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100-425: From these sites have been obtained fossils of large mammals, mainly elephants ( Straight-tusked elephant ), with remains of nearly fifty individuals from each site, in addition to large bovines and horses. The sites can be seen as an elephants' graveyard , although the use of the term for a concentration of bones raises a problem of definition. The sites show evidence of successive occupations by human beings, who had

200-611: A hunting station or, more likely, scavanged carrion and carried out quartering. Known since the end of the 19th century, they were excavated first by the Marquis of Cerralbo between 1909 and 1914, later, in the early '60s and early '80s, by the American Francis Clark Howell with the collaboration of the paleontologist Emiliano Aguirre and later, in the '90s, new campaigns were carried out by Manuel Santonja and Alfredo Pérez-González . The remains from

300-457: A close common ancestry with Asian elephants and other species of Elephas , which was based on a number of morphological similarities between the two groups. In 2016, a mitochondrial DNA sequence analysis instead found that the mitochondrial genome of P. antiquus was nested within those of the African forest elephant ( Loxodonta cyclotis ), with analysis of a partial nuclear genome supporting

400-491: A closer relationship to L. cyclotis than to the African bush elephant ( L. africana ). A subsequent study published in 2018 that includes some of the same authors presented a complete nuclear genome sequence, indicating a more complicated relationship between straight-tusked elephants and other species of elephants. According to this study, the lineage of Palaeoloxodon antiquus was the result of reticulate evolution , with

500-499: A granulometry higher than the previous ones, corresponds again to alluvial fan facies, in which neither fossils nor lithic industry have been found. The Torralba site, in turn, is integrated into the Torralba Formation . It would be more recent than Ambrona, formed at a later time, not determinable, of the encasement of the fluvial network in the valley, although Aguirre it supposes a probable temporary overlap between

600-494: A great bovid ( Aurochs ), 25 deer ( Cervus elaphus ) and 3 rhinoceros ( Stephanorhinus hemitoechus ), and the lithic industry accounted for a total of 557 specimens, including hand axes , cleavers , flakes , cores and Chopping tools . Cerralbo was accompanied in his excavations by the archaeologist Juan Cabré , the geologist Pedro Palacios and the French paleontologist Édouard Harlé . The international diffusion of

700-410: A large male specimen from Neumark Nord that has a deep puncture hole wound in its forehead with surrounding bone growth indicating that it had healed, as well as another large male from the same locality with a healed puncture hole wound in its scapula. Like modern elephants, the herds would have been restricted to areas with available fresh water due to the greater hydration needs and lower mobility of

800-461: A large number of archeology students, reaching over fifty in one of the campaigns. In Ambrona a total of 688 m were excavated and some surveys and control tastings were carried out in Torralba. Some 975 lithic industry specimens were obtained, however, most paleontological remains were left unexploded, consolidated, covered again and protected to prevent spoilage and looting, in anticipation of

900-439: A large number of scientific publications on all related aspects, paleontology, archeology, geology, paleoclimatology, etc., but highlighting, due to its social impact, those related to the presumed hunting activities of primitive man. As a result of the results of Howell and collaborators, in the following years, extensive discussions took place on some conclusions related to human behavior, mainly those related to active hunting or

1000-408: A medium-high salinity is maintained, but with a lower water layer, decreasing until disappearing towards the roof of the unit. The ostracod Leucocythere mirabilis , present throughout the sequence (AS1 to AS6), indicates oligotrophic water conditions. On the other hand, the salinity of the waters could not be very high, as indicated by the presence of certain ostracods. The set of frogs indicates

1100-513: A more benign environment than the current one, with less dry summers and less cold winters. The age (stage of ontogenetic development) of some specimens indicates that death occurred between March and summer, most likely in the spring. The bermejuela , the only fish found in these sites, indicates a small fluvial course or lagoon not very extensive. Some of the birds found are typical of lacustrine areas with thick marginal vegetation, but no remains of any diving species have appeared, indicating that

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1200-479: A possible future extension of the museum exhibition in situ to a much larger extension of the site. The works were followed by numerous publications, highlighting an extensive monographic volume of the magazine Zona Arqueológica (vol.5, 2005). The sediments in which the deposits are integrated correspond to ancient fluvium - Lake , Discordant deposits, by a soft local pale-relief, on the Triassic materials of

1300-610: A result of the natural break during the activity of the animals in the barking of trees or the digging of the ground looking for water, as is the case with today's elephants. On the other hand, Aguirre, among others, maintains that it is very probable a certain rudimentary industry of trihedral and other bone elements endowed with points and edges. Likewise, Aguirre interprets the tips of ivory of defenses as possible soft hammerstones , selected and purposely prepared by man for use in lithic carving. These affirmations are based on direct experimentation with current elephant bones and comparing

1400-486: A straight-tusked elephant were collected from travertine deposits near Burgtonna in what is now Thuringia , Germany. While these remains were originally declared to be purely mineral in nature by the Collegium Medicum in the nearby city of Gotha , Wilhelm Ernst Tentzel, a polymath in the employ of the ducal court of Saxe-Gotha-Altenburg , correctly identified them as elephant remains. The Burgtonna skeleton

