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''Homo heidelbergensis'' (also ''H. sapiens heidelbergensis'') is an extinct
species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individu ...

species
or
subspecies In biological classification In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interacti ...
of
archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (dependin ...
which existed during the
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocene Epoch withi ...
. It was subsumed as a subspecies of '' H. erectus'' in 1950 as ''H. e. heidelbergensis'', but towards the end of the century, it was more widely classified as its own species. It is debated whether or not to constrain ''H. heidelbergensis'' to only Europe or to also include African and Asian specimens, and this is further confounded by the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Bi ...
( Mauer 1) being a jawbone, because jawbones feature few diagnostic traits and are generally missing among Middle Pleistocene specimens. Thus, it is debated if some of these specimens could be split off into their own species or a subspecies of ''H. erectus''. Because the classification is so disputed, the Middle Pleistocene is often called the "muddle in the middle." ''H. heidelbergensis'' is regarded as a
chronospecies A chronospecies is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organ ...

chronospecies
, evolving from an African form of ''H. erectus'' (sometimes called '' H. ergaster''). By convention, ''H. heidelbergensis'' is placed as the
most recent common ancestor In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanis ...
between modern humans (''H. sapiens'' or ''H. s. sapiens'') and
Neanderthal Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, phys ...
s (''H. neanderthalensis'' or ''H. s. neanderthalensis''). Many specimens assigned to ''H. heidelbergensis'' likely existed well after the modern human/Neanderthal split. In the Middle Pleistocene, brain size averaged about 1,200
cubic centimetre A cubic centimetre (or cubic centimeter in US English) (SI unit symbol: cm3; non-SI abbreviations: cc and ccm) is a commonly used unit Unit may refer to: Arts and entertainment * UNIT Unit may refer to: Arts and entertainment * UNIT, a fictio ...
s (cc), comparable to modern humans. Height in the Middle Pleistocene can only be estimated off remains from 3 localities: Sima de los Huesos, Spain, for males and for females; for a female from Jinniushan, China; and for a specimen from
Kabwe Kabwe is the capital of the Zambian Central Province, Zambia, Central Province with a population estimated at 202,914 at the 2010 census. Named Broken Hill until 1966, it was founded when lead and zinc deposits were discovered in 1902. Kabwe also ...
, Zambia. Like Neanderthals, they had wide chests and were
robustRobustness is the property of being strong and healthy in constitution. When it is transposed into a system, it refers to the ability of tolerating perturbations that might affect the system’s functional body. In the same line ''robustness'' can be ...
overall. The Middle Pleistocene of Africa and Europe features the advent of Late
Acheulian Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the type site of Saint-Acheul (Amiens), Saint-Acheul, is an archaeological industry of stone tool manufacture characterized by distinctive oval and pear-shaped "han ...
technology, diverging from that of earlier and contemporary ''H. erectus'', and probably issuing from increasing intelligence. Fire likely became an integral part of daily life after 400,000 years ago, and this roughly coincides with more permanent and widespread occupation of Europe (above
45°N
45°N
), and the appearance of
hafting Hafting is a process by which an artifact, often bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitutes part of the vertebrate skeleton in animals. Bones protect the various organs of the body, produce red blood cell, red ...
technology to create spears. ''H. heidelbergensis'' may have been able to carry out coordinated hunting strategies, and consequently they seem to have had a higher dependence on meat.


Taxonomy


Research history

The first fossil, Mauer 1 (a jawbone), was discovered by a worker in Mauer, southeast of
Heidelberg Heidelberg () is a university town in the German state The Federal Republic of Germany, as a federal state, consists of sixteen partly sovereign federated states (german: Land (state), plural (states); commonly informally / federated s ...

Heidelberg
, Germany, in 1907. It was formally described the next year by German anthropologist Otto Schoetensack, who made it the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Bi ...
of a new species, ''Homo heidelbergensis''. He split this off as a new species primarily because of the mandible's archaicness—in particular its enormous size—and it was the then-oldest human jaw in the European fossil record at 640,000 years old. The mandible is well preserved, missing only the left
premolar The premolars, also called premolar teeth A tooth (plural teeth) is a hard, calcification, calcified structure found in the jaws (or mouths) of many vertebrates and used to Mastication, break down food. Some animals, particularly carnivores, al ...
s, part of the 1st left molar, the tip of the left coronoid process (at the jaw hinge), and fragments of the mid-section as the jaw was found in 2 pieces and had to be glued together. It may have belonged to a young adult based on slight wearing on the 3rd molar. In 1921, the skull
Kabwe 1 Kabwe 1 (also called the Broken Hill skull, Rhodesian Man) is a Middle Paleolithic The Middle Paleolithic (or Middle Palaeolithic) is the second subdivision of the Paleolithic or Old Stone Age as it is understood in Europe Europe is a c ...
was discovered by Swiss miner Tom Zwiglaar in
Kabwe Kabwe is the capital of the Zambian Central Province, Zambia, Central Province with a population estimated at 202,914 at the 2010 census. Named Broken Hill until 1966, it was founded when lead and zinc deposits were discovered in 1902. Kabwe also ...
, Zambia (at the time Broken Hill,
Northern Rhodesia Northern Rhodesia was a protectorate in southern Africa, south central Africa, formed in 1911 by Amalgamation (politics), amalgamating the two earlier protectorates of Barotziland-North-Western Rhodesia and North-Eastern Rhodesia.''Commonwealth ...
), and was assigned to a new species, "'' H. rhodesiensis''", by English palaeontologist
Arthur Smith Woodward Sir Arthur Smith Woodward, FRS (23 May 1864 – 2 September 1944) was an English English usually refers to: * English language English is a West Germanic languages, West Germanic language first spoken in History of Anglo-Saxon England ...

