Homo heidelbergensis

Classification Raciology On 21 October 1907, miners recovered a large human mandible (lower jaw) about down the Grafenrain sand pit near the village of Mauer — southeast of Heidelberg. German geologists and had earlier characterised the site as diluvial deposits (remnants of the Great Flood) dating to the Tertiary. Mauer 1 was the oldest European human fossil at the time. German anthropologist Otto Schoetensack made the first report of the skull in 1908, classifying it as a new human species, Homo heidelbergensis. After Neanderthals (H. neanderthalensis), it was the second-named fossil species in the genus Homo. (Qafzeh 9 pictured) were thought to directly descend from Mauer 1. Modern evolutionary synthesis By the middle of the century with the formulation of modern evolutionary synthesis, the common convention was to relegate all ancient human specimens into the genus Homo, and designate only a single species of Homo at any point in time: H. erectus which evolved into H. sapiens (anagenesis). The many defined species of archaic humans, including heidelbergensis, were generally lumped as subspecies of either H. erectus or H. sapiens. This left H. erectus and H. sapiens considerably polytypic and anatomically variable. In 1963, Russian-American geneticist Theodosius Dobzhansky noted that Middle Pleistocene Western and Eastern Eurasia — the former represented by H. e. heidelbergensis and the Tunisian H. e. mauritanicus; and the latter by the Chinese H. e. pekinensis and H. e. erectus — had significant anatomical and technological differences (Movius Line). He forwarded the possibility that these two groups represent distinct but contemporary species (cladogenesis), with H. erectus in the East, and the ancestors of H. sapiens in the West. grouped the Tighennif jaw ("H. e. mauritanicus", pictured) with Mauer 1 ("H. e. heidelbergensis"). Cladistics In 1974, British physical anthropologist Chris Stringer noted that the Greek Petralona 1 was anatomically more comparable to the Zambian Kabwe 1, Mauer 1, and Vértesszőlős 2 than to East Asian Middle Pleistocene H. erectus. He proposed classifying them as H. s. heidelbergensis — a widespread Euro-African clade, and the last common ancestor of modern humans (H. sapiens sapiens) and Neanderthals (H. sapiens neanderthalensis). At the time, he was hesitant to revive entire species for fear of recluttering human taxonomy, but in 1983, he proposed classifying them as a unique species as either H. heidelbergensis or H. rhodesiensis (named in 1921 with Kabwe 1), depending on the inclusion of Mauer 1, as the common ancestor of H. sapiens and H. neanderthalensis. The utility of H. heidelbergensis is complicated by its definition on a jaw, which is an uncommon find in Middle Pleistocene deposits, and additionally has few diagnostic traits. Nonetheless, Kabwe 1, Petralona 1, the Ethiopian Bodo cranium, and the French Arago have normally been discussed altogether as representatives of H. heidelbergensis, united most evidently by their brow ridge anatomy. Though H. heidelbergensis became a popular designation, in 2000, American anthropologists Sally McBrearty and Alison S. Brooks argued that H. heidelbergensis should be reserved for only the direct ancestors of Neanderthals in Europe. They recommended reviving H. rhodesiensis to house African Middle Pleistocene fossils they believed were directly ancestral to modern humans. Spanish palaeoanthropologist Juan Luis Arsuaga and colleagues made a similar opinion while studying the Spanish Sima de los Huesos hominins — which comprise the vast majority of the Middle Pleistocene human fossil record. They opted to classify every Middle Pleistocene European fossil as a Neanderthal ancestor under the name H. heidelbergensis, and placed the 1 million year old Spanish H. antecessor as the last common ancestor of modern humans and Neanderthals. Stringer disagreed with the inclusion of the Sima de los Huesos hominins in H. heidelbergensis, preferring to classify them as Neanderthals. In 2011, Arsuaga and colleagues failed to identify distinctly Neanderthal traits in Mauer 1 — unlike in the Sima de los Huesos hominins and some other Middle Pleistocene Europeans. They recognised two distinct groups occupying Middle Pleistocene Europe: one that was evolving into Neanderthals (pre-Neanderthals), and one that was not (maybe best designated as H. heidelbergensis). In Africa, Stringer noted that some of the specimens he assigned to H. heidelbergensis have similarities with modern humans in the face (such as the Tanzanian Ndutu cranium) while others do not (Kabwe 1 or Bodo). Congruently, in 2020, the Kabwe 1 skull was dated to roughly 300,000 years ago, living at the same time as the earliest recognised modern human fossils at the Moroccan Jebel Irhoud site. Late-surviving H. heidelbergensis populations may have interbred with modern humans. While some East Asian Middle Pleistocene fossils have some anatomical similarities to material typically classified as H. heidelbergensis, genetic sequencing of human fossils in the 2010s identified an enigmatic group of archaic humans called the Denisovans — closely related to Neanderthals — dispersed across East Asia. This opened the possibility that these East Asian specimens belong to a different, unique species (one that was also interbreeding with modern humans, as well as Neanderthals), In a 2024 interview with Cell Biology Magazine, Stringer expressed doubts with his earlier conceptions of H. heidelbergensis. ==Anatomy==