1500-400: A subgenus of Loxodonta (which includes the living African elephants). The species has a confused taxonomic history, with at least 21 named synonyms . In publications in the 1930s and 1940s, Henry Fairfield Osborn assigned the species to its own genus Hesperoloxodon, which was followed by some later authors, but is now rejected. In his widely cited 1973 work, Origin and evolution of

1600-568: Is 3.9 metres (13 ft) in length, has a circumference of around 77 centimetres (30 in) where it would have exited the skull, and is estimated to have weighed over 190 kilograms (420 lb) in life. Females reached shoulder heights and weights rarely exceeding 3 metres (9.8 ft) and 5.5 tonnes (12,000 lb) respectively, for comparison, female African bush elephants reach an average shoulder height of 2.6 metres (8.5 ft) and body mass of 3 tonnes (6,600 lb) under optimal growth conditions. Newborn and young calves were likely around

1700-578: Is a mandible (lower jaw) with a second molar (M2006). The exact provenance of the specimen is unknown, though it probably originates from Britain, and possibly the site of Grays in Essex , southeast England. The common name "straight-tusked elephant" was used for the species as early as 1873 by William Boyd Dawkins . In 1924, the Japanese paleontologist Matsumoto Hikoshichirō assigned E. antiquus to his new taxon Palaeoloxodon , which he classified as

1800-774: Is believed to derive from the migration of a population of Palaeoloxodon recki out of Africa, suggested to have occurred around 800,000 years ago, approximately at the boundary between the Early Pleistocene and Middle Pleistocene . P. antiquus first appeared during the Middle Pleistocene, with the earliest record of Palaeoloxodon in Europe being from the Slivia site in Italy, dating to around 800-700,000 years ago. Its earliest known appearance in northern Europe

1900-591: Is in England around 600,000 years ago. The arrival of Palaeoloxodon in Europe coincided with the extinction of the temperate-adapted European mammoth species Mammuthus meridionalis and the migration of Mammuthus trogontherii (the steppe mammoth) into Europe from Asia. The arrival of Palaeoloxodon in Europe was part of a larger faunal turnover event around the transition between the Early and Middle Pleistocene, where many European mammal species that characterised

2000-578: Is little evidence that straight-tusked elephants were targeted preferentially over smaller animals. Most individuals at these sites are subadult to adult and primarily male in sex. The male sex bias likely both represents the fact that adult males, despite their larger size, were more vulnerable targets due to their solitary nature, as well as the tendency of adult male elephants to engage in risky behavior causing them to more frequently die in natural traps, as well as being weakened or killed by injuries caused by combat with other male elephants during musth. At

2100-507: Is part of the genus Palaeoloxodon (whose other members are also sometimes called straight-tusked elephants), which emerged in Africa during the Early Pleistocene , before dispersing across Eurasia at the beginning of the Middle Pleistocene, with the earliest record of Palaeoloxodon in Europe dated to around 800–700,000 years ago. The straight-tusked elephant is the ancestor of a number of species of dwarf elephants that inhabited islands in

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2200-543: Is uncertain. The extinction was part of the Late Pleistocene megafauna extinctions , which resulted in the extinction of most large terrestrial mammals globally. The extinction of P. antiquus and other temperate adapted European megafauna has resulted in a severe loss of functional diversity in European ecosystems. Cleaver (tool) In archaeology , a cleaver is a type of biface stone tool of

2300-478: Is unclear if it was used for heavy digging or not. More experiments were shown that the cleaver in Africa was more used as a butchering instrument. They were also helpful in skinning large game, and breaking bones. In Africa, cleavers first appeared from the late Oldowan to the Acheulean . In Peninj on the western shore of Lake Notron, cleavers constitute 16% of all findings. One type of cleaver made in Africa

2400-480: Is very controversial. On the one hand, Villa and collaborators, based on the taphonomic analysis of the types of breaks and superficial alterations of the bones deny the existence of bone industry, at least for the Ambrona site, leaving Torralba with indetermination. In the deposits, more than fifty tips of young elephant tusks have been collected, after almost a century of excavations, which these authors interpret as

2500-552: The Geological and Mining Institute of Spain , with the designations "VP -07: Loma del Saúco, Torralba" and "VP-07b: Loma de los Huesos, Ambrona", within the category "vertebrate sites of the Pliocene-Spanish Pleistocene". In the late 19th century it was decided that a new railway line should link Torralba and Soria . The contract was awarded to a Belgian company. The first remains appeared in 1888, with

2600-486: The Lower Palaeolithic . Cleavers resemble hand axes in that they are large and oblong or U-shaped tools meant to be held in the hand. But, unlike hand axes, they have a wide, straight cutting edge running at right angles to the axis of the tool. Acheulean cleavers resemble handaxes but with the pointed end truncated away. Flake cleavers have a cutting edge created by a tranchet flake being struck from

2700-743: The Mediterranean . The species became extinct during the latter half of the Last Glacial Period , with the youngest remains found in the Iberian Peninsula , dating to around 44,000 years ago. Possible even younger records include a single tooth from the Netherlands that has been dated to around 37,000 years ago, and footprints from the southern part of the Iberian peninsula dated to 28,000 years ago. The body, including

