|Cranium 5 (skull with jawbone) from Sima de los Huesos (cast at Natural History Museum, London)|
† Homo rhodesiensis
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 2] known from fossils found in Southern Africa, East Africa and Europe. African H. heidelbergensis has several subspecies. The subspecies are Homo heidelbergensis heidelbergensis, Homo heidelbergensis daliensis, Homo rhodesiensis, and Homo heidelbergensis steinheimensi. The derivation of Homo sapiens from Homo rhodesiensis has often been proposed, but is obscured by a fossil gap from 400–260 kya.[note 3] The species was originally named Homo heidelbergensis due to the skeleton's first discovery near Heidelberg, Germany.
The first discovery—a mandible—was made in 1907 by Otto Schoetensack. The skulls of this species share features with both Homo erectus and the anatomically modern Homo sapiens; its brain was nearly as large as that of Homo sapiens. 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.
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). Its exact relation both to the earlier Homo antecessor and Homo ergaster, and to the later lineages of Neanderthals, Denisovans, and modern humans is unclear.
Homo sapiens has been proposed as derived from H. heidelbergensis via Homo rhodesiensis, present in East and North Africa from around 400,000 years ago. 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 Homo erectus, Homo heidelbergensis, Homo rhodesiensis and Neanderthals.
It is uncertain whether H. heidelbergensis is ancestral to Homo sapiens, as a fossil gap in Africa between 400,000 and 260,000 years ago obscures the presumed derivation of H. sapiens from H. rhodesiensis.[note 3] 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 disappearance of Homo antecessor.
Derivation and taxonomy
H. heidelbergensis is thought to be derived from Homo antecessor around 800,000 to 700,000 years ago. The oldest-known fossil classified as H. heidelbergensis dates to around 600,000 years ago, but the flint tools found in 2005 at Pakefield near Lowestoft in Suffolk with teeth from the water vole Mimomys savini, a key dating species, suggest human presence in England at 700,000 years ago, assumed to correspond to a transitional form between H. antecessor and H. heidelbergensis. Fifty prehistoric hominid footprints up to nearly one million years old were discovered in Happisburgh, England. They are likely members of Homo antecessor that lived from 1.2 million to 800,000 years ago.
In Europe, H. heidelbergensis is taken to have given rise to H. neanderthalensis at 240,000 years ago (a conventional date dictated by a fossil gap; late H. heidelbergensis in Europe prior to 240 kya is also called "pre-Neanderthal" or "ante-Neanderthal"). Homo sapiens most likely derived from H. rhodesiensis (African H. heidelbergensis) after around 300,000 years ago.
A morphological separation of a European and an African branch of H. heidelbergensis during the Wolstonian Stage and Ipswichian Stage, the last of the prolonged Quaternary glacial periods, has been argued[by whom?] based on the evidence of the Atapuerca skull in Spain and the Kabwe skull in modern-day Zambia.
Neither the derivation of H. heidelbergensis from H. erectus, nor the derivation of anatomically modern humans and Neanderthals from H. heidelbergensis, are clear-cut and are the object of debate. Both H. erectus and H. heidelbergensis are described as polytypic species, which went through a number of population bottlenecks and associated events.[note 4] In the summary of Hublin (2013), Middle Pleistocene humans in Eurasia underwent a succession of population bottlenecks due to glaciations. The "Western Eurasian clade" derived form H. rhodesiensis or H. heidelbergensis sensu lato (i.e. the Neanderthals) diverge at MIS 12 (480 kya) but coalesce as late as MIS 5 (130 kya), suggesting a division between Eurasian H. heidelbergensis and H. neanderthalensis before MIS 11 (424 kya). A fossil gap in Africa between 400 and 260 kya obscures the presumed derivation of H. sapiens from H. rhodesiensis.
Chris Stringer (2012) argues for Homo heidelbergensis as an independent chronospecies. A 2013 genetic study on the Sima de los Huesos fossils classified them as H. heidelbergensis or "early Neanderthal".[note 5]
For more than half a century, many experts were reluctant to accept Homo heidelbergensis as a separate taxon due to the rarity of specimens, which prevented sufficient informative morphological comparisons and the distinction of H. heidelbergensis from other known human species.[note 6] The species name "heidelbergensis" only experienced a renaissance with the many discoveries of Middle Pleistocene fossils since the 1990s.
The paleontology institute at Heidelberg University, where the type specimen has been kept since 1908, as late as 2010 still classified it as Homo erectus heidelbergensis, i.e. categorizing it as a Homo erectus subspecies. This was reportedly changed to Homo heidelbergensis, accepting the categorization as separate species, in 2015.
"Rhodesian Man" (Kabwe 1) is now mostly classified as Homo heidelbergensis, though other designations such as Homo sapiens arcaicus and Homo sapiens rhodesiensis have also been proposed. White et al. (2003) suggested Rhodesian Man as ancestral to Homo sapiens idaltu (Herto Man).
