Homo heidelbergensis

Homo heidelbergensis is an extinct species or subspecies of archaic humans in the genus Homo, which radiated in the Middle Pleistocene from about 700,000 to 300,000 years ago.[note 1] The derivation of H. sapiens from H. rhodesiensis has often been proposed, but is obscured by a fossil gap from 400–260 kya.[note 2] The species was originally named Homo heidelbergensis due to the skeleton's first discovery near Heidelberg, Germany.[2]

Homo heidelbergensis Temporal range: 0.7–0.2 Ma PreꞒ Ꞓ O S D C P T J K Pg N ↓ Middle Pleistocene
Cranium 5 (skull with jawbone) from Sima de los Huesos (cast at Natural History Museum, London)
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Suborder:	Haplorhini
Infraorder:	Simiiformes
Family:	Hominidae
Subfamily:	Homininae
Tribe:	Hominini
Genus:	Homo
Species:	†H. heidelbergensis
Binomial name
†Homo heidelbergensis Schoetensack, 1908
Synonyms
† Homo rhodesiensis (Woodward, 1921)

The first discovery—a mandible—was made in 1907 by Otto Schoetensack.[2][3] The skulls of this species share features with both H. erectus and modern humans; its brain was nearly as large as that of modern humans.[4] The Sima de los Huesos cave at Atapuerca in northern Spain holds particularly rich layers of deposits where excavations were still in progress as of 2018.[5][6][7][8]

H. heidelbergensis was dispersed throughout Eastern and Southern Africa (Ethiopia, Namibia, Southern Africa) as well as Europe (England, France, Germany, Greece, Hungary, Italy, Portugal, Spain).[9] Its exact relation both to the earlier H. antecessor and H. ergaster, and to the later species Neanderthals, Denisovans, and modern humans is unclear.[10][11][12]

Modern humans have been proposed to derive from H. heidelbergensis via H. rhodesiensis, present in East and North Africa from around 400 kya.[13][14] The correct assignment of many fossils to a particular chronospecies is difficult and often differences in opinion ensue among paleoanthropologists due to the absence of universally accepted dividing lines (autapomorphies) between H. erectus, H. heidelbergensis, and Neanderthals.

It is uncertain whether H. heidelbergensis is ancestral to modern humans, as a fossil gap in Africa between 400–260 thousand years ago obscures the presumed derivation of H. sapiens from H. rhodesiensis.[note 2] Genetic analysis of the Sima de los Huesos fossils (Meyer et al. 2016) seems to suggest that H. heidelbergensis in its entirety should be included in the Neanderthal lineage, as "pre-Neanderthal" or "archaic Neanderthal" or "early Neanderthal", while the divergence time between the Neanderthal and modern lineages has been pushed back to before the emergence of H. heidelbergensis, to about 600,000 to 800,000 years ago, the approximate time of the disappearance of Homo antecessor.[15][16]

The delineation between early H. heidelbergensis and H. erectus is also unclear.[8][17][18]

Taxonomy

Research history

Mauer 1, the type specimen

The Ciampate del Diavolo near the extinct Roccamonfina volcano in Italy, fossilised hominid footprints dated to around 350,000 years ago and attributed to Homo heidelbergensis.

The type specimen, Mauer 1 (a jawbone), was discovered by worker Daniel Hartmann in Mauer, to the southeast of Heidelberg, Germany, and was described in 1907 by German anthropologist Otto Schoetensack. He noted a lack of a distinct chin, but conceded that it had clearly belonged to a human form due to the humanlike teeth.[19] More fossils were subsequently found in Steinheim an der Murr, Germany (the Steinheim skull); Arago, France (Tautavel Man); Petralona, Greece; and Ciampate del Diavolo, Italy.

In 1921, a skull, Kabwe 1, was discovered by Swiss miner Tom Zwiglaar in Kabwe, Zambia (Zambia at the time was called Northern Rhodesia), and was tentatively assigned to a new species, H. rhodesiensis, by English palaeontologist Arthur Smith Woodward.[20] Kabwe 1 dates to around 300,000 years ago.[21] In 1976, the 600,000 year old Bodo cranium was discovered in the Middle Awash, Ethiopia, but was classified into H. heidelbergensis as H. rhodesiensis was beginning to fall out of favour.[22][23] The 400,000 year old Tanzanian Ndutu cranium has been classified as an "archaic H. sapiens".[24] The South African Saldanha cranium, or Elandsfontein cranium, discovered in 1954 was assigned to H. heidelbergensis.[25]

Replica of the Kabwe 1 skull

The Chinese Dali Man and Maba Man could represent Asian H. heidelbergensis.