2800-927: The Middle and Late Pleistocene . One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of 4 metres (13 ft) and a weight of 13 tonnes (29,000 lb). Straight-tusked elephants likely lived very similarly to modern elephants, with herds of adult females and juveniles and solitary adult males. The species was primarily associated with temperate and Mediterranean woodland and forest habitats, flourishing during interglacial periods, when its range would extend across Europe as far north as Great Britain and Denmark and eastwards into Russia, while persisting in southern Europe during glacial periods . Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors. The species

2900-1025: The Palaeoloxodon antiquus large-mammal assemblage, along with other temperate adapted megafauna species, including the hippopotamus ( Hippopotamus amphibius ), rhinoceroses belonging to the genus Stephanorhinus ( Merck's rhinoceros S. kirchbergensis and the narrow-nosed rhinoceros S. hemitoechus ), the European water buffalo ( Bubalus murrensis ), bison ( Bison spp.), Irish elk ( Megaloceros giganteus ), aurochs ( Bos primigenius ), European fallow deer ( Dama dama ), roe deer ( Capreolus capreolus), red deer ( Cervus elaphus ), moose ( Alces alces ), wild horse ( Equus ferus ) and wild boar ( Sus scrofa ). Carnivores included European leopards ( Panthera pardus spelaea ), cave hyenas ( Crocuta spelaea ), cave/steppe lions ( Panthera spelaea ), wolves ( Canis lupus ) and brown bears ( Ursus arctos ). Some authors have argued, in accordance with

3000-555: The Real Academia de la Historia – and established his paleontological workshop in Santa María de Huerta , a relatively close location where he owned an estate. Cerralbo excavated between 1000 and 2000 m of the Torralba site and an unknown, but much smaller, area of Ambrona. Paleontological elements recovered accounted for 525 elephant remains ( Straight-tusked elephant ), 86 horse ( Equus caballus torralbae ), 37 of

3100-598: The Tagus river ) in central-eastern Portugal, originally dated to around 34–33,000 years ago, but later revised to around 44,000 years ago. A late date of around 37,400 years ago has been reported from a single molar found in the North Sea off the coast of the Netherlands, but it has been suggested that this date needs independent confirmation, due to only representing a single sample. Some authors have suggested that P. antiquus likely survived until around 28,000 years ago in

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3200-528: The Vera/wood-pasture hypothesis , that the effects of straight-tusked elephants and other extinct megafauna on vegetation likely resulted in increased openness of woodland habitats, though this conclusion has been disputed by other authors. Potential gnaw marks suggested to have been made by hyenas and lions on the bones of straight-tusked elephants have been reported at some localities, which suggests that these species likely at least scavenged on

3300-458: The occipital bone of the skull roof which projects forwards and overhangs the rest of the skull. The crest was probably an anchor for muscles, including the splenius , as well as an additional muscle layer that wrapped around the top of the head, called the "extra splenius". The latter was likely similar to the "splenius superficialis" found in Asian elephants . The crest likely developed to support

3400-783: The Dirección General de Bellas Artes), Francisco Jordá Cerdá (professor at the University of Salamanca, delegate of Fine Arts in 1962–63), Desmond Collins (University of Cambridge, United Kingdom), Peter Taylor, Richard G. Klein, Blanca Izquierdo, José Viloria (MNCN, preparation and restoration of fossils), Karl W. Butzer (University of Wisconsin, edaphology and geology ), Josefina Menéndez Amor (MNCN) and F. Florschutz ( palynology and paleobotany , Leslie Gordon Freeman (University of Chicago, record), Thomas Lynch (University of Chicago), Susan Tax (cartoonist), several Spanish and American students denses and more than twenty workers in

3500-625: The Elephantidae , Vincent J. Maglio sunk P. antiquus into the South Asian P. namadicus , as well as Palaeoloxodon back into Elephas (which contains the living Asian elephant ). While the sinking of Palaeoloxodon into Elephas (with Palaeoloxodon sometimes being treated as a subgenus of Elephas ) gained considerable traction in the following decades, today both P. antiquus and Palaeoloxodon are considered distinct. Traditionally, Palaeoloxodon species were thought to share

3600-557: The Lehringen site in north Germany, dating to the Eemian/Last Interglacial (around 130-115,000 years ago) a skeleton of P. antiquus was found with a spear made of yew wood between its ribs, with flint artifacts found close by, providing unequivocal evidence that this specimen was hunted (though the elephant may have already been mired prior to being killed ). Studies in 2023 proposed that in addition to Lehringen,

3700-471: The Marquis of Cerralbo vacationed in the area, he had news of the appearance of "colossal" elephant carcasses; after visiting the place and aware, from the beginning, of the antiquity of the remains, he decided to undertake and pay for the excavations himself, hoping to find evidence of its synchrony with the "most primitive" man. He initiated them in 1909 – a year after taking office as Permanent Academician of

3800-422: The Marquis of Cerralbo. His study, scientific and methodical, will continue without rest. He could discover the skeletons of some elephant hunters! The first excavations were carried out by Enrique de Aguilera y Gamboa , Marquis of Cerralbo, first in Torralba from 1909 to 1913 and later in Ambrona from 1914 to 1916, and have been considered as the best performed of the first half of the 20th century. In 1907, when

3900-541: The Museum of Ambrona, built ten years earlier, necessary to correct the humidity that endangered it, recovering more fossils and lithic industry. The last campaigns of Howel were realized in 1980, 1981 and 1983. The possibility of finding some human fossil facilitated new economic supports, even of the National Geographic Society . For the excavations and analysis of samples of these campaigns, he had