Arthur Smith Woodward
. These were two of the many putative species of
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocene Epoch withi ...
''Homo'' which were described throughout the first half of the 20th century. In the 1950s,
Ernst Mayr Ernst Walter Mayr (; 5 July 1904 – 3 February 2005) was one of the 20th century's leading evolutionary biologists File:Francesco Redi.jpg, Francesco Redi, the founder of biology, is recognized to be one of the greatest biologists of all ti ...
had entered the field of anthropology, and, surveying a "bewildering diversity of names," decided to define only three species of ''Homo'': "''H. transvaalensis''" (the
australopithecine Australopithecina or human taxonomy, Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' (cladistically including the genus, genera ''Homo'', ''Paranthropus'', and ''Kenyanthropus''), and ...
s), '' H. erectus'' (including the Mauer mandible, and various putative African and Asian taxa) and ''Homo sapiens'' (including anything younger than ''H. erectus'', such as modern humans and
Neanderthal Neanderthals (, also Neandertals, ''Homo neanderthalensis'' or ''Homo sapiens neanderthalensis'') are an extinct species In biology Biology is the natural science that studies life and living organisms, including their anatomy, phys ...
s). Mayr defined them as a sequential lineage, with each species evolving into the next (
chronospecies A chronospecies is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organ ...

chronospecies
). Though later Mayr changed his opinion on the australopithecines (recognising ''
Australopithecus ''Australopithecus'' (, ; ; singular: australopith) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genera ''Homo'' (which includes modern humans), ''Paranthropus'', and ''Kenyanthropus'' ev ...

Australopithecus
''), his more conservative view of
archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (dependin ...
diversity became widely adopted in the subsequent decades. Though ''H. erectus'' is still maintained as a highly variable, widespread and long-lasting species, it is still much debated whether or not sinking all Middle Pleistocene remains into it is justifiable. Mayr's lumping of ''H. heidelbergensis'' was first opposed by American anthropologist
Francis Clark Howell Francis Clark Howell (November 27, 1925 – March 10, 2007), generally known as F. Clark Howell, was an American anthropologist. Born in Kansas City, Missouri, F. Clark Howell grew up in Kansas, where he became interested in natural history. He ...
in 1960. In 1974, British physical anthropologist
Chris Stringer Christopher Brian Stringer (born 1947) is a British physical anthropologist noted for his work on human evolution. Biography Growing up in a working-class family in the East End of London The East End of London, often referred to withi ...

Chris Stringer
pointed out similarities between the Kabwe 1 and the Greek
Petralona Petralona ( el, Πετράλωνα, ) is a neighborhood of Athens , image_skyline = File:Athens Montage L.png, center, 275px, alt=Athens montage. Clicking on an image in the picture causes the browser to load the approp ...

Petralona
skulls to the skulls of modern humans (''H. sapiens'' or ''H. s. sapiens'') and Neanderthals (''H. neanderthalensis'' or ''H. s. neanderthalensis''). So, Stringer assigned them to ''
Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and large, complex brains. This has enabled the development of advanced tools, culture, and language. Humans are highl ...

Homo sapiens
sensu lato ''Sensu'' is a Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Roman ...
'' ("in the broad sense"), as ancestral to modern humans and Neanderthals. In 1979, Stringer and Finnish anthropologist
Björn KurténBjörn Olof Lennartson Kurtén (19 November 1924 – 28 December 1988) was a Finnish vertebrate paleontologist, belonging to the Swedish-speaking Finns, Swedish-speaking minority of his country. Early life and education Kurtén was born at Vaasa. ...
found that the Kabwe and Petralona skulls are associated with the
CromerianThe Cromerian Stage or Cromerian Complex, also called the Cromerian (german: Cromerium), is a stage in the Pleistocene glacial history of north-western Europe, mostly occurring more than half a million years ago. It is named after the East Anglian to ...
industry Industry may refer to: Economics * Industry (economics) In macroeconomics, an industry is a branch of an economy that produces a closely related set of raw materials, goods, or services. For example, one might refer to the wood industry ...
like the Mauer mandible, and thus postulated these three populations might be allied with each other. Though these fossils are poorly preserved and do not provide many comparable possible diagnostic traits (and likewise it was difficult at the time to properly define a unique species), they argued that at least these Middle Pleistocene specimens should be allocated to ''H. (s.?) heidelbergensis'' or "''H. (s.?) rhodesiensis''" (depending on, respectively, the inclusion or exclusion of the Mauer mandible) to formally recognise their similarity. Further work most influentially by Stringer, palaeoanthropologist
Ian Tattersall Ian Tattersall (born 1945) is a British-born American paleoanthropologist and a curator emeritus with the American Museum of Natural History in New York City, New York New York City (NYC), often simply called New York, is the List of Unit ...

Ian Tattersall
, and human evolutionary biologist Phillip Rightmire reported further differences between Middle Pleistocene Afro-European specimens and ''H. erectus'' ''
sensu stricto ''Sensu'' is a Latin Latin (, or , ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken in the area around Rome, known as Latium. Through the power of the Ro ...
'' ("in the strict sense", in this case specimens from East Asia). Consequently, Afro-European remains from 600 to 300 thousand years ago—most notably from Kabwe, Petralona,
BodoBodo may refer to: Ethnicity * Bodo people, an ethno-linguistic group mainly from Northwest Assam, India * Bodo-Kachari people, an umbrella group from Nepal, India and Bangladesh that includes the Bodo people Culture * Bodo culture, the cul ...

Bodo
and Arago—are often classified as ''H. heidelbergensis''. In 2010, American physical anthropologist Jeffrey H. Schwartz and Tattersall suggested classifying all Middle Pleistocene European as well as Asian specimens—namely from Dali and Jinniushan in China—as ''H. heidelbergensis''. This model is not as universally accepted. After the 2010 identification of the genetic code of some unique archaic human species in Siberia, termed "
Denisovan The Denisovans or Denisova hominins ) are an extinct species or subspecies of archaic human that ranged across Asia during the Lower Paleolithic, Lower and Middle Paleolithic. Denisovans are known from few remains, and, consequently, most of wha ...
s" pending diagnostic fossil finds, it is postulated that the Asian remains could represent that same species. Thus, Middle Pleistocene Asian specimens, such as Dali Man or the Indian Narmada Man, remain enigmatic. The palaeontology institute at
Heidelberg University } Heidelberg University, officially the Ruprecht Karl University of Heidelberg, (german: Ruprecht-Karls-Universität Heidelberg; la, Universitas Ruperto Carola Heidelbergensis) is a public In public relations Public relations (PR) is ...

