Homo heidelbergensis

Please do not edit this article unless this tag is removed, to avoid edit clashes. If the page has not been edited in a while, ask an admin what you should do.
The text on this page may have grammatical errors, incomplete data, broken images features and/or incorrect formatting.

## Homo heidelbergensis

A portrait of Arago 21 and 47, a possible member of this species.

## Statistics

### Diet

Omnivorous

*Homo* *heidelbergensis* is an extinct species of hominin that lived in Europe during the MIddle Pleistocene.

## History

* Bronze statue by Louis Mascré and Aimé Rutot, dated 1909-14.
* Bust by Elisabeth Daynès, 2010s.
1/2
    On October 21, 1907, Daniel Hartmann, worker at a sand mine in the Grafenrain open-field system in Mauer, excavated a hominin mandible found 24.63 meters (80.1 feet) in depth and about 10 kilometers (6.2 miles) from Heidelberg, Germany. However, he was aware fossils might appear here by the instruction of Otto Schoetensack over 20 years. After a well-preserved *Palaeoloxodon* skull was discovered in 1887, he had asked other workers to keep a lookout and taught them the characteristics of human material on regular visits to the mine where he searched for "traces of mankind". When dig out, the fossil was flung into the air and broke in two, and the which found out later, and a process from the left side was never recovered. A thick, cemented crust of coarse sand adhered to the canines and molars, typical of the Mauer site, and was caused by carbonation of calcium. A chunk of limestone, probably Muschelkalk, 6 centimeters (2.4 inches) long and about 4 centimeters (1.6 inches) wide was firmly attached to the sandy crust of the two frontal molars and the left side of the fossil. The contractor reported the discovery to Schoetensack immediately, who examined the site and fossil. He would present his studied in autumn of the next year by publishing a monograph. In Novermeber 19, 1907, he stated in a legal document that the contractor, Josef Rösch, lent the specimen to the Heidelberg University as a gift, which their Geological-Palaeontological Institute still thinks is their most important item. The mine also holds Hornstein artifacts that were excavated by Karl Friedrich Hormuth in 1924, which was interpreted to been made by the same species. Wilhelm Freudenberg (1933) discovered a frontal fragment that is also attributed to the species. Said species is often referred to as *Homo heidelbergensis*.

    Marcel de Serres (1828) first reported faunal remains from Caune de l'Arago, considering them antediluvian. Jean Abélanet (1963) recovered stone tools, which inspired Henry de Lumley to excavate for human remains; in 1964, this search was validated through the discovery of Arago 21. Marie-Antoinette de Lumley, his wife, described them that year, dated to the Riss glaciation and thus before *Homo neanderthalensis*. They compared the material to *Homo erectus* and suggested they were intermediary between the two species, referring to them as "Pre-Neanderthals" to avoid assigning a species. In 1979, they suggested the name "*Homo erectus tautavelensis*", but other authors reassigned them to *Homo heidelbergensis* (which is typically called a direct ancestor to Neanderthals. The only skull elements is the distorted face Argo 21 and 47, a parietal. It is often reconstructed using typical *Homo heidelbergensis* remains, with one done by moulder René David with M.-A. de Lumely's oversight and was completed in 1982. In 1982 and 1986, Antonio Ascenzi made another reconstruction; Éliane Spitery reconstructed in 1982 and 1984; as did Emanuel Vlček in 1986; Dominique Grimaud-Hervé did in 1982 and 1991; Gaspard Guipert did in 2005; and M.-A. de Lumley completed the most recent reconstruction in 2015. Guiper considered it a *Homo heidelbergensis* and defined the species as polymorphic. M.-A. de Lumley (2015) redescribed the material as *Homo erectus tautavelensis* by restricting *Homo heidelbergensis* and comparing directly with *Homo erectus* sensu stricto. They found the remains are similar with Ceprano and Petralona and also found similarities with early Neanderthal specimens such as Atapuerca, Swanscombe and Vértesszőlős. Arago 21 and 47 may represent the same individual in 2014, 148 hominin fossils were discovered, including 123 teeth, 5 mandibles, 9 upper limb elements and 19 lower limb elements representing a total of 18 adults and 12 juveniles (30 individuals). Despite being related to Mauer, the description of *Homo longi* (*Homo daliensis*) in 2021 recovers Arago 21 with Kabwe suggesting that it is a *H. rhodesiensis*, and Chris Stringer classifies it as 'near' Petralona.