4000-583: The Neumark Nord, Taubach and Gröbern sites, which show evidence of systematic butchery, provided evidence of widespread hunting of straight-tusked elephants by Neanderthals during the Eemian in Germany. The remains of at least 57 elephants were found at Neumark Nord; the study authors estimated that they accumulated over a time span of around 300 years and that one elephant was hunted once every 5–6 years at

4100-455: The anthropologists F. Clark Howell (University of Chicago) and Pierre Biberson (Museum of the Man of Paris), in the works that the Marquis of Cerralbo had done in Torralba and his concept of "station", very similar to the one they were discussing. Howell visited Ambrona and Torralba in 1960. He obtained funding and permits to excavate, helped by Biberson, who was also able to provide some funding for

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4200-424: The area (limestone and quartz). Between 20% and 65% of the pieces, according to the levels, they seem not to be eroded, while the rest show signs of light bearing, secondary accumulation, with few elements very rolled. There is no evidence of in situ manufacture, only of secondary retouching , and mainly on quartzite elements. The existence or not of bone industry from bones and elephant defenses in these sites

4300-547: The area (locally constituted by lutites and gypsums facies of type Keuper ). They were located in the flat and impermeable bottom of an elongated valley from an ancient polje , then open and in the process of erosion. The Ambrona site is located in the "Ambrona formation", whose sediment thickness, as a whole, would not exceed eight meters. Seven levels or litic facies have been identified, grouped into three members: one lower (AS1 to AS5 levels) – gravels, gray silts and clays, another medium (AS6) – sands and gray limes, and

4400-408: The area. Grids were drawn, stratigraphic profiles were raised and each extracted remainder was labeled. As an example of thoroughness, samples of pollen were taken from the clay adhered between the teeth of the elephants, to get as close as possible to the existing environment during the accumulation of the remains. In 1973 Aguirre directed the systematic excavation of more than 200 m around

4500-539: The association and characteristics typical of mid-Middle Pleistocene , between 400 000 and 300  of Ionian age, which lasted from 781 000 to 126 000 years before the present). The most characteristic temporal marker is the Arvicolinae Microtus ( I. ) brecciensis , which in Ambrona presents some primitive characters, as does Palaeoloxodon antiquus and Bos primigenius , prior to populations recorded in late-Middle Pleistocene sites. On

4600-403: The bones of straight-tusked elephants and in at least one case their ivory were used to make tools. There is evidence that exploitation of straight-tusked elephants in Europe increased and became more systematic from the mid-Middle Pleistocene (around 500,000 years ago) onwards. Based on analysis of sites of straight-tusked elephants with cut marks and/or artifacts, it has been argued that there

4700-696: The cave of around 66–65,000 years ago. Another late Italian record has been reported from Mousterian layers in Barma Grande cave in northwest Italy, probably dating to around the same time as Grotta Guattari, which has been suggested to display evidence of butchery by Neanderthals. Other late remains have been reported from several sites in the Iberian Peninsula , including El Castillo cave in northern Spain, which were initially radiocarbon dated to 43,000 years ago, with later calibration suggesting an age of around 49,570–44,250 years Before Present , and Foz do Enxarrique (a sequence of terrace deposits of

4800-663: The central part of the Peloponnese peninsula in southern Greece was known for its enormous bones, which Pausanias reported were considered to be those of giants who died during the Gigantomachy , a mythic climactic battle between the giants and the Greek gods. Given that this region is today known for its straight-tusked elephant fossils, it is plausible that at least some of the giant bones to which Pausanias referred were those of straight-tusked elephants. In 1695, remains of

4900-470: The continent during warm interglacial periods. Fossils are also known from Israel, western Iran and probably Turkey in West Asia . Some remains of the species have also been reported from Central Asia in northeastern Kazakhstan and Tajikistan . Outside of Eastern Europe, the northernmost records of the species are known from Great Britain, Denmark and Poland, around the 55th parallel north . Some of

5000-455: The crest being more pronounced in older individuals, as well as due to distortion during fossilisation ( taphonomic variation). P. antiquus differs from P. namadicus in having a less stout cranium and more robust limb bones, and in lacking a teardrop-shaped indentation behind the eye socket (infraorbital depression). The premaxillary bones (which contain the tusks) are fan-shaped and very broad in front view. The tusks are very long relative to

5100-407: The current one. For the lower sections of the sequence (AS1 to AS5) an initial environment is inferred with grasses, riparian trees (alder, willow and elm) and few pines, after which an environment of moorland with junipers, alternating, with an increase towards the end, with pine forests. The AS6 level is characterized by the almost exclusive domain of the pine forests, but on the roof, the return of

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5200-442: The deposits have been changing over time, depending on the discovery of better diagnostic elements or the vagaries of the systematic ; we try to present the most up-to-date relation. The vegetation, identified by the pollen at different levels, shows the evolution of the paleoenvironment during the sedimentation of the same, which in general corresponds to fluvio-lacustrine media under a temperate climate, softer and more humid than