Homo heidelbergensis is intermediate between Homo erectus and Homo neanderthalensis, with a typical cranial volume of approximately 1,250 cm3 (76 cu in). "The anatomy [of H. heidelbergensis] is clearly more primitive than that of Neanderthal, but the harmoniously rounded dental arch and the complete row of teeth...already typically human."
In general, the findings show a continuation of evolutionary trends that are emerging from around the Lower into Middle Pleistocene. Along with changes in the robustness of cranial and dental features, a remarkable increase in brain size from H. erectus towards H. heidelbergensis is noticeable.
Male H. heidelbergensis averaged about 1.75 m (5 ft 9 in) tall and 62 kg (136 lb). Females averaged 1.57 m (5 ft 2 in) and 51 kg (112 lb). A reconstruction of 27 complete human limb bones found in Atapuerca (Burgos, Spain) has helped to determine the height of H. heidelbergensis compared with H. neanderthalensis; the conclusion was that these H. heidelbergensis averaged about 170 cm (5 ft 7in) in height and were only slightly taller than Neanderthals.
According to Lee R. Berger of the University of the Witwatersrand, tibia and femora remains indicate that populations of H. heidelbergensis between 350,000 and 400,000 years ago were routinely over 2.1 m (7 ft) tall. According to him, this was a short-lived experiment that lasted during a grassland expansion, which lead to very large ungulates and antelopes.
The "nature of our object" reveals itself "at first sight" since "a certain disproportion between the jaw and the teeth" is obvious: "The teeth are too small for the bone. The available space would allow for a far greater flexibility of development" and "It shows a combination of features, which has been previously found neither on a recent nor a fossil human mandible. Even the scholar should not be blamed if he would only reluctantly accept it as human: Entirely missing is the one feature, which is regarded as particularly human, namely an outer projection of the chin portion, yet this deficiency is found to be combined with extremely strange dimensions of the mandibular body. The actual proof that we are dealing with human parts here only lies within the nature of the dentition. The completely preserved teeth bear the stamp 'human' as evidence: The canines show no trace of a stronger expression in relation to the other groups of teeth. They suggest a moderate and harmonious co-evolution, as it is the case in recent humans."
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. Dental wear analysis suggests they were as likely to be right-handed as modern people. Steven Mithen believes that H. heidelbergensis, like its descendant H. neanderthalensis, acquired a pre-linguistic system of communication. No forms of art have been uncovered, although 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 excavations in the south of France.
An archeological 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. This find could mean that modern humans and Neanderthals inherited the stone-tipped spear, rather than developing the technology independently.
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 direct evidence for active hunting by H. heidelbergensis (pre-Neanderthals).
Mauer 1, the first fossil discovery of this species, was found on 21 October 1907, at Mauer, near Heidelberg, Germany. However, it was not until 1908 that the discovery gained public interest. It is a jaw in good condition except for the missing premolar teeth, which were eventually found near the jaw. Otto Schoetensack, from the University of Heidelberg, identified and named the fossil.
Boxgrove Man is the name associated with a lower tibia discovered in 1994 at the Boxgrove Quarry site, close to the English Channel. The fossil was found along with hundreds of hand axes, and has been dated between 478,000 and 524,000 years old. Several H. heidelbergensis teeth were found at the same site in subsequent seasons.
Beginning in 1992, a Spanish team has located more than 5,500 human bones dated to an age of at least 350,000 years in the Sima de los Huesos site in the Sierra de Atapuerca in northern Spain. The pit contains fossils of perhaps 32 individuals together with remains of Ursus deningeri and other carnivores and a biface nicknamed Excalibur. It is hypothesized that this Acheulean axe made of red quartzite was some kind of ritual offering for a funeral. If it is so, it would be the oldest evidence of known of funerary practices. Ninety percent of the known H. heidelbergensis remains have been obtained from this site. The fossil pit bones include:
- A complete cranium (skull 5), nicknamed Miguelón, and fragments of other crania, such as skull 4, nicknamed Agamemnón and skull 6, nicknamed Rui (from El Cid, a local hero).
- A complete pelvis (pelvis 1), nicknamed Elvis, in remembrance of Elvis Presley.
- Mandibles, teeth, and many postcranial bones (femurs, hand and foot bones, vertebrae, ribs, etc.)
Nearby sites contain the only known and controversial Homo antecessor fossils.
There is current debate among scholars whether the remains at Sima de los Huesos are those of H. heidelbergensis or early H. neanderthalensis. In 2015, the study of mitochondrial DNA samples from three caves Sima de los Huesos revealed that they are "distantly related to the mitochondrial DNA of Denisovans rather than to that of Neanderthals."