Heidelberg University
, where the Mauer mandible has been kept since 1908, changed the label from ''H. e. heidelbergensis'' to ''H. heidelbergensis'' in 2015. In 1976 at Sima de los Huesos (SH) in the Sierra de Atapuerca, Spain, Spanish palaeontologists Emiliano Aguirre, José María Basabe and Trinidad Torres began to excavate archaic human remains. Their investigation of the site was prompted by the finding of several
bear Bears are carnivora Carnivora is an order of placental Placentalia is one of the three extant subdivisions of the class of animals Mammalia Mammals (from Latin Latin (, or , ) is a classical language belonging to the Ita ...

bear
remains ('' Ursus deningeri'') since the early 20th century by amateur cavers (which consequently destroyed some of the human remains in that section). By 1990, about 600 human remains were reported, and by 2004 the number had increased to roughly 4,000. These represent at least 28 individuals, of which possibly only one is a child, and the rest teenagers and young adults. The fossil assemblage is exceptionally complete, with whole corpses buried rapidly, with all bodily elements represented. In 1997, Spanish palaeoanthropologist assigned these to ''H. heidelbergensis'', but in 2014, he retracted this, stating that Neanderthal-like features present in the Mauer mandible are missing in the SH humans.


Classification

In palaeoanthropology, the
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocene Epoch withi ...
is often termed the "muddle in the middle" because the species-level classification of
archaic human A number of varieties of ''Homo ''Homo'' () is the that emerged in the (otherwise extinct) genus ' that encompasses the extant species ' (), plus several extinct species classified as either to or closely related to modern humans (dependin ...
remains from this time period has been heavily debated. The ancestors of modern humans (''Homo sapiens'' or ''H. s. sapiens'') and Neanderthals (''H. neanderthalensis'' or ''H. s. neanderthalensis'') diverged during this time period, and, by convention, ''H. heidelbergensis'' is typically considered the
last common ancestor In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Biochemistry, chemical processes, Molecular biology, molecular interactions, Physiology, physiological mechanis ...
(LCA). This would make ''H. heidelbergensis'' a member of a
chronospecies A chronospecies is a species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organ ...

chronospecies
. It is much debated if the name ''H. heidelbergensis'' can be extended to Middle Pleistocene humans across the Old World, or if it is better to restrict it to just Europe. In the latter case, Middle Pleistocene African remains can be split off into "''H. rhodesiensis''". In the latter view, "''H. rhodesiensis''" can either be seen as the direct ancestor of modern humans, or of "'' H. helmei''" which evolved into modern humans. In 2021, Canadian palaeoanthropologist Mirjana Roksandic and colleagues suggested renaming "''H. rhodesiensis''" as "'' H. bodoensis''", after the Bodo cranium, as the name ''rhodesiensis'' honours English diamond magnate
Cecil Rhodes Cecil John Rhodes (5 July 1853 – 26 March 1902) was a British mining magnate A magnate, from the late Latin ''magnas'', a great man, itself from Latin ''magnus'', "great", is a noble or a man in a high social position, by birth, wealth or ...
who disenfranchised the black population in southern Africa. Regarding the Middle Pleistocene European remains, some are more firmly placed on the Neanderthal line (namely SH, Pontnewyyd, Steinheim and
Swanscombe Swanscombe (/ˈswɒnzkəm/) is a town in the Borough of Dartford in Kent Kent is a Counties of England, county in South East England and one of the home counties. It borders Greater London to the north-west, Surrey to the west and East Susse ...
), whereas others seem to have few uniquely Neanderthal features (Arago,
CepranoCeprano ( Ciociaro: ) is a town and ''comune The (; plural: ) is a basic Administrative division, constituent entity of Italy, roughly equivalent to a township or municipality. Importance and function The provides many of the basic civil ...
, Vértesszőlős,
Bilzingsleben Bilzingsleben is a village and a former Municipalities in Germany, municipality in the Sömmerda (district), Sömmerda Districts of Germany, district of Thuringia, Germany. Since 1 January 2019, it is part of the municipality Kindelbrück. The vil ...
, and Aroeira). Because of this, it is suggested there were multiple lineages (or species) in this region and time period, but French palaeoanthropologist
Jean-Jacques Hublin Jean-Jacques Hublin (born 30 November 1953) is a French paleoanthropologist Paleoanthropology or paleo-anthropology is a branch of paleontology Paleontology, also spelled palaeontology or palæontology (), is the scientific study of life that ...
considers this an unjustified extrapolation as they may have simply been different but still interconnected populations of a single, highly variable species. In 2015, Marie Antoinette de Lumley suggested the less derived material can also be split off into their own species or a subspecies of ''H. erectus s. l.'' (for example, the Arago material as "''H. e. tautavelensis''"). In 2018, Mirjana Roksandic and colleagues revised the hypodigm of ''H. heidelbergensis'' to include only the specimens with no Neanderthal-derived traits (namely Mauer, , Ceprano,
HaZore'a HaZore'a ( he, הַזּוֹרֵעַ, ''lit.'' The Sower, named after the neighbouring Abu Zurayq, Tel Zariq, ) is a kibbutz in northern Israel established in 1936 by German Jews. It is the only kibbutz that was established by members of the moveme ...
and Nadaouiyeh Aïn Askar). There is no defined distinction between latest potential ''H. heidelbergensis'' material—specifically Steinheim and SH—and the earliest Neanderthal specimens— Biache, France; Ehringsdorf, Germany; or Saccopastore, Italy. The use of the Mauer mandible, an isolated jawbone, as the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism In biology Biology is the natural science that studies life and living organisms, including their anatomy, physical structure, Bi ...
for the species has been problematic as it does not present many diagnostic features, and in addition it is missing from several Middle Pleistocene specimens. Anthropologist William Straus said on this topic that, "While the skull is the creation of God, the jaw is the work of the devil." If the Mauer mandible is actually a member of a different species than the Kabwe skull and most other Afro-European Middle Pleistocene archaic humans, then "''H. rhodesiensis''" would take priority as the name of the LCA. In 2021, Canadian anthropologist Mirjana Roksandic and colleagues recommended the complete dissolution of ''H. heidelbergensis'' and "''H. rhodesiensis''". They classified all European ''H. heidelbergensis'' as ''H. neanderthalensis'', and synonymised ''H. rhodesiensis'' with a new species they named "'' H. bodoensis''" which includes all African specimens, and potentially some from the Levant and the Balkans which have no Neanderthal-derived traits (namely Ceprano, Mala Balanica, HaZore'a and Nadaouiyeh Aïn Askar). ''H. bodoensis'' is supposed to represent the immediate ancestor of modern humans, but does not include the LCA of modern humans and Neanderthals. They suggested the confusing morphology of the Middle Pleistocene was caused by periodic ''H. bodoensis'' migration events into Europe following population collapses after glacial cycles, interbreeding with surviving indigenous populations.