5300-518: The desired end product that is now known as the flake-cleaver. There are collections of these flakes in the Institute of Human Paleontology in Paris. They include larger hand axes and smaller hand axes ranging from 6–26 cm, and large flake cleavers that were commonly found in this area. Another way to make the tools was known as the "Victoria West" technique, termed by Goodwin in 1934. This process

5400-419: The diet of P. antiquus was highly variable, ranging from almost completely grazing to almost completely browsing (feeding on leaves, stems and fruits of high-growing plants). However, microwear only reflects the diet in the last few days or weeks before death, so the observed dietary variation may be seasonal, as is the case with living elephants. Isotopic analysis of a specimen from Greece suggests that it

5500-636: The different excavations are scattered, mainly, between the Museo Nacional de Ciencias Naturales , the Museo Arqueológico Nacional , the Museo Numantino de Soria and the museum in situ of Ambrona. They were declared Bien de Interés Cultural in the category of "archaeological zone" on September 7, 1995. They are also declared as "a place of international geological interest of international relevance" ("Geosite") by

5600-527: The elephants were hunted or scavenged, though both scavenging of already dead elephants and active hunting are likely to have occurred. Straight-tusked elephant butchery sites have been found in Israel, Spain, Italy, Greece, Britain, and Germany. These sites are likely attributable to Homo heidelbergensis and Neanderthals . Stone tools used at these sites include flakes , choppers , bifacial tools like handaxes , as well as cores . At some sites,

5700-447: The end of the Eemian interglacial around 115,000 years ago due to climatic conditions becoming unfavourable, and fossils after that time during the Last Glacial Period are rare. A molar from the cave deposits of Grotta Guattari in central Italy has been suggested to date to around 57,000 years ago, though other studies have found it to have an older early Late Pleistocene age, and later dating done in 2023 suggested an age of deposition in

5800-468: The findings of Torralba, originally published in Spanish in 1916, with a second edition expanded in 1925 that was translated into English. The American anthropologist Francis Clark Howell conducted six excavation campaigns in Torralba and Ambrona, between 1961 and 1963 and in 1980, 1981 and 1983. The results of his studies seemed to have demonstrated the practice of active hunting by the human groups of

5900-847: The following team: co-directors: Leslie Gordon Freeman ( lithic industry ) and Martín Almagro Basch (director of the Museo Arqueológico Nacional de España ); researchers: Emiliano Aguirre, Karl W. Butzer, Richard G. Klein, M. Teresa Alberdi, A. Azzaroli, J. Bischoff, T. E. Cerling, Katherine Cruz-Uribe, Ignacio Doadrio, Frank Harrold, Manuel Hoyos, P. Preece, Antonio Sánchez-Marco (birds), F. Borja Sanchiz (amphibians), H. P. Schwarcz, Carmen Sesé (micromammals), Kathy Schick, N. P. Toth and Charles Turner. In all its campaigns, Howell excavated more than 1000 m in Torralba, recovering about 700 lithic instruments and more than 2100 fossils, and about 2700 m in Ambrona, with more than 4400 lithic instruments and several thousand fossils (of them more of 2000 elephants). The investigations of these years gave rise to

6000-426: The holotype of Palaeoloxodon turkmenicus , a skull found in western Turkmenistan, to P. antiquus , but later analysis found that P. turkmenicus represented a morphologically distinct and valid species. The straight-tusked elephant is primarily associated with temperate and Mediterranean forest and woodland habitats, as opposed to the colder open steppe environments inhabited by contemporary mammoths, though

6100-577: The juveniles. Fossil tracks of newborns, calves and adults, which are likely of a herd of P. antiquus , have been found in dune deposits in southern Spain, dating to the early Late Pleistocene ( Marine Isotope Stage (MIS) 5 , around 130–80,000 years ago). Due to their larger size, straight-tusked elephants are thought to have finished growing 10 to 15 years later than living elephants, continuing to grow after 50 years of age. They may also have lived longer than extant elephants, with lifespans perhaps in excess of 80 years. Dental microwear studies suggest that

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6200-476: The lagoon would be shallow, of shallow depth. Mammals indicate forest environments ( Cervus , Dama , Capreolus ) and open areas with meadows ( Microtus brecciensis , Palaeoloxodon , Stephanorhinus , Bos , Dama , Capreolus ), with areas of abundant water ( Arvicola , Palaeoloxodon ) and more temperate and humid climate than the current one. Some Pleistocene mammals may provide some biochronologic resolution, and those represented in Torralba and Ambrona have

6300-405: The largest known land mammals to have ever lived. Under optimal conditions where individuals were capable of reaching full growth potential, 90% of mature fully grown straight-tusked elephant bulls are estimated to have had shoulder heights in the region of 3.8–4.2 m (12.5–13.8 ft) and a weight between 10.8–15 tonnes (24,000–33,000 lb). For comparison, 90% of mature fully grown bulls of

6400-684: The largest living elephant species, the African bush elephant under optimal growth conditions have heights between 3.04 to 3.36 metres (10.0 to 11.0 ft) and masses between 5.2–6.9 tonnes (11,000–15,000 lb). Extremely large bulls, such as those represented by a now lost pelvis and tibia collected from Spain in the late 19th century, may have reached shoulder heights of 4.6 m (15.1 ft) and body masses of over 19 tonnes (42,000 lb). Adult males had tusks typically around 3.5–4 metres (11–13 ft) long, with masses comfortably exceeding 100 kilograms (220 lb). The preserved portion of one particularly large and thick tusk from Aniene, Italy,