In 2016 nuclear DNA analysis determined the Sima hominins are Neanderthals and not Denisova hominins, and the divergence between Neanderthals, Denisovans and anatomically modern humans predates 430,000 years ago.
Recent studies have hypothesized that homo sapiens and Neanderthals separated from the Homo heidelbergensis branch. They also proposed that "[a]s there are potential H. heidelbergensis fossils from Asia, it is possible they could represent the ancestors of the Denisovans."
A number of morphologically-comparable fossil remains came to light in East Africa (Bodo, Ndutu, Eyasi, Ileret) and North Africa (Salé, Rabat, Dar-es-Soltane, Djbel Irhoud, Sidi Aberrahaman, Tighenif) during the 20th century. The Saldanha cranium, found in 1953 in South Africa was subject to at least three taxonomic revisions from 1955 to 1996.
Kabwe 1, also called the Broken Hill skull, was assigned by Arthur Smith Woodward in 1921 as the type specimen for Homo rhodesiensis; it is today mostly assigned to Homo heidelbergensis. It was found in a lead and zinc mine in Broken Hill, Northern Rhodesia (now Kabwe, Zambia) in 1921 by Tom Zwiglaar, a Swiss miner. In addition to the cranium, an upper jaw from another individual, a sacrum, a tibia, and two femur fragments were also found. The skull was dubbed "Rhodesian Man" at the time of the find, but is now commonly referred to as the Broken Hill skull or the Kabwe cranium. Cranial capacity of the Broken Hill skull has been estimated at 1,230 cm3 (75 cu in). Bada, & al., (1974) published the direct date of 110 ka for this specimen measured by aspartic acid racemization. The destruction of the paleoanthropological site has made layered dating impossible. The skull suggests an extremely robust individual with the comparatively largest brow-ridges of any known hominin. It was described as having a broad face similar to that of Homo neanderthalensis (i.e. large nasal bones and thick protruding brow ridges). Consequently, researchers came up with interpretations such as "African Neanderthal". However, with regard to the skull's extreme robustness, recent research has highlighted several intermediate features between modern Homo sapiens and Neanderthal. The skull has cavities in ten of the upper teeth and is considered one of the oldest known occurrences of cavities. Pitting indicates significant infection before death and implies that the cause of death may have been due to dental disease infection or possibly chronic ear infection. The skull is kept in the Natural History Museum, London. There is a replica in the Museum in Livingstone, Zambia.
The 600,000-year-old Bodo cranium was found in 1976 by members of an expedition led by Jon Kalb at Bodo D'ar in the Awash River valley of Ethiopia. The initial discovery—a lower face—was made by Alemayhew Asfaw and Charles Smart. Two weeks later, Paul Whitehead and Craig Wood found the upper portion of the face. The skull is 600,000 years old. Although the skull is most similar to those of Kabwe, Woodward's nomenclature was discontinued and its discoverers attributed it to H. heidelbergensis. It has features that represent a transition between Homo ergaster/erectus and Homo sapiens.
Another specimen, "the hominid from Lake Ndutu" in northern Tanzania, around 400,000 years old. In 1976 R. J. Clarke classified it as Homo erectus and it has generally been viewed as such since, although points of similarity to H. sapiens have also been recognized. After comparative studies with similar finds in Africa allocation to an African subspecies of H. sapiens seems most appropriate. An indirect cranial capacity estimate suggests 1,100 ml (39 imp fl oz; 37 US fl oz). Its supratoral sulcus morphology and the presence of protuberance as suggested by Philip Rightmire "give the Nudutu occiput an appearance which is also unlike that of Homo erectus", but Stinger (1986) pointed out that a thickened iliac pillar is typical for Homo erectus. In a 1989 publication Clarke concludes: "It is assigned to archaic Homo sapiens on the basis of its expanded parietal and occipital regions of the brain".
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. Furthermore, there is evidence that Homo heidelbergensis was right-handed. Handedness is associated with the development of language among hominins. Considering this evidence, scientist 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."
Homo heidelbergenis firsts
Several attributes of human species appear first in H. heidelbergensis. According to the Smithsonian National Museum of Natural History, it was the first species of the genus Homo to build permanent shelters. Furthermore, H. heidlebergensis was the first Homo with a body shape that enabled it to withstand colder temperatures, paving the way for its successors to adapt for even colder landscapes: H. heidlebergensis could conserve more body heat to endure harsher climates because its body was comparatively wide in relation to its height. The apparently earliest documented case of odontogenic orbital cellulitis, a severe eye infection that develops from an abscess in the mouth, occurred in H. heidelbergensis.
- The "Galilee skull", found in 1925/6 at Mugharet el-Zuttiyeh, now in Israel, has been described as "the most likely Heidelberg candidate from Western Asia".
- Petralona 1, discovered in Petralona cave, Greece, in 1960, dated to between roughly 350,000 and 150,000 years old. It has been classified as either H. heidelbergensis or H. neanderthalensis.
