{"id":234,"date":"2017-09-13T20:22:41","date_gmt":"2017-09-13T20:22:41","guid":{"rendered":"https:\/\/courses.lumenlearning.com\/suny-history-of-our-tribe\/chapter\/33-homo-heidelbergensis\/"},"modified":"2017-10-03T16:27:11","modified_gmt":"2017-10-03T16:27:11","slug":"33-homo-heidelbergensis","status":"publish","type":"chapter","link":"https:\/\/courses.lumenlearning.com\/suny-history-of-our-tribe\/chapter\/33-homo-heidelbergensis\/","title":{"raw":"33. Homo heidelbergensis","rendered":"33. Homo heidelbergensis"},"content":{"raw":"
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Homo heidelbergensis <\/i>(<600 kya)<\/b><\/h1>\r\n

(\u201csame\u201d \/ Heidelberg, Germany)<\/h2>\r\n[caption id=\"attachment_320\" align=\"aligncenter\" width=\"235\"]\"8.25\" Figure 33.1\u00a0<\/em>Model of Homo heidelbergensis head and shoulders. \u201cHomo heidelbergensis adult male \u2013 head model \u2013 Smithsonian Museum of Natural History \u2013 2012-05-17<\/a>\u201d by Tim Evanson is licensed under CC BY-SA 2.0<<\/a>.[\/caption]\r\n\r\n
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SITES<\/b><\/h3>\r\n

Europe:<\/b><\/p>\r\nSpain:<\/b> Sima de los Huesos\r\n\r\nFrance:<\/b> Arago\r\n\r\nGermany:<\/b> Mauer, Steinheim, Ehringsdorf, Sch\u00f6ningen, Bilzingsleben\r\n\r\nEngland:<\/b> Boxgrove, Swanscombe\r\n\r\nItaly:<\/b> Ciampate del Diavolo<\/a> (footprints)\r\n\r\nHungary:<\/b> V\u00e9rtessz\u00f6ll\u00f6s\r\n\r\nGreece:<\/b> Petralona, Apidima\r\n

Africa:<\/b><\/p>\r\nMorocco:<\/b> Jebel Irhoud\r\n\r\nEthiopia:<\/b> Bodo, Omo (Kibish Beds)\r\n\r\nZambia:<\/b> Kabwe\r\n\r\nTanzania:<\/b> Lake Eyasi, Laetoli (Ngaloba Beds)\r\n\r\nSouth Africa: <\/b>Florisbad, Elandsfontein\r\n

Asia:<\/b><\/p>\r\nChina:<\/b> Dali, Maba, Jinniushan\r\n

PEOPLE<\/b><\/h3>\r\n<\/b>Otto Schoetensack<\/a> and many more\r\n\r\n<\/div>\r\n

INTRODUCTION<\/h2>\r\nFor many years, fossil material from ~500\u2013200 kya from Africa, Asia, and Europe that was more human- or sapiens<\/i>-like was included in our own genus and species but was distinguished as \u201cEarly Archaic\u201d Homo sapiens<\/i> (EAHS). There was much debate as to when to draw the line between more erectus<\/i>-like forms and more sapiens<\/i>-like forms. The prevailing view was that material on all three continents was descended from H. erectus<\/i>. (The various geographic species distinctions for H. erectus<\/i> had not yet come into use.) Again, this is referred to as the Regional Continuity Model<\/b> (RCM<\/b>) for the origin of modern humans. In this scenario, erectus<\/i>-like forms on each of the continents slowly evolved into modern humans via gene flow between populations. This is in contrast to the Recent African Origin<\/b> (RAO<\/b>) model whereby modern humans evolved in Africa and moved out to eventually replace archaic forms elsewhere, e.g. H. erectus<\/i> in Asia.\r\n\r\nThe RAO model has gained in popularity due to a combination of the reevaluation of fossil material and especially DNA methods aimed at evaluating genetic distance between species, in terms of number of years since divergence from a common ancestor. The material from Asia that had previously been assigned to EAHS has been relegated to H. erectus<\/i>, with very little evidence for an intermediate form bridging the gap between H. erectus<\/i> and AMH.\r\n\r\nFossil material from Europe and Africa that was formerly assigned to EAHS is now termed Homo heidelbergensis<\/i>, with some researchers using Homo rhodesiensis<\/i> for some of the African material. It is now well accepted that H. heidelbergensis<\/i> was ancestral to both humans and neandertals (see Figure 33.2\u2014note that in this scenario, H. antecessor<\/i> is not considered to be ancestral to H. heidelbergensis<\/i>).\r\n\r\n[caption id=\"attachment_321\" align=\"aligncenter\" width=\"630\"]\"8.26\" Figure 33.2<\/em> Human phylogeny. <\/b>\u201cHomo-Stammbaum, Version Stringer<\/a>\u201d by Chris Stringer is licensed under CC BY-SA 3.0 DE<\/a>.[\/caption]\r\n