The Montmaurin mandible was discovered by Raoul Cammas on June 18, 1949 in Grotte de la Niche in the oldest layer prior to Acheulean bifaces and close to occurrences of *Machairodus*. Quarrymen mined cliffside limestone and revealed a cave system that was visited by Father Henri Breuil and Count Henri Bégouën, who entrusted Louis Méroc with excavation from 1946 to 1961 (though he was not very interested with Niche since he thought it did not bear stratigraphy and that the mandible was from La Terrasse). Cammas disputed this, considering the 2 vertebrae and a tibia discovered from here. Vallois (1955) described it, first in French and later in English in 1956. It is from the Mindel-Riss interglacial 370-410 kya and was first assigned to *Homo erectus*. Mounier et al. (2009) assign an age closer to 400 kya and find relations with *Homo heidelbergensis*. Crégut-Bonnoure et al. (2010) later assigned it to 190-240 kya, an examination of lithic material gave 344-126 kya and finally, Martinex et al. (2020) place it between the age of the Sima de los Huesos Neanderthals and Arago.

## Classification

The anatomy of the specimen is mosaic, with the rounded dental arch and complete row of teeth being derived and the chronological delineation from *Homo neanderthalensis* and earlier forms showing basal traits. Stringer believes that it is separable from *Homo ergaster* and derived species such as Neanderthals and *Homo sapiens*, possibly being the LCA from the latter two. Others believe that development in Africa and Europe was gradual from *Homo erectus* sensu lato to *H. heidelbergensis* and then *Homo neanderthalensis*. Some thing that segregation is arbitrary and drop the name all together, classifying Mauer 1 as a late local *Homo erectus*. Most agree, however, that this fossil is connected to the derived line and may be an early descendant of a European and Asian migration that is indicated by *Homo georgicus*.

A 2021 study suggests *H. heidelbergensis* should be disused, with all known material referred to other species. They suggest most European specimens should be referred to pre-Neanderthals and African specimens to be referred to *Homo rhodesiensis*. The Maba Man is often included as an Asian extension to this species. However, others have suggested Denisovan affinities, which would make sense at it seems to fit the variation of Denisovans. Additionally, no other Asian *H. heidelbergensis* have ever been discovered, so it is likely not *H. heidelbergensis*. Chris Stringer states that Mauer, the holotype, still bears traits that may be seen as distinctive or idiosyncratic and takes blame for the species' descent into wastebasket status. If the species were to be considered valid, it would need extreme revision.

Sheela Athreya and Allison Hopkins (2021) note that *H. heidelbergensis* is a poorly-defined wastebasket taxon. They also find that this makes it difficult to determine which specimens may actually be of a distinct species and which should be reassigned. This is further found difficult by the fact that Mauer, a mandible, is the holotype rather than a skull. To combat this, they state that less species should be proposed and that they should be given better definitions. Another 2021 study by Suzanna White et al. compares "*H. heidelbergensis*" with other cattarhines, finding it to be a successful method for testing specimens who may be anomalous. However, they also find that *H. heidelbergensis* can be contained within *Homo erectus* sensu lato, *Homo sapiens* and Neanderthals. They note that the identity and status of *H. heidelbergensis* is still unknown. Because of a restricted hypodigm, it's temporal range is reduced to the Middle Pleistocene of Europe here until further notice. Another study suggests that mandibles from Montmaurin, Visogliano and possibly Arago 13 may belong to this species. It was first believed that the Montmaurin mandible was a Neanderthal, but it only bears 2-3 derived traits (similar enamel molar surfaces, enamel-dentin junction and the pulp cavity proportions) at most and is unfit for inclusion; It is more archaic and more basal than the Sima de los Hueso hominins that are supposedly older.