- Tautavel Man (Arago 21) is a human skull discovered on 22 July 1971 near the village of Tautavel in Pyrénées-Orientales, dated at 450,000 years old. The fossil was classified as Homo erectus tautavelensis, and as such would not belong to H. heidelbergensis but to a different lineage of H. erectus which occupied Europe at the same time as H. heidelbergensis.
- H. heidelbergensis likely speciated into H. sapiens and H. neanderthalensis before c. 250 ka, but late survival of H. heidelbergensis in Africa is suggested by the tentative dating of Kabwe 1, the type specimen of H. rhodesiensis, at 110 ka.
- 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, via H. rhodesiensis, possibly H. sapiens) around 0.4–0.3 Ma.
- "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."
- "the picture emerging is one of Homo erectus as a widespread, polytypic species, with groups persisting longer in some regions than in others. The pattern documented in China and especially in Java contrasts with that in the West, where Homo erectus seems to disappear from the record at a relatively early date".
- "The  fossils' identity suddenly became complicated when a study of the maternally inherited mitochondrial DNA (mtDNA) from one of the bones revealed that it did not resemble that of a Neanderthal. Instead, it more closely matched the mtDNA of a Denisovan...". "Indeed, the Sima de los Huesos specimens are early Neanderthals or related to early Neanderthals," after his team had scanned this DNA for markers found only in Neanderthals, Denisovans or modern humans, they found that the nuclear genomes of those specimens were significantly more similar to Neanderthals. "And that suggests the Neanderthal-Denisovan split happened before 430,000 years ago".
- Until the 1990s it was common to place these specimens either in H. erectus or into a broad category along with Neanderthals that was often called archaic H. sapiens.
- Manzi, Giorgio (2016-08-08). "Humans of the Middle Pleistocene: The controversial calvarium from Ceprano (Italy) and its significance for the origin and variability of Homo heidelbergensis". Quaternary International. 411: 254–261. doi:10.1016/j.quaint.2015.12.047. ISSN 1040-6182.
- Hublin, J.-J. (2013), "The Middle Pleistocene Record. On the Origin of Neandertals, Modern Humans and Others" in: R. David Begun (ed.), A Companion to Paleoanthropology, John Wiley, pp. 517-537 (summary 529–531). (p. 532).
- Buck, Laura T.; Stringer, Chris B. (2014-03-17). "Homo heidelbergensis". Current Biology. 24 (6): R214–R215. doi:10.1016/j.cub.2013.12.048. ISSN 0960-9822. PMID 24650901.
- Kjærgaard, Peter C. (2014-04-29). "Inventing Homo gardarensis: Prestige, Pressure, and Human Evolution in Interwar Scandinavia". Science in Context. 27 (2): 359–383. doi:10.1017/s0269889714000106. ISSN 0269-8897.
- "Homo heidelbergensis (600,000 to 100,000 years ago)- Species Description". WGBH Educational Foundation and Clear Blue Sky Productions, Inc. Retrieved November 29, 2015.
- "Archaeological Site of Atapuerca". UNESCO World Heritage Centre. Retrieved November 29, 2018.
- "Homo heidelbergensis". Natural History Museum, London. Retrieved 18 March 2013.
- "Homo heidelbergensis: Evolutionary Tree information". Smithsonian National Museum of Natural History. 2010-02-14. Retrieved 18 March 2013.
- Mounier, Aurélien; Marchal, François; Condemi, Silvana (2009). "Is Homo heidelbergensis a distinct species? New insight on the Mauer mandible". Journal of Human Evolution. 56 (3): 219–46. doi:10.1016/j.jhevol.2008.12.006. PMID 19249816.
- "Der Homo heidelbergensis - Vor etwa 700 000 Jahren taucht der Homo heidelbergensis in Afrika auf und wandert dann über eine noch unbekannte Route ebenfalls nach Europa aus..." Homo heidelbergensis von Mauer e.V. Retrieved November 29, 2015.
- Lenton, Tim; Watson, Andrew J. (20 January 2011). Revolutions that Made the Earth. Oxford University Press. p. 364. ISBN 978-0-19-958704-9. Retrieved 7 February 2018.
- Dusseldorp, Gerrit Leendert (2009). A View to a Kill: Investigating Middle Palaeolithic Subsistence Using an Optimal Foraging Perspective. Sidestone Press. p. 14. ISBN 978-90-8890-020-4. Retrieved 7 February 2018.
- Minelli, Alessandro; Contrafatto, Giancarlo (10 November 2009). BIOLOGICAL SCIENCE FUNDAMENTALS AND SYSTEMATICS - Volume IV. EOLSS Publications. p. 248. ISBN 978-1-84826-189-1. Retrieved 7 February 2018.