6500-448: The late Acheulean primitive or middle-ancient Acheulian, dated in the «full» Middle Pleistocene . The typology is diverse: hand axes , cleavers , racloirs , denticulates , perforators , back knife or burins , among others. The materials are very varied: flint , quartzite , limestone and quartz , from different origins, some transported from long distances (flint and quartzite) and others taking advantage of those available in

6600-730: The majority of the genome of straight-tusked elephants deriving from a lineage of elephants that was most closely related but basal to the common ancestor of forest and bush elephants (~60% of total genomic contribution), which had significant introgressed ancestry from African forest elephants (>33%) and to a lesser extent from mammoths (~5%). The African forest elephant ancestry was more closely related to modern West African forest elephants than to other African forest elephant populations. This hybridisation likely occurred in Africa, prior to migration of Palaeoloxodon into Eurasia, and appears to be shared with other Palaeoloxodon species. Like other Eurasian Palaeoloxodon species , P. antiquus

6700-455: The methods of electron paramagnetic resonance (EPR) and uranium-thorium (U-Th), on the enamel and dentin of horse teeth , a dating has been obtained absolute not inferior to 350,000 years, contemporary of OSI 9 or final of OSI 11. The studies of paleomagnetism have given in all Ambrona samples a normal polarity value, consistent with the cron Brunhes , the current one, which started 779 000 years ago . The lists of taxa identified in

6800-502: The moorlands is finally reflected with junipers and grasses (Poaceae). The diatoms indicate that during the sedimentation of the AS4 and AS5 units the salinity increased in the lagoon and that the water layer was somewhat higher with respect to the previous units -the salts would be contributed by the sediments of the underwater Keuper facies that surround all area-. For the AS6 unit they show that

6900-411: The most represented mammals in all the fossiliferous levels, except in the AS6 levels of Ambrona (middle member), in which only horse remains appear, and Va de Torralba (upper complex), in which the horse is clearly predominant over the rest. The lithic industry recovered from these sites has been very numerous, although in relation to the excavated volumes it can be considered scarce. It corresponds to

7000-702: The northernmost reported fossils of the species are from the banks of the Kolva river in the Russian Urals at around the 60th parallel north . During glacial periods P. antiquus permanently resided in the Mediterranean region. Many of the West Asian remains have been assigned to the species primarily on the basis of geography, and it has been suggested that some of these, such as those from Israel, actually belong to P. recki . A 2004 study attributed

7100-946: The other hand, Arvicola aff. sapidus , from Ambrona, presents traits that indicate an age after the beginning of the Middle Pleistocene, coinciding with what is indicated by the presence of Palaeoloxodon . Other sites, with a similar faunal association of mammals or the evolutionary status of the significant species, are those of Áridos ( Arganda del Rey ) and some terraces of the Manzanares River ( Madrid ), Pinedo ( Toledo ), El Higuerón ( Rincón de la Victoria , Province of Málaga ) or Solana del Zamborino ( Fonelas , Province of Granada ). The bone remains of large mammals are, in general, dispersed, eroded and fragmented, evidencing trawling by fluvial transport, although in clay or silty sediments there are usually elements in anatomical connection, with little or no transport (primary accumulation), and without predators or scavengers, even

7200-414: The pelvis, of P. antiquus was broad relative to extant elephants . The forelimbs, particularly the humerus, and the scapula are proportionally longer than those of living elephants, resulting in a high position of the shoulder. The head represents the highest point of the animal, with the back being somewhat sloped though irregular in shape. The spines of the back vertebrae are noticeably elongate. The tail

7300-512: The preceding late Villafranchian became extinct, along with the dispersal of immigrant species into Europe from Asia and Africa. There appears to be no overlap between M. meridionalis and P. antiquus , which suggests that the latter might have outcompeted the former. During P. antiquus 's hundreds of thousands of years of existence, its tooth morphology remained relatively static, unlike European mammoth populations. A number of species of dwarf elephants that are thought to have evolved from

7400-417: The primary surface. Cleavers, found in many Acheulean assemblages such as Africa, were similar in size and manner of hand axes. The differences between a hand axe and a cleaver is that a hand axe has a more pointed tip, while a cleaver will have a more transverse "bit" that consists of an untrimmed portion of the edge oriented perpendicular to the long axis of the tool. These were used in lithic technology. It

7500-415: The remains of straight-tusked elephants like lions and hyenas do on elephants in Africa today. Remains of juvenile straight-tusked elephants are also known from Kirkdale Cave in northern England, a well known cave hyena den. Remains of straight-tusked elephants at numerous sites are associated with stone tools and/or bear cut and percussion marks indicative of butchery. At most sites it is unclear whether

7600-496: The removal of large flakes. Cleavers can also be found made from different raw materials such as flint or limestone, but these are not nearly so common. There is also a chronological gap between the two lithic assemblages of the early core and flake techniques to the later Acheulian. Central France shows that both of these existed. One of the cleavers found in Central France was a 20 cm long cleaver. This particular one