Evolution

''H. heidelbergensis'' is thought to have descended from African ''H. erectus''—sometimes classified as ''
Homo ergaster ''Homo ergaster'' is an extinct species In biology, a species is the basic unit of biological classification, classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the larges ...

Homo ergaster
''—during the first
early expansions of hominins out of Africa Several expansions of populations of archaic humans (genus ''Homo ''Homo'' () is the genus Genus (plural genera) is a taxonomic rank Taxonomy (general) is the practice and science of classification of things or concepts, including the pr ...
beginning roughly 2 million years ago. Those that dispersed across Europe and stayed in Africa evolved into ''H. heidelbergensis'' or speciated into ''H. heidelbergensis'' in Europe and "''H. rhodesiensis''" in Africa, and those that dispersed across East Asia evolved into ''H. erectus s. s.'' The exact derivation from an ancestor species is obfuscated by a long gap in the human fossil record near the end of the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, being the earliest (or lowest) division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently ...
.
Convenience link
In 2016, Antonio Profico and colleagues suggested that 875,000-year-old skull materials from the Gombore II site of the
Melka Kunture Melka Kunture (መልካ ቁንጥሬ) is a Paleolithic The Paleolithic or Palaeolithic or Palæolithic (), also called the Old Stone Age (from Greek palaios - old, lithos - stone), is a period in prehistory Prehistory, also known as p ...
Formation, Ethiopia, represent a transitional morph between ''H. ergaster'' and ''H. heidelbergensis'', and thus postulated that ''H. heidelbergensis'' originated in Africa instead of Europe. According to genetic analysis, the LCA of modern humans and Neanderthal split into a modern human line, and a Neanderthal/Denisovan line, and the latter later split into Neanderthal and Denisovans. According to
nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA Deoxyribonucleic acid (; DNA) is a molecule A scanning tunneling microscopy image of pentacene molecules, which consist of linear chains of five carbon rings. A mole ...
analysis, the 430,000-year-old SH humans are more closely related to Neanderthals than Denisovans (and that the Neanderthal/Denisovan, and thus the modern human/Neanderthal split, had already occurred), suggesting the modern human/Neanderthal LCA had existed long before many European specimens typically assigned to ''H. heidelbergensis'' did, such as the Arago and Petralona materials. In 1997, Spanish archaeologist , Arsuaga, and colleagues described the roughly million-year-old '' H. antecessor'' from Gran Dolina, Sierra de Atapuerca, and suggested supplanting this species in the place of ''H. heidelbergensis'' for the LCA between modern humans and Neanderthals, with ''H. heidelbergensis'' descending from it and being a strictly European species ancestral to only Neanderthals. They later recanted. In 2020, Danish geneticist Frido Welker and colleagues analysed ancient proteins collected from an ''H. antecessor'' tooth found that it was a member of a sister lineage to the LCA rather than being the LCA itself (that is, ''H. heidelbergensis'' did not derive from ''H. antecessor''). Human dispersal beyond seems to have been quite limited during the
Lower Palaeolithic 250px, Four views of an Acheulean handaxe The Lower Paleolithic (or Lower Palaeolithic) is the earliest subdivision of the Paleolithic The Paleolithic or Palaeolithic or Palæolithic (), also called the Old Stone Age, is a period in human pr ...

Lower Palaeolithic
, with evidence of short-lived dispersals northward beginning after a million years ago. Beginning 700,000 years ago, more permanent populations seem to have persisted across the line coinciding with the spread of
hand axe A hand axe (or handaxe or Acheulean hand axe) is a prehistoric Prehistory, also known as pre-literary history, is the period of human history Human history, or world history, is the narrative of Human, humanity's past. It is under ...
technology across Europe, possibly associated with the dispersal of ''H. heidelbergensis'' and behavioural shifts to cope with the cold climate. Such occupation becomes much more frequent after 500,000 years ago.


Anatomy


Skull

In comparison to Early Pleistocene ''H. erectus''/''ergaster'', Middle Pleistocene humans have a much more modern human-like face. The nasal opening is set completely vertically in the skull, and the anterior nasal sill can be crested or sometimes a prominent spine. The
incisive canals In the opening of the incisive foramen, the orifices of two lateral canals are visible; they are named the incisive canals or foramina of Nicolas Steno, Stensen. Through each of them ascends the terminal branch of the greater palatine artery while ...
(on the ) open near the teeth, and are orientated like those of more recent human species. The
frontal bone The frontal bone is a bone A bone is a rigid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesitata'', a species of geometer moth found in Nor ...

frontal bone
is broad, the
parietal bone The parietal bones () are two bones in the skull The skull is a bone A bone is a rigid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific function * ''Triphosa haesi ...

parietal bone
can be expanded, and the squamous part of temporal bone is high and arched, which could all be related to increasing brain size. The
sphenoid bone The sphenoid bone is an unpaired bone of the neurocranium. It is situated in the middle of the skull towards the front, in front of the basilar part of occipital bone, basilar part of the occipital bone. The sphenoid bone is one of the seven bones ...