- "Prehistoric World Hominid Chronology by Peter Kessler Homo neanderthalis". Kessler Associates. July 26, 2005. Retrieved December 9, 2015.
- "What was Homo heidelbergensis? - DNA studies on both species indicate that the two were certainly distinct from each other, although related through their common Homo heidelbergensis ancestors". InnovateUs Inc. Retrieved November 29, 2015.
- Matthias Meyer, Juan-Luis Arsuaga, Cesare de Filippo, Sarah Nagel, Ayinuer Aximu-Petri, Birgit Nickel, Ignacio Martínez, Ana Gracia, José María Bermúdez de Castro, Eudald Carbonell, Bence Viola, Janet Kelso, Kay Prüfer & Svante Pääbo, "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins", Nature 531, pages 504–507 (24 March 2016), doi:10.1038/nature17405
- Ewen Callaway, "Oldest ancient-human DNA details dawn of Neanderthals" Sequence of 430,000-year-old DNA pushes back divergence of humans and Neanderthals", Nature News, 14 March 2016.
- "Homo heidelbergensis - The evolutionary dividing line between Homo erectus and modern humans was not sharp". Dennis O'Neil. Retrieved November 29, 2015.
- Lieberman, Daniel E.; McBratney, Brandeis M.; Krovitz, Gail (2002). "The evolution and development of cranial form" (PDF). Proceedings of the National Academy of Sciences. 99 (3): 1134–1139. Bibcode:2002PNAS...99.1134L. doi:10.1073/pnas.022440799. PMC 122156. PMID 11805284. Retrieved December 9, 2015.
- Parfitt, Simon A.; Barendregt, René W.; Breda, Marzia; Candy, Ian; Collins, Matthew J.; Coope, G. Russell; Durbidge, Paul; Field, Mike H.; Lee, Jonathan R. (2005). "The earliest record of human activity in northern Europe". Nature. 438 (7070): 1008–12. Bibcode:2005Natur.438.1008P. doi:10.1038/nature04227. PMID 16355223.
- Roebroeks, Wil (2005). "Archaeology: Life on the Costa del Cromer". Nature. 438 (7070): 921–2. Bibcode:2005Natur.438..921R. doi:10.1038/438921a. PMID 16355198.
- Parfitt, Simon; Stuart, Tony; Stringer, Chris; Preece, Richard (January–February 2006). "700,000 years old: found in Suffolk". British Archaeology. 86. Archived from the original on 2013-09-28.
- Good, Clare; Plouviez, Jude (2007). The Archaeology of the Suffolk Coast (PDF). Suffolk County Council Archaeological Service.[page needed]
- Kinver, Mark (14 December 2005). "Tools unlock secrets of early man". BBC News. Retrieved November 16, 2012.
- Ashton N, Lewis SG, De Groote I, Duffy SM, Bates M, Bates R, et al. (2014) Hominin Footprints from Early Pleistocene Deposits at Happisburgh, UK.PLoS ONE 9(2): e88329. https://doi.org/10.1371/journal.pone.0088329
- D. Dean; J.-J. Hublin; R. Holloway; R. Ziegler (1998). "On the phylogenetic position of the pre-Neandertal specimen from Reilingen, Germany". Journal of Human Evolution. 34 (5). pp. 485–508. doi:10.1006/jhev.1998.0214.
- "Homo heidelbergensis - Homo heidelbergensis began to develop regional differences that eventually gave rise to two species of humans". Australian Museum. Retrieved October 29, 2015.
- Rightmire, G Philip (1998). "Human Evolution in the Middle Pleistocene: The Role of Homo heidelbergensis" (PDF). Evolutionary Anthropology. doi:10.1002/(SICI)1520-6505(1998)6:6<218::AID-EVAN4>3.0.CO;2-6.
- Stringer, Chris (2012). "Comment: What makes a modern human". Nature. 485 (7396): 33–35 . Bibcode:2012Natur.485...33S. doi:10.1038/485033a. PMID 22552077.
- "DNA from Neandertal relative may shake up human family tree". American Association for the Advancement of Science. September 11, 2015. Retrieved November 29, 2015.
- "Researchers Sequenced 430,000-Year-Old DNA From Neanderthal Relative". IFLScience. September 13, 2015. Retrieved November 29, 2015.
- McHenry, Henry. "Homo heidelbergensis". Encyclopaedia Britannica. Retrieved 2017-11-10.
- Mounier, M Aurélien (October 28, 2009). "Homo heidelbergensis is supported as a valid taxon. - Validité du taxon Homo heidelbergensis Schoetensack, 1908" (PDF). Université de la Méditerranée - Faculté de Médicine de Marseille. Retrieved 2017-11-10.
- "Die Evolution des Menschen - Homo heidelbergensis" (in German). evolution-mensch.de. Retrieved 2017-11-10.