PHYLOGENY<\/b><\/h2>\r\nH. heidelbergensis<\/i> is thought to have evolved from a more derived form of H. ergaster<\/i>. Some researchers believe that the material from Tighenif (formerly Ternifine), Algeria, known by some as Homo mauritanicus<\/i>, is a possible transitional form between the two species. Mandibles from Tighenif are very similar to the type specimen, the Mauer mandible (see Figure 33.3) from the Heidelberg area of Germany, from which the species name is derived. In addition to the material from North Africa, the oldest material is from the Bodo site in Ethiopia, dated to 600 kya. Thus while an African origin is favored, some believe that H. heidelbergensis<\/i> is descended from H. antecessor<\/i> in Europe. Whether H. heidelbergensis<\/i> evolved in Europe or Africa, they had to have migrated from one continent to the other. Thus if H. heidelbergensis<\/i> evolved in Europe from H. antecessor<\/i>, a portion of the population migrated to Africa and evolved toward AMH, leaving some regional population of the remaining European stock to evolve into H. neanderthalensis<\/i>. The other more popular scenario has H. heidelbergensis<\/i> evolving in Africa from some descendant form of Homo ergaster<\/i>. A portion of the population then emigrated to Europe and evolved into neandertals while a portion of the remaining African stock evolved toward AMH.\r\n\r\nIn addition, a new species of hominin is also thought to be descended from H. heidelbergensis<\/i>. The Denisovans<\/b> (see this chapter), as they have come to be known due to their discovery in the Denisova Cave in the Altai Mountains of Russia, are thought to have branched off from the H. heidelbergensis<\/i> lineage that led to neandertals. DNA analyses show that Denisovans interbred with neandertals, as well as the first wave of AMH that left Africa, possibly around 125 kya and subsequently settled Melanesia and Australia.\r\n\r\n[caption id=\"attachment_322\" align=\"aligncenter\" width=\"300\"]\"8.27\" Figure\u00a033.3\u00a0<\/em>Type specimen: Mauer mandible. <\/b>\u201cUnterkiefer von Mauer (Replika)<\/a>\u201d by Gerbil is licensed under CC BY-SA 3.0<\/a>.[\/caption]\r\n\r\nContrary new DNA evidence suggests that H. heidelbergensis<\/i> branched off from some common ancestors\u00a0prior to 800 kya and is thus not part of our recent lineage. H. heidelbergensis<\/i> then divided into the neandertals and the Denisovan lineages, but interbreeding continued between them in areas of geographic overlap.\r\n\r\nThere is good evidence that H. heidelbergensis<\/i> gave rise to neandertals. There are skulls from the sites of Erhingsdorf, Germany; Fontechavade, France; and Sima de los Huesos, Spain, that appear to be transitional in that they exhibit characteristics seen in neandertals, such as similar brow ridge morphology, enlarged facial sinuses, occipital bun, etc. It is rare in paleoanthropology to have such good support for continuity in physical characteristics between ancestor and descendent species. However, the fact that those species lived more recently in time and in developed and populated areas certainly helps, in terms of facilitating discoveries.\r\n\r\n[caption id=\"attachment_323\" align=\"aligncenter\" width=\"264\"]\"8.28\" Figure 33.4\u00a0<\/em>Homo heidelbergensis from Steinheim, Germany. <\/b>\u201cHomo steinheimensis, holotype<\/a>\u201d by Dr. G\u00fcnter Bechly is licensed under CC BY-SA 3.0<\/a>.[\/caption]\r\n