## Description

Analysis of Mauer 1 by Schoetensack was largely based on the expert opinion of Prof. Germann Klaatsch, which was only briefly acknowledged in the preface. Scheotensack notes now the teeth are disproportionately small compared to the jaw and would allow for more flexible development. He also finds a combination of features that had not been yet observed in human mandibles; the developed chin, dimensions, no stronger expression in relation to other tooth groups in the canines and other strange dentitions. He also suggests moderate and harmonious co-evolution. The specimen was estimated to be 20-30 years old at death. Dentition and similar anatomical features led Schoetensack to assign the specimen to *Homo*, making it still the earliest fossil of this genus from Germany. He decided to erect *H. heidelbergensis* based on the lacking chin, which led to classification elements that made him reluctantly suggest it was ancestral to modern Europeans and gave the vague descriptor "Pre-Neanderthaloid (which would, though given a terminology change, would remain accurate). He believed that the mandible advocates for an ape-like human ancestor, but is not the LCA itself. He clearly denotes his Darwinist beliefs elsewhere in the same publication. Mauer 1, which is noted by Stringer to be possibly distinct, is now diagnosed by a reduced chin, a notch in the submental space, parallel upper/lower boundaries of the mandible in the side view, mental foramina near the cheek teeth, a horizontal retromolar space, a gutter between molars and the ramus, a long jaw, deep fossa for the masseter muscle, a small gonial angle, an extensive planum alveolare, a planum triangulare that is developed and a mylohyoid line starting on level with the 3rd molar.

Reconstruction of the Arago crania indicates similarity with Asian *Homo erectus* (strong brows, a receding forehead, a low face, a depression between the brows, postorbital constriction, defined ridges under the orbits, a weak chin and developed prognathism, strong and thick jaws, U-shaped tooth rows and supposed sexual dimorphism). However, Arago are differentiated by a wider base, cheekbones oriented forwards, a larger and thicker brow and more defined postorbital constriction. Based on the fronto-parietal suture and robustness, Arago 21 is assumed to be a young man about 20 years old. Reconstructions find a 199 millimeter length, however, further measurements are negated here as they are based on presumably unrelated species. Material suggests strong jaw muscles. The mandibles have strongly developed tori, subhorizontal mylohyoid lines, deep and narrow submandibular forvea and a narrow-convex plane that merges with the tori. The teeth are proportionally large for a hominin of this time, with the P4 and M2 being especially large. The molar cusps retain anterior forvea, a mesial and distal trigonid crest, a cusp of Carabelli and 1-2 accessory cusps, which are all basal. Ralph Holloway (1983) estimated a cranial capacity of 1166 using Arago 21 and 46 with the help of the Swanscombe occipital (a *Homo neanderthalensis*), which is comparable to Peking and is the lower range of derived *Homo sapiens*. Holloway described this individual as "garrulous" based on the expanded Broca's area, but admitted it was pure speculation in 2004. Hyoids from *Homo* [aff.] *erectus* are quite archaic, which indicates a "proto-language", but a hyoid from Atapuerca assignable to *Homo neanderthalensis* suggests that later erectines may have had speech. The spine and torso are represents an atlas and axis, 2 clavicles, 4 humeri, an ulna, pelves and a sacrum, 2 iliac wings are known, 7 femora, 2 tibiae and 7 fibulae. The arms and legs are notably massive, the acetabulum is ovular (typical of erectines). Leg thickening may be a result to colder climates, greater physical activity, or both. Using femora and fibulae, an average height of 166 centimeters (5 feet 5 inches) is recovered.