- "Hierzu zählte noch im Jahr 2010 auch das Geologisch-Paläontologische Institut der Universität Heidelberg, das den Unterkiefer seit 1908 verwahrt und ihn als Homo erectus heidelbergensis auswies. Inzwischen wird er jedoch auch in Heidelberg als Homo heidelbergensis bezeichnet, siehe" (in German). Sammlung des Instituts für Geowissenschaften. Retrieved November 29, 2015.
- H. James Birx (10 June 2010). 21st Century Anthropology: A Reference Handbook. SAGE Publications. p. 48. ISBN 978-1-4522-6630-5.
- Bernard Wood (31 March 2011). Wiley-Blackwell Encyclopedia of Human Evolution, 2 Volume Set. John Wiley & Sons. pp. 761–762. ISBN 978-1-4443-4247-5.
- White, Tim D.; Asfaw, B.; DeGusta, D.; Gilbert, H.; Richards, G. D.; Suwa, G.; Howell, F. C. (2003). "Pleistocene Homo sapiens from Middle Awash, Ethiopia". Nature. 423 (6491): 742–747. Bibcode:2003Natur.423..742W. doi:10.1038/nature01669. PMID 12802332
- Asfaw, Berhane (2005). "A new hominid parietal from Bodo, middle Awash Valley, Ethiopia". American Journal of Physical Anthropology. 61 (3): 367–371. doi:10.1002/ajpa.1330610311. PMID 6412559.
- Dorey, Fran (2017-09-25). "Homo heidelbergensis - Key physical features". Australian Museum. Retrieved 2017-11-10.
- Johanna Kontny u. a.: Reisetagebuch eines Fossils. In: Günther A. Wagner u. a., S. 44.
- "Die Evolution des Menschen - Homo heidelbergensis - Die Tatsache, dass es keine klaren Übergänge zu geben scheint, macht es schwierig, eine Liste eindeutiger Merkmale des Homo heidelbergensis aufzustellen..." evolution-mensch de. Retrieved November 29, 2015.
- "Evolution of Modern Humans: Homo heidelbergensis". Behavioral Sciences Department, Palomar College. Retrieved 2012-12-01.
- "Homo heidelbergensis was only slightly taller than the Neanderthal". American Association for the Advancement of Science (AAAS). June 6, 2012. Retrieved November 29, 2015.
- Carretero, José-Miguel; Rodríguez, Laura; García-González, Rebeca; Arsuaga, Juan-Luis; Gómez-Olivencia, Asier; Lorenzo, Carlos; Bonmatí, Alejandro; Gracia, Ana; Martínez, Ignacio (2012). "Stature estimation from complete long bones in the Middle Pleistocene humans from the Sima de los Huesos, Sierra de Atapuerca (Spain)" (PDF). Journal of Human Evolution. 62 (2): 242–55. doi:10.1016/j.jhevol.2011.11.004. PMID 22196156. Lay summary – ScienceDaily (June 6, 2012).
- Burger, Lee (November 2007). "Our Story: Human Ancestor Fossils". The Naked Scientists.
- "Homo heidelbergensis essay". Institute of Human Origins. Retrieved October 29, 2015.
- "Scientists Determine Height of Homo Heidelbergensis". Sci-News. June 6, 2012. Retrieved November 29, 2015.
- "100 years of Homo heidelbergensis – life and times of a controversial taxon" (PDF). Max Planck Institute for Evolutionary Anthropology. Archived from the original (PDF) on March 4, 2016. Retrieved December 9, 2015.
- The Mystery of the Pit of Bones, Atapuerca, Spain: Species Homo heidelbergensis. Smithsonian Institution. Retrieved December 15, 2011.
- "Homo heidelbergensis - Eudald Carboneli et al. reported in the March 27, 2008 issue of Nature that a human jaw with a tooth dating 1.2-1.1 million years ago has been found in Sima del Elefante cave in the Atapuerca Mountains of Northern Spain". palomar.edu. Retrieved October 29, 2015.
- Martínez I, Rosa M, Arsuaga JL, Jarabo P, Quam R, Lorenzo C, Gracia A, Carretero JM, Bermúdez de Castro JM (2004). "Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain". Proceedings of the National Academy of Sciences. 101 (27): 9976–81. Bibcode:2004PNAS..101.9976M. doi:10.1073/pnas.0403595101. JSTOR 3372572. PMC 454200. PMID 15213327.
- Lozano, Marina; Mosquera, Marina; De Castro, José María Bermúdez; Arsuaga, Juan Luis; Carbonell, Eudald (2009). "Right handedness of Homo heidelbergensis from Sima de los Huesos (Atapuerca, Spain) 500,000 years ago". Evolution and Human Behavior. 30 (5): 369–76. doi:10.1016/j.evolhumbehav.2009.03.001.
