Homo heidelbergensis

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Homo heidelbergensis
Temporal range: 0.7–0.2 Ma
Middle Pleistocene[note 1]
Skull, Natural History Museum, London - DSCF0431.JPG
Cranium 5 (skull with jawbone) from Sima de los Huesos (cast at Natural History Museum, London)
Scientific classification edit
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Genus: Homo
H. heidelbergensis
Binomial name
Homo heidelbergensis

Homo rhodesiensis
(Woodward, 1921)

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] African H. heidelbergensis has 4 subspecies: H. h. heidelbergensis, H. h. daliensis, H. h. rhodesiensis, and H. h. steinheimensis.[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 3] The species was originally named Homo heidelbergensis due to the skeleton's first discovery near Heidelberg, Germany.[3]

The first discovery—a mandible—was made in 1907 by Otto Schoetensack.[3][4] 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.[5] 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.[6][7][8][9]

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).[10] Its exact relation both to the earlier H. antecessor and H. ergaster, and to the later species Neanderthals, Denisovans, and modern humans is unclear.[11][12][13]

Modern humans have been proposed to derive from H. heidelbergensis via H. rhodesiensis, present in East and North Africa from around 400 kya.[14][15] 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 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 the disappearance of Homo antecessor.[16][17]

The delineation between early H. heidelbergensis and H. erectus is also unclear.[9][18][19]

Derivation and taxonomy[edit]

Homo heidelbergensis: Steinheim skull replica

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.[20][21][22][23][24] 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.[25]

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").[26] 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.[27]

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.[2]

Chris Stringer (2012) argues for Homo heidelbergensis as an independent chronospecies.[29] 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.[33][34]

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.[35]

"Rhodesian Man" (Kabwe 1) is now mostly classified as Homo heidelbergensis, though other designations such as Homo sapiens arcaicus[36] and Homo sapiens rhodesiensis[37] have also been proposed. White et al. (2003) suggested Rhodesian Man as ancestral to Homo sapiens idaltu (Herto Man).[38][39]


H. heidelbergensis adult male, reconstruction by Élisabeth Daynès (2010) based on fragments from Sima de los Huesos

Homo heidelbergensis is intermediate between Homo erectus and Homo neanderthalensis, with a typical cranial volume of approximately 1,250 cm3 (76 cu in).[40] "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."[41]

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.[42]

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).[43] 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.[44][45][46]

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.[47][disputed ]

Otto Schoetensack described the mandible Mauer 1 in his original species description in 1907:[48]

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."


One of hundreds of handaxes found at Boxgrove

Discoveries in a pit in Atapuerca (Spain) of 28 human skeletons suggest that H. heidelbergensis might have been the first species of the genus Homo to bury its dead.[49][50]

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.[51] Dental wear analysis suggests they were as likely to be right-handed as modern people.[52]

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.[53] This find could mean that modern humans and Neanderthals inherited the stone-tipped spear, rather than developing the technology independently.[53][54][55]

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.[56]

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).[57][58][59]


Steven Mithen[60] believes that H. heidelbergensis, like 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.[61] Furthermore, there is evidence that Homo heidelbergensis was right-handed. Handedness is associated with the development of language among hominins.[62] 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."[63]


Original type specimen from Mauer


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.[64]

The next H. heidelbergensis remains were found in Steinheim an der Murr, Germany (the Steinheim skull, 350kya), Arago, France (Arago 21), Petralona, Greece, and Ciampate del Diavolo, Italy.

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.[65] 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.[66] 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.[66] Ninety percent of the known H. heidelbergensis remains have been obtained from this site. The fossil pit bones include:

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.[67] 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."[68]

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.[69][70]

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."[3]


Replica of the Kabwe skull
Facial reconstruction based on the Kabwe skull by John Gurche (2010), Smithsonian Museum of Natural History
Reconstruction of Homo rhodesiensis based on the Kabwe skull, by Élisabeth Daynès (2010), Museum of Human Evolution, Burgos

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.[71] The Saldanha cranium, found in 1953 in South Africa was subject to at least three taxonomic revisions from 1955 to 1996.[72]

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.[73][74] 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).[75] Bada, & al., (1974) published the direct date of 110 ka for this specimen measured by aspartic acid racemization.[76][77] 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".[citation needed] However, with regard to the skull's extreme robustness, recent research[citation needed] 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.[citation needed] The skull is kept in the Natural History Museum, London.[78] There is a replica in the Museum in Livingstone, Zambia.

The 600,000-year-old[79] 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.[80] 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.[81] Although the skull is most similar to those of Kabwe, Woodward's nomenclature was discontinued and its discoverers attributed it to H. heidelbergensis.[82] It has features that represent a transition between Homo ergaster/erectus and Homo sapiens.[83]

Another specimen,[84] "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.[84] 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.[75] 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".[85]

The Saldanha cranium, or Elandsfontein cranium, are fossilized remains later identified as Homo heidelbergensis, found in 1954 in Elandsfontein, located in the Hopefield of South Africa.[86]

New evidence[edit]

Homo heidelbergensis firsts

The skull of the Homo heidelbergensis with odontogenic orbital cellulitis[87]

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.[88] 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.[88][89] 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.[90][91]


  • 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".[92]
  • 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.[93]
  • 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.[94]

See also[edit]


  1. ^ 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.
  2. ^ 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.
  3. ^ a b "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."[2]
  4. ^ "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".[28]
  5. ^ "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...".[30] "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".[31]
  6. ^ 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.[32]


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Further reading[edit]

  • 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. http://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. http://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. http://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. http://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: 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. http://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.

External links[edit]

Media related to Homo heidelbergensis at Wikimedia Commons

  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. 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.