In 1992, the 350,000 year old Sima de los Huesos site in the Sierra de Atapuerca in northern Spain was discovered, which has since yielded 5,500 bones belonging to perhaps 32 individuals, about 90% of known H. heidelbergensis remains. These individuals are thought to represent the immediate ancestors to Neanderthals or are themselves early Neanderthals.[26] In 1994, Boxgrove Man was discovered near the English Channel in association with hundreds of hand axes dating to 524–478,000 years ago.[27]

Classification

H. heidelbergensis is generally considered to have been the direct ancestor to modern humans[2] (though African members are alternatively classified as H. rhodesiensis or archaic H. sapiens[28][29][24]), Neanderthals,[30][2] and Denisovans.[31] This would make H. heidelbergensis a chronospecies.[32] H. heidelbergensis has also been described as a potential polytypic species, with different populations possibly being sufficiently different enough to qualify as being different subspecies.[33] The transitional forms for both species is obscured by a large gap in the fossil record about 400–300,000 years ago.[1] Thus, the timing of the replacement of H. heidelbergensis by descendant species is generally a matter of convention.[30]

"Miguelón" from Sima de los Huesos

Similarly, the timing of the derivation of H. heidelbergensis from an ancestor species is also unclear.[34] The oldest identified H. heidelbergensis fossils date to about 600,000 years ago. Stone tools discovered in 2005 from Pakefield, Suffolk, indicate human activity in England 700,000 years ago, and at the time these tools were assumed to have been created by a transitional form between H. heidelbergensis and the older European human H. antecessor.[35][36][37][38] However, enamel proteome analysis suggests that H. antecessor is not ancestral to H. heidelbergensis, but closely related nonetheless.[39] H. heidelbergensis may instead derive from African or European H. erectus, which also existed contemporaneously.

Until the 1990s, H. heidelbergensis was commonly classified as a subspecies of H. erectus (H. e. heidelbergensis). With the discovery of more complete remains, H. heidelbergensis has since become accepted as a unique species.[40][41] The paleontology institute at Heidelberg University, where the type specimen has been kept since 1908, changed the label from the subspecies to the species classification in 2015.[42]

Morphology

Reconstruction of Kabwe 1 (left) and a Sima de los Huesos individual (right) by Élisabeth Daynès

The average brain capacity of 10 H. heidelbergensis specimens from across Africa and Eurasia was 1,206 cc (73.6 cu in).[43] Kabwe 1 has been estimated at 1,230 cm³ (75 cu in).[44] For comparison, the average brain capacity for contemporaneous H. erectus was 973 cc (59.4 cu in),[43] and modern humans 1,270 cc (78 cu in) for males and 1,130 cc (69 cu in) for females.[45]

Male H. heidelbergensis averaged about 175 cm (5 ft 9 in) tall and 62 kg (136 lb) in males, and 1.57 m (5 ft 2 in) and 51 kg (112 lb) in females.[46] An analysis of 27 limb bones Sima de los Huesos calculated an average size of about 170 cm (5 ft 7 in) in height, which is similar to the average for Neanderthals and pre-industrial modern humans.[47] According to South African palaeoanthropologist Lee Rogers Berger, tibiae and femora remains indicate that the average height 400–350,000 years ago reached 2.1 m (7 ft) during a grassland expansion which supported larger prey items.[48]

Behavior

Further information: Pleistocene human diet and Origin of language

One of hundreds of handaxes found at Boxgrove

A biface nicknamed Excalibur from Sima de los Huesos[49] is hypothesised to have been a grave good, which, if correct, would be the oldest evidence of funerary practices.[49][50][51]

The morphology of the outer and middle ear suggests they had an auditory sensitivity similar to modern humans. They were probably able to differentiate between many different sounds.[52] Dental wear analysis suggests they were as likely to be right-handed as modern people.[53]

An archaeological site in Schöningen, Germany contained eight exceptionally well-preserved roughly 400,000-year-old spears for hunting, and various other wooden tools. Five-hundred-thousand-year-old hafted stone points used for hunting are reported from Kathu Pan 1 in South Africa, tested by way of use-wear replication.[54] This find could mean that modern humans and Neanderthals inherited the stone-tipped spear, rather than developing the technology independently.[54][55][56]

No forms of art have been uncovered. Red ochre, a mineral that can be used to mix a red pigment which is useful as a paint, has been found at Terra Amata in France and Bečov in the Czech Republic, but the dating of these pigments to the Middle Pleistocene is contested.[57]

The Schöningen spears are eight wooden throwing spears, dated to before 300,000 years ago, discovered between 1994 and 1998 in the open-cast lignite mine, in Schöningen, county Helmstedt, Germany, together with thousands of animal bones. They are regarded as the first direct evidence for active hunting by H. heidelbergensis (pre-Neanderthals).[58][59][60]

Language

Steven Mithen[61] believes that H. heidelbergensis and its descendant H. neanderthalensis, acquired a pre-linguistic system of communication.