7700-436: The results with the breaking and polishing marks of some bone elements of the sites, with sharpness and with the percussion polishing marks at the tips of the defenses, as well as in the relative abundance of these last ones against complete defenses. Straight-tusked elephant The straight-tusked elephant ( Palaeoloxodon antiquus ) is an extinct species of elephant that inhabited Europe and Western Asia during

7800-534: The same size as those of modern elephants. A largely complete 5 year old calf from Cova del Rinoceront in Spain was estimated to have a shoulder height of 178–187 centimetres (5.8–6.1 ft) and a body mass of 1.45–1.5 tonnes (3,200–3,300 lb), which is comparable to a similarly aged African bush elephant. In the second century AD , the Greek geographer Pausanias remarked that the Megalopolis region in

7900-941: The site. At the Bilzingsleben site in Germany and the Stránská Skála 1 site in the Czech Republic, bones of straight-tusked elephants have been found engraved with roughly parallel lines of unclear purpose. There are no cave paintings that unambiguously depict P. antiquus . An outline drawing of an elephant in El Castillo cave in Cantabria, Spain, as well as a drawing from Vermelhosa in Portugal have been suggested to possibly depict it, but these could also potentially depict woolly mammoths . Palaeoloxodon antiquus retreated from northern Europe after

8000-419: The size of the body and vary from straight to slightly curved. The teeth are high crowned ( hypsodont ), with each third molar having approximately 16–21 lamellae (ridges). The species was sexually dimorphic , with males being substantially larger than females; this size dimorphism was more pronounced than in living elephants. P. antiquus was on average considerably larger than any living elephant, and among

8100-685: The skeleton of an elephant specimen (in the so-called "α" concentration) is practically complete. On the other hand, there is evidence of human manipulation in some elephant bones: certain fractures and cut marks by lytic instruments for dying. The abundant accumulations of elephant remains in Ambrona are similar to the modern sites popularly known as " Elephants' graveyards ". In these places, close to water sources, corpses tend to accumulate, mainly elephants, but also other mammals, and have been well studied in African parks. In this environment trampling and fragmentation of exposed or semi-buried remains by elephants or other mammals occurs, which continue to go to

8200-488: The southern Iberian Peninsula based on footprints found in Southwest Portugal and Gibraltar. While some authors have argued that climate change was primarily responsible for the extinction of the straight-tusked elephant, others have argued that climate change alone cannot account for the species extinction. Human hunting has been suggested to have possibly played a contributory role, but the importance of this

8300-633: The species is also known to have inhabited open grasslands , and is thought to have been tolerant of a range of environmental conditions. As with modern elephants, female and juvenile straight-tusked elephants are thought to have lived in matriarchal herds of related individuals, with males leaving these groups to live solitarily upon reaching adolescence around 14–15 years of age. Adult males likely engaged in combat with each other during musth similar to living elephants. Some straight-tusked elephant specimens appear to document injuries obtained in fights with conspecifics; particularly notable specimens include

8400-418: The straight-tusked elephant are known from many Mediterranean islands , spanning from Sicily and Malta in the west to Cyprus in the east. The responsible factors for the dwarfing of island mammals are thought to include the reduction in food availability, predation and competition from other herbivores. Palaeoloxodon antiquus is known from abundant finds across Europe, reaching its widest distribution on

8500-646: The subfamily Lactuceae ). Straight-tusked elephants rarely coexisted alongside mammoths, although they occasionally did so, like at the Ilford locality in Britain that dates to the Marine Isotope Stage (MIS) 7 interglacial (~200,000 years ago) and where both steppe mammoths and P. antiquus are found. At this locality, the two species appear to have engaged in dietary niche partitioning . During interglacial periods, P. antiquus existed as part of

8600-615: The superior members of Ambrona and the inferior ones of Torralba. The maximum recognized thickness of this formation is about 15 meters. In 1965, Butzer differentiated thirteen units in the stratigraphic column , grouped into two "complexes", both with fossils and lithic industry: Above the Torralba Formation, the Sahuco Formation, of the Upper Pleistocene, absent in Ambrona, is superimposed. Combining

8700-488: The time-hypothesis discussed later, in favor of occasional scavenging. Also carbonaceous remains to indicate the presence of homes: the intentional and controlled use of fire. In 1959, during the Pan-African Congress of Prehistory and Quaternary Studies, in which John Desmond Clark was presenting the concept of "occupation sites", the Spanish archaeologist Lluís Pericot (University of Barcelona) interested

8800-510: The upper one (AS7) – graves and red sands. The AS3 level (silt and clays) is locally eroded, supporting AS4 directly on AS2 at some points. The base, discordant, corresponds to facies of alluvial fans channel (AS1 and AS2). The AS3 level has been interpreted as lacustrine environment. The rest of the lower and middle members (AS4 to AS6) are interpreted as deposited in low-energy shallow fluvial - lacustrine environments, with some sediments of channel and overflow . The upper member (AS7), with

8900-649: The use of bone instruments. In order to establish a precise formation model of the deposits, the archaeologist Manuel Santonja and the geologist Alfredo Pérez-González proposed and co-directed a new stage of excavations, focused mainly to establish with precision the geology and the detailed stratigraphy of the same. The approach was based on the realization, prior to the systematic excavation, of trial pits and sections for detailed stratigraphic analysis, since simultaneous excavation in large areas could lead to confusion among very similar facies , mixing levels that should be differentiated with this other method. The works began in

