels. The lion probably appeared in Europe around 600,000 years ago, and the sabre-tooth cat disappeared shortly thereafter, most likely due to this new competition for the place of top predator in the ecosystem. It seems then, that there were felids of varying sizes in the Atapuerca ecosystem, ranging from the large sabre-tooth cat to the small wildcat.

Among the canids, remains of two species have been recovered: 'Vulpes praeglacialis', an ancestor of the arctic fox which was not yet adapted to periglacial environments, and 'Canis mosbachensis', a small wolf, not much larger than the living jackal. This wolf increased in size, becoming the modern species of wolf, some 300,000 years ago.

Cráneo de hiena manchada ('Crocuta crocuta') de TD4 (Atapuerca) The lower levels of Gran Dolina have also yielded the oldest remains of the spotted hyena ('Crocuta crocuta') in Europe, a social carnivore and powerful competitor of hominids in both hunting and scavenging carrion. With their specialized dentition, spotted hyenas were capable of breaking the long bones of large herbivores to access the bone marrow inside which is rich in nutrients and fat, and is a prized resource.

Numerous cave bear remains have also been found in the lower levels of Gran Dolina, and, based on these fossils, a new species has been identified, 'Ursus dolinensis'. This primitive species represents the ancestor of the cave bear and is also evolutionarily very close to the precursor of the brown bear.

The deposits which are younger than 500,000 years ago also contain a large number of fossils. At the site of Galería, remains of the cuon, the same species as the modern-day Asiatic wild dog, or dhole, have been recovered. Due to the scarcity of the bison remains, it is difficult to identify the species represented at Atapuerca. It could be either 'Bison schoetensacki', the so-called forest buffalo, or 'Bison priscus', the larger steppe buffalo. Some of the bovine remains could represent the auroch ('Bos primigenius'), or even the water buffalo, which today is found only in Asia but inhabited Europe as well in the past. Unfortunately, it is difficult to distinguish between these different species based on isolated bones of the skeleton.

During this time there were also rhinoceroses belonging to the species 'Stephanorhinus hemitoechus'. This rhino grazed the steppes of Europe over a long time period, together with another larger species, Merck’s rhinoceros ('Stephanorhinus kirchbergensis)'. This latter species was truly impressive, measuring up to 2.5 meters tall, much taller than any living species. Both rhino species were adapted to milder climates and disappeared from Central Europe at the beginning of the last glaciation, during the time of the Neandertals. Finally, elephant remains have been found at Gran Dolina, a milk tooth from an undetermined species, and at the Sima del Elefante, for which the site is named.

Among the large rodents, marmots and porcupines ('Hystrix vinogradovi') continue to be present, as do horses, deer, giant deer, bison and rhinoceroses among the herbivores.

Atapuerca foto Javier Trueba. Mandíbula de León ('Panthera leo') procedente del nivel TG11 del yacimiento de Galería The carnivore assemblage from the Sima de los Huesos is comprised of many species and families. By far the most abundant, however, is the ancestor of the cave bear, Ursus deningeri. Wolves and foxes are also present, as are lynxes, evolutionarily related to the present-day Iberian lynx, and wildcats. Remains of lions have also been found, as well as an enigmatic felid species represented by a fragment of metatarsal (foot bone) which, judging by its size, could correspond to either a leopard or a European jaguar. Thus, there are four different sized felids in the Sima de los Huesos. Much smaller carnivores, the mustelids, are represented at the site by four different species: two larger forms, similar to the sable or marten and the badger, a smaller form, similar to the weasel or the stoat, and a medium-sized form, the polecat.

The absence of hyena remains in the Sima de los Huesos, Galería and in what has been excavated to date in the upper levels of Gran Dolina is surprising. One hypothesis to explain this absence is that the hominids competed with them, driving them out of the Sierra, or at least reducing their presence. Their earlier presence in the lower levels of Dolina could be attributed to several possible factors including a scarce human presence in the Sierra, poor social organization or short-term occupation of the Sierra on the part of hominids. Nevertheless, hyenas were abundant on the Iberian Peninsula in more recent Pleistocene times. It is possible that as hominids became more active hunters rather than scavengers, the ecological niche for which they had been competing was taken over again by hyenas.

Mandíbula de perro jaro ('Cuon alpinus') del yacimiento de la Galería (Atapuerca) The case of birds is quite different from that of the mammals just described. In the first place because the species which lived during Pleistocene times were practically the same as those found in Europe today. Thus, we can’t speak of evolutionary changes or extinctions, as with the mammals. In TD6, species which prefer open habitats, those which can be found today in high plateaus or grain-producing steppes on the Iberian Peninsula, are abundant, including various species of larks. However, other species also occur, such as the horned lark ('Eremophila alpestris'), which currently inhabits Finland and the Caucasus. Together with these species, others appear which are clearly associated with water, including the teal ('Anas crecca') or the black-tailed godwit ('Limosa limosa'), which indicate the existence of nearby lakes or marshes. Both ducks and other species typical of wetlands are abundant in unit GIII in Galería. From this same unit, we have also found remains of mallards ('Anas plathyrhyncos'), teal ('Anas querquedula'), widgeons ('Anas penelope'), Baillon’s crakes ('Porzana pusilla'), great snipes ('Gallinago media'), common snipes ('Gallinago gallinago') and terns ('Sterna albifrons'), among others. This abundance of birds related with rivers and lakes indicates the existence, some 300,000 years ago, of important wetlands in the region, although we also find birds typical of steppe environments in unit GIII, such as the great bustard ('Otis tarda').

It’s evident from the list of fossil species that there was a great faunal and floral diversity in the sierra de Atapuerca during the Pleistocene and that, as mentioned previously, the alternating climatic cycles didn’t drastically change the faunal composition of the region. This latter observation is not surprising in and of itself, since mammals in general, and large mammals in particular, are very plastic species that can adapt well to climatic changes of the magnitude that occurred in the Sierra throughout the Pleistocene.

Asta de ciervo del nivel TD10 de la Gran Dolina (Sierra de Atapuerca) The best explanation for this enormous diversity is the existence of a large number of diverse habitats in the Sierra and the surrounding region. These include ample plains, waterways, limestone bluffs and the nearby mountain tops, which would explain the coexistence in the fossil record of species typical of a forest habitat, such as deer and wild boar with others more adapted to open spaces such as giant deer and horses.

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