Homo heidelbergensis is thought to have been the first ancestor of modern humans not to have air sacs, which are laryngeal diverticula involved in vocalization. The loss of air sacs may have contributed to humans' ability to develop vocal language. Ancestors such as Australopithecus afarensis did have air sacs, as do other great apes.[62] Furthermore, there is evidence that Homo heidelbergensis was right-handed. Handedness is associated with the development of language among hominins.[63] Considering this evidence, scientists have hypothesized about the speaking capabilities of the species. A recent study that compared the speech frequency of humans and chimpanzees reported that H. heidelbergensis speech abilities more closely resemble those of modern-day humans. More specifically, "the Atapuerca SH hominins show[ed] a bandwidth that [wa]s slightly displaced and considerably extended to encompass the frequencies that contain relevant acoustic information in human speech."[64]

Pathology

The skull of the Homo heidelbergensis with odontogenic orbital cellulitis[65]

The earliest documented case of odontogenic orbital cellulitis, a severe eye infection that develops from an abscess in the mouth, occurred in H. heidelbergensis.[66][67]

See also

• Altamura man
• Homo cepranensis
• Homo rhodesiensis
• Human timeline
• List of fossil sites (with link directory)
• List of human evolution fossils
• Swanscombe Heritage Park

Notes

1. The fossil range spans about 0.6 to 0.4 Ma; cladistically, H. heidelbergensis is estimated to have developed from H. erectus (or H. antecessor) around 0.8–0.7 Ma, and given rise to H. neanderthalensis (and possibly H. sapiens) around 0.4–0.3 Ma.
2. "Most, if not all, of the African specimens assigned to H. rhodesiensis (cf heidelbergensis) seem to predate the divergence between H. neanderthalensis and H. sapiens [viz., assumed at 0.5 Mya prior to the revision by Meyer et al. 2016]. However, a gap in the fossil record, possibly between 400 and 260 ka, blurs the transition or punctuation event that separated H. rhodesiensis and H. sapiens."[1]

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67. Ascaso, F.; Adiego, M.I. (2016-09-14). "Homo heidelbergensis: the oldest case of odontogenic orbital cellulitis?". Acta Ophthalmologica. 94. doi:10.1111/j.1755-3768.2016.0022. ISSN 1755-375X.