sphenoid bone
features a spine extending downwards, and the
articular tubercle The articular tubercle (eminentia articularis) is a bony eminence on the temporal bone The temporal bones are situated at the sides and base of the skull, and lateral to the temporal lobes of the cerebral cortex. The temporal bones are overlaid by ...
on the underside of the skull can jut out prominently as the surface behind the jaw hinge is otherwise quite flat. In 2004, Rightmire estimated the brain volumes of ten Middle Pleistocene humans variously attributable to ''H. heidelbergensis''—from Kabwe, Bodo, Ndutu, Dali, Jinniushan, Petralona, Steinheim, Arago, and two from SH. This set gives an average volume of about 1,206 cc, ranging from 1,100 to 1,390 cc. He also averaged the brain volumes of 30 ''H. erectus''/''ergaster'' specimens, spanning nearly 1.5 million years from across East Asia and Africa, as 973 cc, and thus concluded a significant jump in brain size, though conceded brain size was extremely variable ranging from 727 to 1,231 cc depending on the time period, geographic region, and even between individuals within the same population (the last one probably due to notable sexual dimorphism with males much bigger than females). In comparison, for modern humans, brain size averages 1,270 cc for males and 1,130 cc for females; and for Neanderthals 1,600 cc for males and 1,300 cc for females. In 2009, palaeontologists Aurélien Mounier, François Marchal and Silvana Condemi published the first differential diagnosis of ''H. heidelbergensis'' using the Mauer mandible, as well as material from Tighennif, Algeria; SH, Spain; Arago, France; and Montmaurin, France. They listed the diagnostic traits as: a reduced chin, a notch in the
submental space The submental space is a fascial spaces of the head and neck, fascial space of the head and neck (sometimes also termed fascial spaces or tissue spaces). It is a potential space located between the mylohyoid muscle Anatomical terms of location#Super ...
(near the throat), parallel upper and lower boundaries of the mandible in side-view, several mental foramina (small holes for blood vessels) near the cheek teeth, a horizontal retromolar space (a gap behind the molars), a gutter between the molars and the
ramusRamus can refer to: * A branch (botany) * A portion of a bone (from Latin ''ramus'', "branch"), as in the Ramus of the mandible or Superior pubic ramus * A nerve ramus such as the Dorsal ramus of spinal nerve * A taxonomic rank ("branch" in English) ...
(which juts up to connect with the skull), an overall long jaw, a deep fossa (a depression) for the
masseter muscle In human anatomy, the masseter is one of the muscles of mastication There are four classical muscles of mastication. During mastication, three muscles of mastication (''musculi masticatorii'') are responsible for adduction of the jaw, and one ( ...

masseter muscle
(which closes the jaw), a small gonial angle (the angle between the body of the mandible and the ramus), an extensive planum alveolare (the distance from the frontmost tooth socket to the back of the jaw), a developed planum triangulare (near the jaw hinge), and a mylohyoid line originating at the level of the third molar.


Build

Trends in body size through the Middle Pleistocene are obscured due to a general lack of limb bones and non-skull (post-cranial) remains. Based on the lengths of various
long bone The long bones are those that are longer than they are wide. They are one of five types of bone A bone is a rigid tissue Tissue may refer to: Biology * Tissue (biology), an ensemble of similar cells that together carry out a specific funct ...

long bone
s, the SH humans averaged roughly for males and for females, with maximums of respectively and . The height of a female partial skeleton from Jinniushan is estimated to have been quite tall at roughly in life, much taller than the SH females. A tibia from Kabwe is typically estimated to have been , among the tallest Middle Pleistocene specimens, but it is possible this individual was either unusually large or had a much longer tibia to
femur The femur (; ), or thigh bone, is the proximal Standard anatomical terms of location deal unambiguously with the anatomy of animals, including humans. Terms used generally derive from Latin or Greek language, Greek roots and used to describe s ...

femur
ratio than expected. If these specimens are representative of their respective continents, they would suggest that above-medium to tall people were prevalent throughout the Middle Pleistocene Old World. If this is the case, then most all populations of any archaic human species would have generally averaged to in height.
Early modern human Early modern human (EMH) or anatomically modern human (AMH) are terms used to distinguish ''Homo sapiens Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedality, bipedalism and larg ...
s were notably taller, with the Skhul and Qafzeh remains averaging for males and for females, an average of , possibly to increase the energy-efficiency of long-distance travel with longer legs. A conspicuously massive proximal (upper half) femur recovered from Berg Aukas Mine, Namibia, about east of
Grootfontein Grootfontein ( en, large spring after the nearby hot springs) is a city of 23,793 inhabitants in the Otjozondjupa Region of central Namibia. It is one of the three towns in the Otavi Triangle, situated on the B8 road (Namibia), B8 national road t ...
was originally estimated to have been as much as in life, but the exorbitant size is now attributed to an intense activity level while maturing; the Berg Aukas individual was probably proportionally similar to Kabwe 1. The human
bauplan A body plan, ''Bauplan'' (German plural ''Baupläne''), or ground plan is a set of morphological features common to many members of a phylum In biology, a phylum (; plural The plural (sometimes list of glossing abbreviations, abbreviated ) ...

bauplan
(body plan) had evolved in ''H. ergaster'', and characterises all later ''Homo'' species, but among the more derived members there are 2 distinct morphs: a narrow-chested and gracile build like modern humans, and a broader-chested and robust build like Neanderthals. It was once assumed that the Neanderthal build was unique to Neanderthals based on the gracile ''H. ergaster'' partial skeleton KNM WT-15000 ("
Turkana Boy Turkana Boy, also called Nariokotome Boy, is the name given to fossil KNM-WT 15000,KNM-WT 15000: Kenya National Museum; West Turkana; item 15000 a nearly complete skeleton of a ''Homo ergaster'' (alternatively referred to as African ''Homo erec ...

Turkana Boy
"), but the discovery of some Middle Pleistocene skeletal elements (though generally fragmentary and few and far between) seems to suggest Middle Pleistocene humans overall featured a more Neanderthal morph. Thus, the modern human morph may be unique to modern humans, evolving quite recently. This is most clearly demonstrated in the exceptionally well-preserved SH assemblage. Based on skull robustness, it was assumed Middle Pleistocene humans featured a high degree of
sexual dimorphism Sexual dimorphism is the condition where the sex Sex is either of two divisions, typically male Male (♂) is the sex of an organism that produces the gamete known as sperm. A male gamete can fuse with a larger female gamete, or ovum, in ...
, but the SH humans demonstrate a modern humanlike level. The SH humans and other Middle Pleistocene ''Homo'' have a more basal pelvis and femur (more similar to earlier ''Homo'' than Neanderthals). The overall broad and elliptical pelvis is broader, taller and thicker (expanded anteroposteriorly) than those of Neanderthals or modern humans, and retains an anteriorly located acetabulocristal buttress (which supports the iliac crests during hip abduction), a well defined supraacetabular groove (between the hip socket and the ilium), and a thin and rectangular
superior pubic ramus In vertebrate Vertebrates () comprise all species of animal Animals (also called Metazoa) are multicellular eukaryotic organisms that form the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, c ...
(as opposed to the thick, stout one in modern humans). The foot of all archaic humans has a taller , making the ankle more flexible (specifically dorsiflexion and plantarflexion).