## Paleobiology

30% of the Arago specimens died between 7-12 years of age, 30% of 18-30, 30% of 30-40 and 3% over 40. This gives an average lifespan (assuming survival past infancy) of 20-25 individuals with an infant mortality rate of 11%. If robust material are males, than they are slightly outnumbered by females. Non-tooth skull specimens Argo 21, 47 and 45 are assumed males, as well as 13 (a mandible), which is assumed of the same. Arago 2, 89, 119, 130 and 131 are assumed females. Iliac specimens Arago 44 and 121 are females. Vialet et al. (2017) examined the dental microstructure of Montmaurin, and in 2018 find that is it small for being a young adult, suggesting that it is a female based on a Sima de los Huesos specimen (but could not confirm based on a limited sample size).

### Technology

The de Lumley's categorize the tools from Caune de l'Arago as "Proto-Cherantian", a sub- of a possible subtype of the Neanderthal Mousterian industry, defined as producing few bifaces. They changed the name to "Mediterranean Acheulean" in 2004, and the former label was disused. 63% of tools are large stone shards, 32% are retouched tools, 3% are lithic cores and 2% are macro-tools. Excluding debris and simple chipping, small retouched tools make 90% of the assemblage, with macro-tools making 10%. Of retouched tools, 36% are scrapers, 16% are notches, 11% are Clactonian notches, 12% denticulate tools, 3% denticulated scrapers and 2% convergent scrapers. Macro-tools are 64% elaborate choppers, 13% primary choppers, 9% chopped tools, 7% rabots, 4% bifaces, 3% unifaces and 0.8% polyhedron-to-spherical-shaped tools. Of elaborate choppers, 60% are single edge, 26% have multiple points and 9% are converging points with 2 edges and one point. Bifaces are hypothesized to have been symmetrical on both sides for purely aesthetic reasons. Low quality quartz, sandstone, quartzose sandstone and limestone could be collected from river cobble; higher quality jasper and blue translucent quartz rocks could have been collected 15-30 kilometers (9.3-18.6 miles) away. Vein quartz would have been the most common material, and could have been common river cobble. It was used because it produced a reliable cutting edge and *H. heidelbergensis* were mostly producing scrapers. 90% of the G assemblage is made of vein quartz from the river below. Macro-tools and hammerstones are common from durable limestone, with more complex retouched tools made from higher quality flint or quartzite and bifaces from hornfel. Fire usage evidence in the upper segment of C dates to ~400 kya, with overall fire usage as a whole being scarce until around this time, possibly due to the invention of fire-starting technology or better maintenance strategies. An analysis of bear fossils from Bilzingsleben and especially Schöningen shows that bears were likely being hunted by *H. heidelbergensis*. Regions of the bear with little flesh, such as the limb extremities, bore precise cuts consistent with defleshing. As northern Europe would have had cold winters, the local population may have used bear skins as insulating clothing 300,000 years ago. The assemblage consists of adults, indicating hunting, and were probably stripped soon after death.

### Diet

Some remains from G and F appear to have been skinned and butchered while fresh based on striations that may suggest cannibalism. This could explain how the chest, hands and feet are absent, since they may have been left behind and eaten by animals. If this is true, than those in G specifically ate brains, tongues, flesh and bone marrow of the limbs of the recently deceased or killed. This would suggest ritual cannibalism rather than for survival, otherwise the entire body would have been used. Human baby teeth in the long-term cycles and seasonally inhabited beds are found, indicating entire families lived in the cave. They are not found in bed L, which could mean that the peopling of the cave was only by a small hunting party.