jawbones When Schoetensack described H. heidelbergensis in 1908 with the jaw Mauer 1, he distinguished it from any other human jaw known at the time by its thickened mandibular body, anteroposteriorly (front to back) widened ramus (where the jaw goes up to connect with the skull), and the lack of a chin. Kabwe 1, Petralona 1, Bodo, and Arago are normally presented altogether as representatives of H. heidelbergensis. The former three lack any jawbone material, but the Arago jawbones share with Mauer 1: • A wide mandibular symphysis (where the left and right halves of the jaw fuse) which arcs up between two tubercles (bony projections where tendons and ligaments attach to) and expands back into a thickened body of the mandible (which creates a horizontal sulcus, or groove, above); • A large and posterior mental foramen (hole for blood vessels); a low mandibular head (where the jaw hinge is) below the level of the coronoid process (which connects with the skull); • A rounded gonial region (where the mandibular body meets the ramus). All four skulls are united by their tall supraorbital tori (brow ridges) which have a generally flat front surface, twist at the superolateral margins (at the top by the edge of the face), and reach maximum height over the middle of the orbit (eye socket). They differ to some degree in the rest of the face. ==Culture==

Diet may have been commonly preyed upon by H. heidelbergensis. and rhinoceroses of the genus Stephanorhinus. 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, Kenya, which has yielded over 50 to 60 butchered baboons (Theropithecus oswaldi); and the Spanish Torralba and Ambrona sites which feature elephants' graveyards. Subsistence on large prey items could indicate group hunting strategies. For instance, at Torralba and Ambrona, the animals may have been encircled and run into swamplands by a coordinated and organised group of hunters before being killed. Some populations seem to have been extensively exploiting plant resources. At the 780,000 year old Gesher Benot Ya'aqov site, Israel, the inhabitants gathered and ate 55 different types of fruits, vegetables, seeds, nuts, and tubers. The inhabitants may have been using fire to roast certain plant materials that otherwise would have been inedible. They also consumed amphibians, reptiles, birds, aquatic and terrestrial invertebrates, in addition to the usual large creatures such as elephant and fallow deer. Technology Stone tools from the Boxgrove site, England The Lower Palaeolithic (Early Stone Age) comprises the Oldowan (a simple chopper and flake industry) which was replaced by the Acheulean, which is characterised by the production of mostly symmetrical hand axes. The Late Acheulean culture spread out across Europe and Africa by the beginning of the Middle Pleistocene, usually associated with the dispersal of H. heidelbergensis. This is distinguished from earlier Acheulean artefacts produced by H. erectus by the thinner and more symmetrical handaxes which bear more flaking scars. Some sites have much smaller handaxes which might fall under the African Middle Stone Age. The Late Acheulean reached Western Europe by the mid-Middle Pleistocene, but some sites — namely Arago — can feature predominantly choppers and flakes instead of handaxes. At the 500,000 year old English Boxgrove site, knappers may have been making prepared platforms for tool making. They were also using bone and antler as hammers. Late Acheulean sites elsewhere preprepared lithic cores ("Large Flake Blanks", LFB) in a variety of ways before shaping them into tools, making prepared platforms unnecessary. LFB Acheulean 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, possibly transitional between the Early Stone Age (Acheulean) and the Middle Stone Age. Some of the points may have been hafted onto spears. In Africa, the earliest evidence of this comes from the 500,000 year old Kathu Pan 1 site in South Africa. A horse scapula from the 500,000 year old Boxgrove site 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 ago. The Kapthurin Formation, Kenya, has yielded the oldest evidence of small blade and bladelet technology, dating to 509,000 to 545,000 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 300,000 to 400,000 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 in Africa, thus possibly (more or less) regularly using fire. The oldest established continuous fire site beyond Africa is at Gesher Benot Ya'aqov. 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, 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 Palaeolithic. Art Engravings tibia from Bilzingsleben, Germany 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). Colouring 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. Beads File:Porosphaera globularis necklace.webp|thumb|Hypothetical Porosphaera globularis necklace, probably inaccurate In 2009, French anthropologist Solange Rigaud and colleagues noticed that the modified areas are lighter in colour than the unmodified, suggesting they were inflicted much more recently such as during excavation. They were also unconvinced that the fossils could be confidently associated with the Acheulean 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. In 2023, Italian archaeologist Gabriele Luigi Francesco Berruti and colleagues demonstrated that the abrasion identified by Bednarik could have occurred naturally from dirt, and reaffirmed that they probably did not come from the same layer as the Acheulean artefacts. They found no evidence of any human modification, and rejected the interpretation of them as decorative beads. == See also ==