9000-494: The very large size of the head, as the skulls of Palaeoloxodon are the largest proportionally and in absolute size among proboscideans . Two morphs of P. antiquus were previously thought to exist in Europe on the basis of differences in the parieto-occipital crest, one more similar to the South Asian Palaeoloxodon namadicus . These differences were shown to be age-related ( ontogenetic variation), with

9100-450: The water sources. In Ambrona, the elephant mortality curve also fits into this model, seems to indicate deaths due to natural causes and does not present the typical bias due to selective hunting. From the excavations of Howell in Ambrona, the alignment of a defense and five long elephant bones of difficult interpretation have been described and in which it has been wanted to see, without any justification, some type of ritual. Elephants are

9200-443: The work. An international multidisciplinary team and a modern work methodology were proposed. In the different campaigns from 1961 to 1963 they were part of Howell's teams: Pierre Biberson (Museum of the Man of Paris, deputy director of the excavations and head of the Ambrona area), Emiliano Aguirre ( Museo Nacional de Ciencias Naturales , paleontology of vertebrates), Dolores Echaide (University of Zaragoza, representative in 1961 of

9300-606: The works in Torralba was due, on the one hand, to the communication that Marquis of Cerralbo himself presented at the International Congress of Prehistory that was held in Geneva in 1912, which he accompanied with a sample of his discoveries, and, on the other hand, to the book by the German paleontologist Hugo Obermaier , The fossil man -reference work during the first third of the 20th century-, in which he describe

9400-522: The works of canalization of the water that the company was carrying out for the first railway station of Torralba (moved later twice, before 1926 and in 1959). Part of that material was acquired by the Escuela Superior de Minas de Madrid and the rest distributed among individuals. The Torralba site is infinitely precious for Spanish prehistory and it is a joy that is in the hands of someone so enlightened and with such powerful means of action as

9500-989: The years 1990 and 1991, with the elaboration of surface geological studies complemented with some soundings, and the main excavation campaigns were carried out, this time only in Ambrona, the summers from 1993 to 2000, without interruption, taking place some complementary sampling and other trials between 2001 and 2002. The team had numerous specialists: Carmen Sesé and Enrique Soto (mammals), Paola Villa ( taphonomy ), Blanca Ruiz Zapata (palynology), Rafael Mora (area of Torralba, registry and cartography), Josep María Parés ( paleomagnetism ), Ángel Baltanás ( ostracods ), Ignacio Doadrio ( fishes ), Ascensión Pinilla ( phytoliths ), Borja Sanchiz (amphibians and reptiles), Antonio Sánchez Marco (birds), Juan M. Rodríguez de Tembleque, Joaquín Panera and Susana Rubio (archeology), Christophe Falguères (dating), Alfonso Benito Calvo (geology), C. Álvaro Chirveches, M. Vilà Margalef and Alexandra Vicent (consolidation and restoration). The excavations were carried out by

9600-548: Was formed by the Tavelbala Tachengit technique. This procedure was used in the north-west Sahara desert. It was done by detachment of large flakes for biface manufacture. This was a significant technological advance because the cleavers normally produced were made on large flakes, which was impossible to produce with using the nodular flint of the size used on the European continent. Once the large flakes were detached, there were only minor modifications needed to get

9700-480: Was one of the specimens that Johann Friedrich Blumenbach described in his publication naming the woolly mammoth ( Mammuthus primigenius , originally Elephas primigenius ) in 1799. The remains of straight-tusked elephants continued to be attributed to woolly mammoths until the 1840s. The straight-tusked elephant was scientifically named in 1847 by British palaeontologists Hugh Falconer and Proby Cautley as Elephas ( Euelephas ) antiquus . The type specimen

9800-877: Was primarily browsing during the dry (presumably summer) months and consumed more grass during the wet (presumably winter) months. Dental mesowear analysis suggests that the diet also varied according to local environmental conditions, with individuals occupying more grass-dominated open environments having a greater grazing-related wear signal. Preserved stomach contents of German specimens found at Neumark Nord suggests that in temperate Europe, its diet included trees such as maple , linden/lime , hornbeam , hazel , alder , beech , ash , oak , elm , spruce and possibly juniper , as well as other plants like ivy , Pyracantha , Artemisia , mistletoe ( Viscum ), thistles ( Carduus and Cirsium ), grass and sedges ( Carex ), as well as members of Apiaceae , Lauraceae , Rosaceae , Caryophyllaceae and Asteraceae (including

9900-438: Was relatively long. Although not preserved, the body was probably only sparsely covered in hair, similar to extant elephants, and probably had relatively large ears. The skull is proportionally both very wide and tall. Like many other members of the genus Palaeoloxodon , P. antiquus possesses a well-developed growth of bone at the top of the cranium above the nasal opening called the parieto-occipital crest, originating from

10000-550: Was used in the Middle Pleistocene of South Africa, and was the chosen method of producing large flakes that was used to create cleavers. This production date was between 285 k.A BP to 510 k.a. BP. Outside Africa where cleavers were most abundant, cleavers have appeared in Southwestern Europe. In these regions, they are more abundant than hand axes where raw material occurs in the form of large quartzite cobbles that do not need extensive decortication and shaping prior to

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