## Paleoecology

Closer dating of Mauer was only done in 2010, finding a range of 609,000-40,000. It was previously referred to as 500-600,000 years old in dated specialist literature based on poorer methods. Contextual fossils are thus best used for dating, and were not found until core drilling in the defunct Grafenrain sand mine took place in 1991. In 1995, several dozen cubic meters of and were screened for fossils, with mice teeth found and proven ineffective for precise dating, since these forms remained mostly unchanged over time. Palynological finding of similar vegetation of the Cromer interglacial period can describe the environment: alluvial forests in river valleys, forests on slopes and open forests on high locations that indicates dryness. Animal fossils from other layers of Grafenrain were identified as belonging to the same interglacial. These include: flying squirrels, roe deer, oak and birch trees, elk, wild boars, moles, shrews, beavers, hares and horses. It is possible that they ate woolly rhinoceros and hippopotamuses, but if they did is doubted. It is suggested that bears, wolves, leopards, hyaenas and sabre-toothed cats were likely predators of the Mauer 1 population, sometimes called "Heidelbergers". The Neckar river would have been juvenile at this time.

Caune de l'Arago is 35 meters long and varies from 5-9 meters in width, but the walls likely caved during the last 100s-1000s of years. Excavation was oversaw by the Institut de Paléontologie Humaine and the Centre Européen de Recherche Préhistorique de Tautavel; the limestone cave opens along an 80-meter-tall cliff face above the Verdouble river and overlooking the Tautavel plain. Fossilferous deposits go 11 meters down, stratified into Lower, Middle and Upper Stratigraphic/Stalagmitic Complexes with hominin remains found in the Middle and the beginning of the Upper Stratigraphic Complexes. These are further subdivided into 4 units and 17 beds with 4 Units. These Units start at the letter Q and reach the letter A at the uppermost Unit. Beds Q-C are specific hominin localities that span oxygen stages 14-10, 550-400 kya, made of sand and aeolian sandy loam that is overlain with thick stalagmitic layers and further by breccia. Nearly all hominin remains are from G (445 kya using uranium-thorium dating), and these populations signify the earliest inhabited caves in the Pyrenees. Throughout occupation, it provided access to mountains, plains and rivers. The plain and plateau swung between temperate and humid forested region that were dominated by pines, deciduous and cyprus trees with mediterranean plants, to a cold-dry grassy steppe that switched from forest steppe about 550 kya and reverting 480 kya and again in 420 kya (and continuing the pattern after occupation). The mammalian fauna of the cave during forested period features red deer, fallow deer, *Ovis ammon antiqua*, *Stephanorhinus hemitoechus*, tahrs, cave lynx, cave lions, dholes, red foxes, *Canis mosbachensis* and *Ursus deningeri*. The latter and *H. rhodesiensis* possibly occupied during different seasons of the year when human occupation was intermittent. In cold events, *Equus mosbachensis*, reindeer, *Bos priscus*, *Praeovibos* and *Stephanorhinus hemitoechus* could have been abundant. beavers make their first appearance as prey in beds G and J. Like other archeological sites, occupations in different layers prefer different prey, such as reindeer in L, fallow deer in J and muskox in G. *Ovis ammon antiqua* are found in all beds and may have also been killed by non-human predators (particularly in O, N and M). Dental development of animals under 2 years old may indicate how long occupation lasted: it was long-term in G; intermittent of a months in P, J, I, F. E and D; and short in L. Human baby teeth in the long-term cycles and seasonally inhabited beds are found, indicating entire families lived in the cave. They are not found in bed L, which could mean that the peopling of the cave was only by a small hunting party.