- Mithen, Steven (2006). The Singing Neanderthals, ISBN 978-0-674-02192-1[page needed]
- Wilkins, Jayne; Schoville, Benjamin J.; Brown, Kyle S.; Chazan, Michael (2012). "Evidence for Early Hafted Hunting Technology" (PDF). Science. 338 (6109): 942–6. Bibcode:2012Sci...338..942W. doi:10.1126/science.1227608. PMID 23161998. Lay summary – The Guardian (November 15, 2012).
- "Homo heidelbergensis Oldest Wooden Spear". Smithsonian Institution. Retrieved October 29, 2015.
- Hartmut Thieme, Reinhard Maier (Hrsg.): Archäologische Ausgrabungen im Braunkohlentagebau Schöningen. Landkreis Helmstedt, Hannover 1995. Hartmut Thieme: Die ältesten Speere der Welt – Fundplätze der frühen Altsteinzeit im Tagebau Schöningen. In: Archäologisches Nachrichtenblatt 10, 2005, S. 409-417.
- Michael Baales, Olaf Jöris: Zur Altersstellung der Schöninger Speere. In: J. Burdukiewicz u. a. (Hrsg.): Erkenntnisjäger. Kultur und Umwelt des frühen Menschen. Veröffentlichungen des Landesamtes für Archäologie Sachsen-Anhalt 57, 2003 (Festschrift Dietrich Mania), S. 281-288.
- Thieme H. 2007. Der große Wurf von Schöningen: Das neue Bild zur Kultur des frühen Menschen in: Thieme H. (ed.) 2007: Die Schöninger Speere – Mensch und Jagd vor 400 000 Jahren. S. 224-228 Konrad Theiss Verlag, Stuttgart ISBN 3-89646-040-4
- Otto Schoetensack: Der Unterkiefer des Homo heidelbergensis aus den Sanden von Mauer bei Heidelberg. Ein Beitrag zur Paläontologie des Menschen. Leipzig, 1908, Verlag von Wilhelm Engelmann
- Streeter et al. 2001, Margret (2001). "Histomorphometric age assessment of the Boxgrove 1 tibial diaphysis". Journal of Human Evolution. 40 (4): 331–338. doi:10.1006/jhev.2001.0460. PMID 11312585. Retrieved 22 March 2015.
- "BBC - Science & Nature - The evolution of man".
- "Sorry, that's a dead link - Natural History Museum". Archived from the original on 2013-12-09. Retrieved 2013-08-23.
- "Nuclear DNA sequences from the hominin remains of Sima de los Huesos, Atapuerca, Spain // 5TH ANNUAL MEETING OF THE European Society for the study of Human Evolution, 10 – 12 SEPTEMBER 2015 LONDON/UK: Nuclear DNA sequences from the hominin" (PDF).
- magazine, Ewen Callaway, Nature. "Oldest Ancient-Human DNA Details Dawn of Neandertals". Scientific American. Retrieved 2016-03-14.
- Meyer, Matthias; Arsuaga, Juan-Luis; de Filippo, Cesare; Nagel, Sarah; Aximu-Petri, Ayinuer; Nickel, Birgit; Martínez, Ignacio; Gracia, Ana; de Castro, José María Bermúdez (2016-03-14). "Nuclear DNA sequences from the Middle Pleistocene Sima de los Huesos hominins". Nature. 531 (7595): 504–507. Bibcode:2016Natur.531..504M. doi:10.1038/nature17405. ISSN 1476-4687. PMID 26976447.
- "The evolution and development of cranial form in Homo" (PDF). Department of Anthropology, Harvard University. Retrieved December 9, 2015.
- Wood, Bernard (2011-03-31). Wiley-Blackwell Encyclopedia of Human Evolution, 2 Volume Set. ISBN 9781444342475. Retrieved December 9, 2015.
- "Kabwe 1". The Smithsonian Institution's Human Origin Program. 2010-01-30. Retrieved 2 November 2010.
- Begun, David R., ed. (2012). "The African Origin of Homo sapiens". A Companion to Paleoanthropology. John Wiley & Sons. ISBN 9781118332375.
- Rightmire, G Philip (1993). The Evolution of Homo erectus: Comparative Anatomical Studies of an Extinct Human Species. Cambridge University Press. ISBN 0-521-44998-7.
- Bada, Jeffrey L., Roy A. Schroeder, Reiner Protsch, and Rainer Berger. Concordance of Collagen-Based Radiocarbon and Aspartic-Acid Racemization Ages PNAS abstract URL.
- Bada, J. L. (1985). "Amino Acid Racemization Dating of Fossil Bones". Annual Review of Earth and Planetary Sciences. 13: 241–268. doi:10.1146/annurev.ea.13.050185.001325.
- "Collections - Natural History Museum". www.nhm.ac.uk.
- "Bodo – Paleoanthropology site information". Fossilized org. Retrieved December 9, 2015.