DISCOVERY AND GEOGRAPHIC RANGE<\/b><\/h2>\r\nThe earliest discoveries of H. heidelbergensis<\/i> are from Germany. The type specimen was discovered in 1907 in Mauer, Germany. The oldest site is Bodo<\/b>, Ethiopia (600 kya). There are numerous H. heidelbergensis<\/i> sites in Europe (e.g.\u00a0Steinheim, see Figure 33.4) that date from as early as 500 kya and range from Spain through Eastern Europe. The greatest number of individuals came from the Sima de los Huesos<\/b> (\u201cPit of Bones<\/b>\u201d) (see Figure 33.5) site in the Atapuerca Mountains of Spain. Both H. antecessor<\/i> and H. heidelbergensis<\/i> sites in that area were discovered when a railway was built. The H. heidelbergensis<\/i> remains were found in a deep chamber inside of a cave, hence the name, \u201cPit of Bones.\u201d More than 32 individuals have been recovered, and most of them are juveniles. While there has been much speculation, the significance of this find in terms of cultural complexity (see ENVIRONMENT AND WAY OF LIFE below) remains elusive. There are a couple of sites in Asia that some researchers believe are also representative of H. heidelbergensis<\/i>, including Dali, China. Sites in Africa (also referred to as H. rhodesiensis<\/i>) include Jebel Irhoud in North Africa; Omo, Bodo, Ndutu, Eyasi, and Ngaloba in East Africa; Kabwe in Zambia; and Florisbad and Elandsfontein in South Africa.\r\n\r\n[caption id=\"attachment_324\" align=\"aligncenter\" width=\"300\"]\"8.29\" Figure 33.5\u00a0<\/em>Homo heidelbergensis from Sima de los Huesos, Spain.\u00a0<\/b>\u201cHomo heidelbergensis-Cranium -5<\/a>\u201d by Jos\u00e9-Manuel Benito \u00c1lvarez is licensed under CC BY-SA 2.5<\/a>.[\/caption]\r\n

PHYSICAL CHARACTERISTICS<\/b><\/h2>\r\nIn general, H. heidelbergensis<\/i> was less robust, both cranially and postcranially, than the erectus<\/i> grade and more robust than AMH. There is intraspecific variability in craniofacial morphology, with some specimens being more robust and others more gracile. While this is not surprising, considering the temporal and geographic range for the species, it has led to disagreement as to how to define it.\r\n\r\nH. heidelbergensis<\/i> is primarily distinguished from erectus<\/i>-like forms by its increased cranial capacity (1100\u20131400 cc\u201493% that of AMH) and more modern skull vault. Cerebral expansion, especially of the parietal lobes, led to increased cranial breadth in the superior aspect of the skull vault and thus a more vertically oriented skull. The occipital region is less angular due to reduced robusticity in the nuchal musculature. Some specimens have very pronounced brow ridges (see Figure 33.7) and some have speculated that those individuals represent males of the species. They share the presence of a canine fossa with AMH. The canine fossa is a depression above the canine on either side of the maxilla. The robusticity of their jaw morphology and tooth wear suggests that they were using their mouths and teeth as tools, for example, pulling meat off bones and working hides. Surprisingly, the middle ear bones were discovered for one specimen, and it could thus be determined that their hearing was comparable with our own (Mart\u00ednez et al. 2004).\r\n\r\n[caption id=\"attachment_575\" align=\"aligncenter\" width=\"882\"]\"image\" Figure 33.6\u00a0<\/em>Homo heidelbergensis by Keenan Taylor.[\/caption]\r\n\r\nSome H. heidelbergensis<\/i> are considered to be transitional and exhibited characteristics seen in the subsequent neandertals. Their midfacial region was (1) prognathic with a corresponding large nose and (2) puffy due to large frontal and maxillary sinuses. Their zygomatics have a \u201cswept back\u201d appearance, and the posterior skull vault exhibited an occipital bun. The outer layer of their long bones (the cortical bone) was thick, and the interior medullary (marrow) cavity was narrow. The species was taller than neandertals, with males and females averaging 5\u2032 9\" (175 cm) and 5\u2032 2\" (157 cm), respectively. Average weights for males and females were 136 lb (62 kg) and 112 lb (51 kg), respectively (Smithsonian Institution 2015).\r\n
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Review of Derived Characteristics<\/h3>\r\n