## Notable Specimens

* Mauer 1/GPIH 1/FM 15547 C "Heidelberg Man" or "Heidelberger": A very well preserved mandible and teeth discovered in 1907 and represents and enigmatic form during the Middle Pleistocene. The American Museum of Natural History classifies it as *Homo erectus* and is called FM 15547 C.
* Bilzingsleben: Fragmented human skulls were identified in 1974. 37 bones and teeth have since been found, representing three individual *Homo erectus bilzingslebenensis*, named by Emanuel Vlcek, from Bilzingsleben, Germany. These skulls were smashed postmortem, possibly as a religious burial ritual. It was found to be distinct from the Steinheim specimen. Some authors classify them as plesiomorphic *Homo sapiens*, *Homo heidelbergensis*, "*Homo steinheimensis*" or pre-Neanderthals. They are most alike to Aroeira 3 and Mauer.
* Arago: Many hominin individuals likely representing a hunting party seasonally occupying the cave.
* Montmaurin, Mandible of Montmaurin:

## Subspecies

### Reassigned subspecies

* *H. h. steinheimensis*
* *H. h. rhodesiensis*
* *H. h. daliensis*

## Synonyms

* *Euranthropus/Europanthropus*
* *Pseudhomo/Pseudohomo*
* *H. (Euranthropus) heidelbergensis*
* *Maueranthropus*
* *Rhenanthropus*
* *Palaeoanthropus heidelbergensis*
* *Pithecanthropus heidelbergensis*
* *Protanthropus*
* *Praehomo heidelbergensis*
* *Praehomo europaeus*
* *Anthropus*
* *H. erectus heidelbergensis*
* *H. erectus bilzingslebenensis*
* *H. erectus tautavelensis*
* *H. tautavelensis*
* *H. sapiens heidelbergensis*
* *H. sapiens bilzingslebenensis*
* *H. h. heidelbergensis*
* *H. heilderbergensis* (lapsus)

## Gallery

Bilzingsleben cranial fragments.Bilzingsleben cranial and mandibular fragments.Arago 13 (left) and 2 (right).Montmaurin mandible.
## References

1. ↑ 1.0 1.1 1.2 1.3 https://en.wikipedia.org/wiki/Homo\_heidelbergensis
2. ↑ 2.0 2.1 2.2 2.3 2.4 https://en.wikipedia.org/wiki/Mauer\_1
3. ↑ https://en.wikipedia.org/wiki/Tautavel\_Man
4. ↑ https://twitter.com/ChrisStringer65/status/1350001899752316928?t=h\_jJR7aaAA9RbnX3sl36Gg&s=19
5. ↑ 5.0 5.1 5.2 https://fr.wikipedia.org/wiki/Mandibule\_de\_Montmaurin
6. ↑ https://twitter.com/ChrisStringer65/status/1453798953644871685
7. ↑ https://onlinelibrary.wiley.com/doi/full/10.1002/ajpa.24330
8. ↑ https://www.sciencedirect.com/science/article/abs/pii/S0047248421001470?via%3Dihub
9. ↑ https://anatomypubs.onlinelibrary.wiley.com/doi/full/10.1002/ar.25190
10. ↑ https://en.wikipedia.org/wiki/Tautavel\_Man
11. ↑ https://en.wikipedia.org/wiki/Tautavel\_Man
12. ↑ https://en.wikipedia.org/wiki/Tautavel\_Man
13. ↑ https://phys.org/news/2022-12-humans-skins-years.html
14. ↑ https://www.heritagedaily.com/2022/12/humans-have-been-wearing-bear-skins-for-at-least-300000-years/145635
15. ↑ 15.0 15.1 https://en.wikipedia.org/wiki/Tautavel\_Man
16. ↑ http://research.amnh.org/paleontology/search.php?action=detail&specimen\_id=168995
17. ↑ https://en.wikipedia.org/wiki/Bilzingsleben\_(Paleolithic\_site)#Human\_remains
18. ↑ https://www.researchgate.net/publication/222412926\_The\_earliest\_settlement\_of\_Germany\_Is\_there\_anything\_out\_there
19. ↑ 19.0 19.1 19.2 19.3 19.4 19.5 19.6 19.7 19.8 https://studylib.net/doc/7690604/europanthropus-heidelbergensis
20. ↑ http://phylopic.org/name/d9ff0b54-8110-45ab-b5f2-c3145f9b59ba