Further reading

• Avery, D. Margaret. 2018. "Micromammals from the Type Site of Broken Hill Man (Homo Rhodesiensis) near Kabwe, Zambia: A Historical Note." Historical Biology 30 (1–2): 276–83. https://doi.org/10.1080/08912963.2017.1297434.[1]
• Back ache: it's been a pain for millions of years - University of Cambridge
• Friess, Martin. 2010. "Calvarial Shape Variation among Middle Pleistocene Hominins: An Application of Surface Scanning in Palaeoanthropology." Comptes Rendus Palevol, Imaging & 3D in palaeontology and palaeoanthropology, 9 (6): 435–43. https://doi.org/10.1016/j.crpv.2010.07.016.[2]
• Godinho, Ricardo Miguel, Laura C. Fitton, Viviana Toro-Ibacache, Chris B. Stringer, Rodrigo S. Lacruz, Timothy G. Bromage, and Paul O'Higgins. 2018. "The Biting Performance of Homo Sapiens and Homo Heidelbergensis." Journal of Human Evolution 118 (May): 56–71. https://doi.org/10.1016/j.jhevol.2018.02.010.[3]
• Hublin, Jean-Jacques, Abdelouahed Ben-Ncer, Shara E. Bailey, Sarah E. Freidline, Simon Neubauer, Matthew M. Skinner, Inga Bergmann, et al. 2017. "New Fossils from Jebel Irhoud, Morocco and the Pan-African Origin of Homo Sapiens." Nature 546 (7657): 289–92. https://doi.org/10.1038/nature22336.[4]
• Murrill, Rupert I. (1975). "A comparison of the Rhodesian and Petralona upper jaws in relation to other Pleistocene hominids". Zeitschrift für Morphologie und Anthropologie. 66 (2): 176–187. PMID 806185..
• Murrill, Rupert Ivan (1981). Ed. Charles C. Thomas (ed.). Petralona Man. A Descriptive and Comparative Study, with New Information on Rhodesian Man. Springfield, Illinois: Thomas. ISBN 978-0-398-04550-0.
• Perner, Josef, and Frank Esken. 2015. "Evolution of Human Cooperation in Homo Heidelbergensis: Teleology versus Mentalism." Developmental Review, Theories of development, 38 (December): 69–88. https://doi.org/10.1016/j.dr.2015.07.005.[5]
• Reich, David (2018). Who We Are And How We Got Here - Ancient DNA and the New Science of the Human Past. Pantheon Books. ISBN 978-1101870327.[6]
• Rice, Stanley (2006). Encyclopedia of Evolution. Facts on File, Inc.
• Sauer, A. (1985). Erläuterungen zur Geol. Karte 1 : 25 000 Baden-Württ. Stuttgart.
• Schoetensack, O. (1908). Der Unterkiefer des Homo heidelbergensis aus den Sanden von Mauer bei Heidelberg. Leipzig: Wilhelm Engelmann.
• Singer Robert R. and J. Wymer (1968). "Archaeological Investigation at the Saldanha Skull Site in South Africa". The South African Archaeological Bulletin. 23 (3): 63–73. doi:10.2307/3888485. JSTOR 3888485.
• Studies on the condition and structure of bone of the Saldanha fossil cranium
• Weinert, Hans (1937). "Dem Unterkiefer von Mauer zur 30jährigen Wiederkehr seiner Entdeckung". Zeitschrift für Morphologie und Anthropologie (in German). 37 (1): 102–13. JSTOR 25749563.
• Woodward, Arthur Smith (1921). "A New Cave Man from Rhodesia, South Africa". Nature. 108 (2716): 371–372. Bibcode:1921Natur.108..371W. doi:10.1038/108371a0.

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1. Avery, D. Margaret (March 2017). "Micromammals from the type site of Broken Hill Man (Homo rhodesiensis) near Kabwe, Zambia: a historical note". Historical Biology. 30 (1–2): 276–283. doi:10.1080/08912963.2017.1297434. ISSN 0891-2963.
2. Friess, Martin (2010-09-01). "Calvarial shape variation among Middle Pleistocene hominins: An application of surface scanning in palaeoanthropology". Comptes Rendus Palevol. 9 (6–7): 435–443. doi:10.1016/j.crpv.2010.07.016. ISSN 1631-0683.
3. Godinho, Ricardo Miguel; Fitton, Laura C.; Toro-Ibacache, Viviana; Stringer, Chris B.; Lacruz, Rodrigo S.; Bromage, Timothy G.; O'Higgins, Paul (2018-05-01). "The biting performance of Homo sapiens and Homo heidelbergensis" (PDF). Journal of Human Evolution. 118: 56–71. doi:10.1016/j.jhevol.2018.02.010. ISSN 0047-2484. PMID 29606203.
4. Hublin, Jean-Jacques; Ben-Ncer, Abdelouahed; Bailey, Shara E.; Freidline, Sarah E.; Neubauer, Simon; Skinner, Matthew M.; Bergmann, Inga; Le Cabec, Adeline; Benazzi, Stefano (2017-06-07). "New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens" (PDF). Nature. 546 (7657): 289–292. Bibcode:2017Natur.546..289H. doi:10.1038/nature22336. ISSN 0028-0836. PMID 28593953.
5. Perner, Josef; Esken, Frank (2015-12-01). "Evolution of human cooperation in Homo heidelbergensis: Teleology versus mentalism". Developmental Review. 38: 69–88. doi:10.1016/j.dr.2015.07.005. ISSN 0273-2297.
6. Diamond, Jared (April 20, 2018). "A Brand-New Version of Our Origin Story". The New York Times. Retrieved April 23, 2018.

External links

• Homo heidelbergensis – The Smithsonian Institution's Human Origins Program
• Theme park: The Grafenrain sand mine
• homepage of Mauer 1 Club
• UNESCO World Heritage Centre - Archaeological Site of Atapuerca
• Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).
• "Kabwe 1". The Smithsonian Institution's Human Origin Program. 2010-01-30. Retrieved 2 November 2010.
• Human Timeline (Interactive) – Smithsonian, National Museum of Natural History (August 2016).