Pathology

On the left side of its face, an SH skull (Skull 5) presents the oldest-known case of
orbital cellulitis Orbital cellulitis is inflammation of human eye, eye tissues behind the orbital septum. It is most commonly caused by an acute spread of infection into the eye socket from either the adjacent Paranasal sinuses, sinuses or through the blood. It may ...

orbital cellulitis
(eye infection which developed from an
abscess An abscess is a collection of pus Pus is an exudate An exudate is a fluid emitted by an organism through pores or a wound, a process known as exuding or exudation. ''Exudate'' is derived from ''exude'', "to ooze", from the Latin Lat ...

abscess
in the mouth). This probably caused
sepsis Sepsis is a life-threatening condition that arises when the body's response to infection An infection is the invasion of an organism's body by , their multiplication, and the reaction of tissues to the infectious agents and the s they pr ...
, killing the individual. A male SH pelvis (Pelvis 1), based on joint degeneration, may have lived for more than 45 years, making him one of the oldest examples of this demographic in the human fossil record. The frequency of 45-plus individuals gradually increases with time, but has overall remained quite low throughout the Palaeolithic. He similarly had the age-related maladies lumbar
kyphosis Kyphosis is an abnormally excessive convex curvature of the spine as it occurs in the thoracic and sacral regions. Abnormal inward concave ''lordotic'' curving of the cervical and lumbar regions of the spine is called lordosis. It can result fr ...

kyphosis
(excessive curving of the
lumbar vertebrae The lumbar vertebrae are, in human anatomy, the five vertebrae In the vertebrate spinal column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic ...

lumbar vertebrae
of the lower back), L5–S1
spondylolisthesis Spondylolisthesis is the displacement of one spinal vertebra In the vertebrate spinal column The vertebral column, also known as the backbone or spine, is part of the axial skeleton. The vertebral column is the defining characteristic of a ...

spondylolisthesis
(misalignment of the last lumbar vertebra with the first
sacral vertebra The sacrum (plural: ''sacra'' or ''sacrums''), in human body, human anatomy, is a large, triangular bone at the base of the vertebral column, spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situ ...
), and Baastrup disease on L4 and 5 (enlargement of the spinous processes). These would have produced lower back pain, significantly limiting movement, and may be evidence of group care. An adolescent SH skull (Cranium 14) was diagnosed with lambdoid single suture craniosynostosis (immature closing of the left
lambdoid suture #REDIRECT Lambdoid suture#REDIRECT Lambdoid suture The lambdoid suture (or lambdoidal suture) is a dense, fibrous connective tissue Connective tissue is one of the four basic types of animal tissue (biology), tissue, along with epithelial tissue, ...

lambdoid suture
, leading to skull deformities as development continued). This is a rare condition, occurring in less than 6 out of every 200,000 individuals in modern humans. The individual died near the age of 5, suggesting it was not abandoned due its deformity as has been done in historical times, and received the same quality of care as any other child.
Enamel hypoplasia Enamel hypoplasia is a defect of the teeth A tooth (plural teeth) is a hard, calcification, calcified structure found in the jaws (or mouths) of many vertebrates and used to Mastication, break down food. Some animals, particularly carnivores, a ...
on the teeth is used to determine bouts of nutritional stress. At a rate of 40% for the SH humans, this is significantly higher than exhibited in the earlier South African
hominin The Hominini form a taxonomic tribe of the subfamily Homininae Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae The Hominidae (), whose members are known as great apes or hominids (), are a t ...
''
Paranthropus robustus ''Paranthropus robustus'' is a species of robustness (morphology), robust australopithecine from the Early Pleistocene, Early and possibly Middle Pleistocene, Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, mo ...
'' at
Swartkrans Swartkrans is a fossil-bearing cave designated as a South African National Heritage Site, located about from Johannesburg Johannesburg (, also ; ; Zulu language, Zulu and xh, eGoli ), informally known as Jozi, Joburg, or "The City of G ...
(30.6%) or
Sterkfontein Sterkfontein (Afrikaans Afrikaans (, ) is a West Germanic language spoken in South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. With over 60 million people, it ...

Sterkfontein
(12.1%). Nonetheless, Neanderthals suffered even higher rates and more intense bouts of hypoplasia, but it is unclear if this is because Neanderthals were less capable of exploiting natural resources, or because they lived in harsher environments. A peak at 3½ years of age may be correlated with weaning age. In Neanderthals this peak was at 4 years, and many modern hunter gatherers also wean at about 4 years of age.


Culture


Food

Middle Pleistocene communities in general seem to have eaten big game at a higher frequency than predecessors, with meat becoming an essential dietary component. Diet could overall be varied—for example the inhabitants of Terra Amata seem to have been mainly eating deer, but also elephants, boar, ibex, rhino and
aurochs The aurochs (''Bos primigenius'') ( or ), also known as urus or ure, is an extinct cattle species that was first described in 1827. With a shoulder height of up to in bulls and in cows, it was one of the largest herbivores in Holocene Europe ...