- "Bodo Skull and Jaw". Skulls Unlimited. Archived from the original on 8 December 2015. Retrieved 15 October 2012.
- "Bodo". Humanorigins.si.edu. 2010-01-30. Retrieved 15 October 2012.
- "Bodo fossil". Britannica Encyclopedia. Retrieved December 9, 2015.
- "Meet Bodo and Herto There is some discussion around the species assigned to Bodo". Nutcracker Man. April 7, 2015. Retrieved December 9, 2015.
- Rightmire, G. Philip (2005). "The Lake Ndutu cranium and early Homo sapiens in Africa". American Journal of Physical Anthropology. 61 (2): 245–254. doi:10.1002/ajpa.1330610214. PMID 6410925.
- "The Ndutu cranium and the origin of Homo sapiens – R. J. Clarke" (PDF). American Museum of Natural History. November 27, 1989. Retrieved December 9, 2015.
- Singer, R (September 1954). "The saldanha skull from hopefield, South Africa". American Journal of Physical Anthropology. 12 (3): 345–362. doi:10.1002/ajpa.1330120309. PMID 13207329.
- De Boer, Bart (2012-01-01). "Loss of air sacs improved hominin speech abilities". Journal of Human Evolution. 62 (1): 1–6. doi:10.1016/j.jhevol.2011.07.007. ISSN 0047-2484. PMID 22078314.
- Lozano, Marina; Mosquera, Marina; De Castro, José María Bermúdez; Arsuaga, Juan Luis; Carbonell, Eudald (2009-09-01). "Right handedness of Homo heidelbergensis from Sima de los Huesos (Atapuerca, Spain) 500,000 years ago". Evolution and Human Behavior. 30 (5): 369–376. doi:10.1016/j.evolhumbehav.2009.03.001. ISSN 1090-5138.
- Martínez, I.; Rosa, M.; Quam, R.; Jarabo, P.; Lorenzo, C.; Bonmatí, A.; Gómez-Olivencia, A.; Gracia, A.; Arsuaga, J.L. (2013-05-08). "Communicative capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain". Quaternary International. 295: 94–101. doi:10.1016/j.quaint.2012.07.001. ISSN 1040-6182.
- Gracia-Téllez, Ana; Arsuaga, Juan-Luis; Martínez, Ignacio; Martín-Francés, Laura; Martinón-Torres, María; Bermúdez De Castro, José-María; Bonmatí, Alejandro; Lira, Jaime (2013-05-08). "Orofacial pathology in Homo heidelbergensis: The case of Skull 5 from the Sima de los Huesos site (Atapuerca, Spain)". Quaternary International. 295: 83–93. doi:10.1016/j.quaint.2012.02.005. ISSN 1040-6182.
- "Homo heidelbergensis". The Smithsonian Institution's Human Origins Program. 2010-02-14. Retrieved 2018-11-17.
- Wroe, Stephen; Parr, William C. H.; Ledogar, Justin A.; Bourke, Jason; Evans, Samuel P.; Fiorenza, Luca; Benazzi, Stefano; Hublin, Jean-Jacques; Stringer, Chris (2018-04-11). "Computer simulations show that Neanderthal facial morphology represents adaptation to cold and high energy demands, but not heavy biting". Proc. R. Soc. B. 285 (1876): 20180085. doi:10.1098/rspb.2018.0085. ISSN 0962-8452. PMC 5904316. PMID 29618551.
- DeCroos, FC; Liao, JC; Ramey, NA; Li, I (2011-08-15). "Management of Odontogenic Orbital Cellulitis". Journal of Medicine and Life. 4 (3): 314–317. ISSN 1844-122X. PMC 3168817. PMID 22567060.
- 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.
- Cartmill, Matt; Smith, Fred H. (2009). The Human Lineage. John Wiley & Sons. ISBN 978-0471214915. Retrieved 2013-03-01.
- "Petralona 1". Smithsonian Institution. Retrieved November 29, 2015.
- "Tautavel Man". Saissac. Retrieved November 29, 2015.
- 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.
- 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.
- 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.
- 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.
- 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: 176–187..
- 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.
- 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.
- 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.
Media related to Homo heidelbergensis at Wikimedia Commons
- Media related to Homo rhodesiensis at Wikimedia Commons
|Wikisource has the text of the 1920 Encyclopedia Americana article Heidelberg Man.|
- 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).
- 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.
- 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.
- 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". Journal of Human Evolution. 118: 56–71. doi:10.1016/j.jhevol.2018.02.010. ISSN 0047-2484. PMID 29606203.
- 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". Nature. 546 (7657): 289–292. doi:10.1038/nature22336. ISSN 0028-0836. PMID 28593953.
- 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.
- Diamond, Jared (April 20, 2018). "A Brand-New Version of Our Origin Story". The New York Times. Retrieved April 23, 2018.