aurochs
. African sites typically commonly yield bovine and horse bones. Though carcasses may have simply been scavenged, some Afro-European sites show specific targeting of a single species, which more likely indicates active hunting; for example:
Olorgesailie Olorgesailie is a geological formation in East Africa containing a group of Lower Paleolithic Archeology, archaeological sites. It is on the floor of the Eastern Rift Valley in southern Kenya, southwest of Nairobi along the road to Lake Magadi ...
, Kenya, which has yielded over 50 to 60 individual baboons ('' Theropithecus oswaldi''); and Torralba and Ambrona in Spain which have an abundance of elephant bones (though also rhino and large hoofed mammals). The increase in meat subsistence could indicate the development of group hunting strategies in the Middle Pleistocene. For instance, at Torralba and Ambrona, the animals may have been run into swamplands before being killed, entailing encircling and driving by a large group of hunters in a coordinated and organised attack. Exploitation of aquatic environments is generally quite lacking, despite some sites being in close proximity to the ocean, lakes or rivers. Plants were probably also frequently consumed, including seasonally available ones, but the extent of their exploitation is unclear as they do not fossilise as well as animal bones. Assuming a diet heavy in lean meat, an individual would have needed a high
carbohydrate A carbohydrate () is a biomolecule , showing alpha helices, represented by ribbons. This poten was the first to have its suckture solved by X-ray crystallography by Max Perutz and Sir John Cowdery Kendrew in 1958, for which they received a ...
intake to prevent protein poisoning, such as by eating typically abundant underground storage organs, tree bark, berries, or nuts. The Schöningen site, Germany, has over 200 plants in the vicinity which are either edible raw or when cooked.


Art

Upper Palaeolithic modern humans are well known for having etched engravings seemingly with symbolic value. As of 2018, only 27 Middle and Lower Palaeolithic objects have been postulated to have symbolic etching, out of which some have been refuted as having been caused by natural or otherwise non-symbolic phenomena (such as the fossilisation or excavation processes). The Lower Palaeolithic ones are: three 380,000-year-old pebbles from Terra Amata; a 250,000-year-old pebble from Markkleeberg, Germany; 18 roughly 200,000-year-old pebbles from Grotte du Lazaret, Lazaret (near Terra Amata); a roughly 200,000-year-old lithic from Jardin Exotique de Monaco, Grotte de l'Observatoire, Monaco; a 370,000-year-old bone from
Bilzingsleben Bilzingsleben is a village and a former Municipalities in Germany, municipality in the Sömmerda (district), Sömmerda Districts of Germany, district of Thuringia, Germany. Since 1 January 2019, it is part of the municipality Kindelbrück. The vil ...
, Germany; and a 200- to 130-thousand-year-old pebble from Quinson, Baume Bonne, France. In the mid-19th century, French archaeologist Jacques Boucher de Crèvecœur de Perthes began excavation at St. Acheul, Amiens, France, (the area where the Acheulian was defined), and, in addition to hand axes, reported perforated sponge fossils (''Porosphaera globularis'') which he considered to have been decorative beads. This claim was completely ignored. In 1894, English archaeologist Worthington George Smith discovered 200 similar perforated fossils in Bedfordshire, England, and also speculated that their function was beads, though he made no reference to Boucher de Perthes' find possibly because he was unaware of it. In 2005, Robert Bednarik reexamined the material, and concluded that—because all the Bedfordshire ''P. globularis'' fossils are sub-spherical and range in diameter, despite this species having a highly variable shape—they were deliberately chosen. They appear to have been bored through completely or almost completely by some parasitic creature (i. e., through natural processes), and were then percussed on what would have been the more closed-off end to fully open the hole. He also found wear facets which he speculated were begotten from clacking against other beads when they were strung together and worn as a necklace. In 2009, Solange Rigaud, Francisco d'Errico and colleagues noticed that the modified areas are lighter in colour than the unmodifed, suggesting they were inflicted much more recently such as during excavation. They were also unconvinced that the fossils could be confidently associated with the Acheulian artefacts from the sites, and suggested that—as an alternative to archaic human activity—apparent size-selection could have been caused by either natural geological processes or 19th-century collectors favouring this specific form. Early modern humans and late Neanderthals (the latter especially after 60,000 years ago) made wide use of red ochre for presumably symbolic purposes as it produces a blood-like colour, though ochre can also have a functional medicinal application. Beyond these two species, ochre usage is recorded at Olduvai Gorge, Tanzania, where two red ochre lumps have been found; Ambrona where an ochre slab was trimmed down into a specific shape; and Terra Amata where 75 ochre pieces were heated to achieve a wide colour range from yellow to red-brown to red. These may exemplify early and isolated instances of colour preference and colour categorisation, and such practices may not have been normalised yet. In 2006, Eudald Carbonell and Marina Mosquera suggested the SH hominins were buried by people rather than being the victims of some catastrophic event such as a cave-in, because young children and infants are absent which would be unexpected if this were a single and complete family unit. The SH humans are conspicuously associated with only a single stone tool, a carefully crafted hand axe made of high-quality quartzite (rarely used in the region), and so Carbonell and Mosquera postulated this was purposefully and symbolically placed with the bodies as some kind of grave good. Supposed evidence of symbolic graves would not surface for another 300,000 years.


Technology


Stone tools

The Lower Palaeolithic (Early Stone Age) comprises the Oldowan which was replaced by the
Acheulian Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the type site of Saint-Acheul (Amiens), Saint-Acheul, is an archaeological industry of stone tool manufacture characterized by distinctive oval and pear-shaped "han ...
characterised by the production of mostly symmetrical
hand axe A hand axe (or handaxe or Acheulean hand axe) is a prehistoric Prehistory, also known as pre-literary history, is the period of human history Human history, or world history, is the narrative of Human, humanity's past. It is under ...
s. The Acheulian has a timespan of about a million years, and such technological stagnation has typically been ascribed to comparatively limited cognitive abilities which significantly reduced innovative capacity, such as a deficit in cognitive fluidity, working memory, or a social system compatible with apprenticeship. Nonetheless, the Acheulian does seem to subtly change over time, and is typically split up into Early Acheulian and Late Acheulian, the latter becoming especially popular after 600 to 500 thousand years ago. Late Acheulian technology never crossed over east of the Movius Line into East Asia, which is generally believed to be due to either some major deficit in cultural transmission (namely smaller population size in the East) or simply preservation bias as far fewer stone tool assemblages are found east of the line. The transition is indicated by the production of smaller, thinner, and more symmetrical hand axes (though thicker, less refined ones were still produced). At the 500,000-year-old Boxgrove Man, Boxgrove site in England—an exceptionally well-preserved site with abundance of tool remains—thinning may have been produced by striking the hand axe near-perpendicularly with a soft hammerstone, hammer, possible with the invention of prepared platforms for tool making. The Boxgrove knappers also left behind large lithic flakes leftover from making hand axes, possibly with the intention of recycling them into other tools later. Late Acheulian sites elsewhere pre-prepared lithic cores ("Large Flake Blanks," LFB) in a variety of ways before shaping them into tools, making prepared platforms unnecessary. LFB Acheulian spreads out of Africa into West and South Asia before a million years ago and is present in Southern Europe after 600,000 years ago, but northern Europe (and the Levant after 700,000 years ago) made use of soft hammers as they mainly made use of small, thick flint nodules. The first prepared platforms in Africa come from the 450,000-year-old Fauresmith (industry), Fauresmith industry, transitional between the Early Stone Age (
Acheulian Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the type site of Saint-Acheul (Amiens), Saint-Acheul, is an archaeological industry of stone tool manufacture characterized by distinctive oval and pear-shaped "han ...
) and the Middle Stone Age. With either method, knappers (tool makers) would have had to have produced some item indirectly related to creating the desired product (hierarchical organisation), which could represent a major cognitive development. Experiments with modern humans have shown that platform preparation cannot be learned through purely observational learning, unlike earlier techniques, and could be indicative of well developed teaching methods as well as self-regulated learning. At Boxgrove, the knappers used not only stone but also bone and antler to make hammers, and the use of such a wide range of raw materials could speak to advanced planning capabilities as stoneworking requires a much different skillset to work and gather materials for than boneworking. The Kapthurin Formation, Kenya, has yielded the oldest evidence of blade and bladelet technology, dating to 545 to 509 thousand years ago. This technology is rare even in the Middle Palaeolithic, and is typically associated with Upper Palaeolithic modern humans. It is unclear if this is part of a long blade-making tradition, or if blade technology was lost and reinvented several times by multiple different human species.


Fire and construction

Despite apparent pushes into colder climates, evidence of fire is scarce in the archaeological record until 400 to 300 thousand years ago. Though it is possible fire remnants simply degraded, long and overall undisturbed occupation sequences such as at Arago or Gran Dolina conspicuously lack convincing evidence of fire usage. This pattern could possibly indicate the invention of ignition technology or improved fire maintenance techniques at this time, and that fire was not an integral part of people's lives before then in Europe. In Africa, on the other hand, humans may have been able to frequently scavenge fire as early as 1.6 million years ago from natural wildfires, which occur much more often on Africa, thus possibly (more or less) regularly using fire. The oldest established continuous fire site beyond Africa is the 780,000-year-old Daughters of Jacob Bridge, Gesher Benot Ya'aqov, Israel. In Europe, evidence of constructed dwelling structures—classified as firm surface huts with solid foundations built in areas mostly sheltered from the weather—has been recorded since the Cromerian Interglacial, the earliest example a 700,000-year-old stone foundation from Přezletice, Czech Republic. This dwelling probably featured a vaulted roof made of thick branches or thin poles, supported by a foundation of big rocks and earth. Other such dwellings have been postulated to have existed during or following the Holstein Interglacial (which began 424,000 years ago) in Bilzingsleben, Germany; Terra Amata (archaeological site), Terra Amata, France; and Fermanville and Saint-Germain-des-Vaux in Normandy. These were probably occupied during the winter, and, averaging only in area, they were probably only used for sleeping in, while other activities (including firekeeping) seem to have been done outside. Less-permanent tent technology may have been present in Europe in the Lower Paleolithic.


Spears

The appearance of repeated fire usage—earliest in Europe from Beeches Pit, England, and Schöningen, Germany—roughly coincides with
hafting Hafting is a process by which an artifact, often bone A bone is a Stiffness, rigid tissue (anatomy), tissue that constitutes part of the vertebrate skeleton in animals. Bones protect the various organs of the body, produce red blood cell, red ...
technology (attaching stone points to spears) best exemplified by the Schöningen spears. These nine wooden spears and spear fragments—in addition to a lance, and a double-pointed stick—date to 300,000 years ago and were preserved along a lakeside. The spears vary from in diameter, and may have been long, overall similar to present day competitive javelins. The spears were made of soft spruce wood, except for spear 4 which was (also soft) pine wood. This contrasts with the Clacton Spear, Clacton spearhead from Clacton-on-Sea, England, perhaps roughly 100,000 years older, which was made of hard yew wood. The Schöningen spears may have had a range of up to , though would have been more effective short range within about , making them effective distance weapons either against prey or predators. Besides these two localities, the only other site which provides solid evidence of European spear technology is the 120,000-year-old Lehringen site, Germany, where a yew spear was apparently lodged in an elephant. In Africa, 500,000-year-old points from Kathu Archaeological Complex, Kathu Pan 1, South Africa, may have been hafted onto spears. Judging by indirect evidence, a horse scapula from the 500,000-year-old Boxgrove shows a puncture wound consistent with a spear wound. Evidence of hafting (in both Europe and Africa) becomes much more common after 300,000 years.


Language

The SH humans had a modern humanlike hyoid bone (which supports the tongue), and middle ear bones capable of finely distinguishing frequencies within the range of normal human speech. Judging by dental striations, they seem to have been predominantly right-handed, and handedness is related to the lateralisation of brain function, typically associated with language processing in modern humans. So, it is postulated that this population was speaking with some origin of language, early form of language. Nonetheless, these traits do not absolutely prove the existence of language and humanlike speech, and its presence so early in time despite such anatomical arguments has been primarily opposed by cognitive scientist Philip Lieberman.


See also

*Altamura man *Ceprano Man *Dmanisi skulls *Early European modern humans *''Homo rhodesiensis'' *''Homo antecessor'' *Swanscombe Heritage Park *Tautavel Man


References


External links

*
Homo heidelbergensis
' – The Smithsonian Institution's Human Origins Program
Homepage of Mauer 1 ClubUNESCO World Heritage Centre - Archaeological Site of AtapuercaHuman Timeline (Interactive)
– Smithsonian Institution, Smithsonian, National Museum of Natural History (August 2016). {{Authority control Homo heidelbergensis, Fossil taxa described in 1908