Homo erectus

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.

## Statistics

### Diet

Omnivorous

*Homo erectus* is an extinct species of hominin that lived in Afro-Asia from the Early Pleistocene to Middle Pleistocene.

## History

—Ralph von Koenigswald discussing the Trinil finds over 50 years after Dubois had discovered them
    In October 1887, Eugène Dubois abandoned his academic career and left for the Dutch East Indies (now Indonesia) to search for fossil hominids, receiving no funding from the Dutch government since no human ancestors had then been discovered. Because nobody else was searching, he joined the local army and became a military surgeon. In July 1888, his work duties ceased and allowed him to resume search in Sumatran caves. He quickly discovered lots of animal bones, but failed for human remains, moving search to Java in 1890. Assisted by convict labourers and two army sergeants, Dubois searched along Solo River near Trinil in August 1891 and excavated Trinil 1, a molar, and a skullcap, Trinil 2, with archaic traits. He chose the name *Anthropopithecus*, which had already been given to a tooth from Siwalik Hills, India, as a species related to chimpanzees. In August 1892, Dubois' team found a femur that looked modern, suggesting bipedalism. It was about 50 feet from the original find; they believed these were from a single, "aged female" individual, renaming it *A. erectus*. In late 1892, he determined the cranium could be up to 900 cubic centimeters in cranial capacity, thus considering it transitional. In 1894, he renamed it *Pithecanthropus erectus*, a generic name that was coined by Ernst Haeckel to represent a "missing link". Thus gives the specimen the ID of *Pithecanthropus* 1. The remains would be later compared to the Peking Man, but classified in different genera. G.H.R. von Koenigswald recovered several early human fossils from Java, including Solo and the Mojokerto child. Von Koenigswald considered the former to be closer to humans, coining *Pithecanthropus modkokertensis*, but Dubois pretested as he viewed his specimens as an "ape-man" and not human. In 1936 and 1941, von Koenigswald discovered a skullcap of similar size at the Trinil 2 site, with discoveries from Sangiran convincing him that these skulls were from early humans. Dubois refused to acknowledge the similarity. Von Koenigswald and Franz Weidenreich compared these to Java and Zhoukoudian, concluding close relation. Dubois died in 1940, refusing to recognize these advancements. The fossils were so controversial that by the end of the 1890s, almost 80 publications covered them. Until the Taung child, these were the oldest human remains, with some suggesting that they were intermediate of the lost common ancestor of hominids and modern forms.

Richard Leakey and Alan Walker (1970s) described hominin remains from the east shore of Lake Turkana; partial skulls KNM ER 3733 and KNM ER 3883. They assigned them to *Homo erectus* and noted differing cranial capacities and a younger age. KNM ER 992 is a mandible described in 1972 as *Homo* indet. sp.. In 1975, Colin Groves and Vratislav Mazák assigned the latter specimen as the holotype as *H. ergaster*, meaning "working" or "workman". They referred KNM ER 803 but did not compare them to Asian *Homo erectus*. This would later cause taxonomic confusion over the definition of both species. Most scholars now lump the two species into the larger *H. erectus* because there is little anatomical difference and the supposed distinction between the two is blurry and is argued to not exist. This would make *H. erectus* a long-spanning species split into African (earlier) and Asian (later) morphs. In 1991, an expedition to Dmanisi uncovered animal bones, stone tools and a mandible (the latter by a team led by Medea Nioradze and Antje Justus). Heads of the expedition, Abesalom Vekua and David Lordkipanidze, were alerted of this the next morning and the mandible was fully freed by the end of the day. The heads and Leo Gabunia described this fossil back at Tbilisi. They recovered it as an indeterminate hominid. Dmanisi was further excavated but hominin remains were rare. In 1997, the right third metatarsal was found in the same layer. In May 1999, more bones were found. Extended rainfall damaged remains. Gocha Kiladze found a coin-sized skull fragment. Further fragments were found when Kiladze, Vekua, Lordkipanidze, Kakha Kakhiani and the head of the 1999 expedition found disarticulate fragments that formed a whole skull. In that same year, a well-preserved skull was recovered intact. In 2022, a isolate tooth was discovered from Orozmani, near Dmanisi, and considered to be of the same taxon.

## Description

The most complete postcranium is the Turkana Boy; with shorter arms (a modern feature), a longer tibia (indicating more bend in the knee while walking) and a cone-shaped torso (an archaic feature). This bauplan may explain how they lived in hot, arid seasonal environments through bodily thinning, which increases heat dissipation through altering the skin area. They were taller, with the Turkana Boy being about 1.62 meters (5 feet, 4 inches) and would have likely reached 1.82 meters (6 feet) if they had survived to adulthood. Adults probably ranged from 1.45-1.85 meters in height. They may have been the earliest species to be hairless based on the temperature of their environment, since a full coat would not have been efficient. Sweating, sexual selection and parasite load reduction are other suggested hair loss explanations. They were adapted for long-distance travel and inhabited lower altitudes than earlier taxa. However, hair loss may have occurred earlier, since it has been estimated that the process began about 3 million years ago. Human ancestors gained pubic lice from gorillas around this time as well, but this would have only been possible if most of the body hair had been lost by this time. Genetic evidence suggests that the melanocortin 1 receptor dates to 1.2 million years ago, which indicates the rise of hairlessness, since the dark skin produced to this would coincide with UV radiation protection. *H. erectus* had a face and skull that is distinct. The Turkana Boy has a cranial capacity of 880, which is close to their final limit, since this element was almost fully developed at death; this is greater than *Homo habilis*, though this change is not as significant as it sounds. *H. erectus* has a cranial capacity ranging from 600 to 910, and smaller samples include a range of 508-580. Postorbital constriction is present. The neurocranium is long, low and the forehead is flat and receding, merging with the brow at an angle. The nose would have projected forewards and had downwards-oriented nostrils like more modern species, possibly being an adaptation to warmer climate. Turkana Boy has a longer from-top-to-bottom face that had prognathic jaws, the latter and teeth are more basal and larger. The jaw is slanted backwards at a sharp angle, which indicates the lack of a chin. Other crania have prognathic faces and brows, receding foreheads, large teeth and projecting nasals. Though the Turkana Boy would have been 12 years old at death, their stature is similar to a 15 year old and their brain is comparable to a 1 year old derived *Homo sapiens*. This would make the species cognitively limited, but the invention of a new industry indicates an increase in function.

* Dubois' holotype.
* Speculative reconstruction of the Trinil skulls by J.H. McGregor in 1922.
1/2
The Trinil population were about 173 centimeters (5 feet 8 inches) tall and were able bipedal walkers. The femur is thickened, which indicates persistent running ability. The skull is thickened with a retreating forehead. A large jaw and teeth caused for prognathism, with the lower lips overhanging the lower mandibular margin. This would have made the chin look more undeveloped. The supraorbital torus was large and robust, supporting a neurocranium with a cranial capacity of 900, which is smaller than later specimens and should expected for such an early population of *Homo erectus*. Generally, the teeth were humanlike, but were larger and with large canines.

Dmanisi *H. erectus* has a cranial capacity of 546-775 with an average of 631. This overlaps *Homo habilis*. Skulls 1-4 have an encephalization quotient of 2.6-3.1, similar to the former and australopiths and the lower end of the latter. Skull 5 has an encephalization quotient of 2.4, within the range of *Australopithecus*. A well-developed brow, sagittal keeling, large orbits, premolars of the upper jaw having single roots and an angulated cranial vault distinguish them from *Homo habilis*. Skull 5 is the only complete skull, having a large prognathic face and a small braincase. These and large teeth make for a combination previously unseen in *Homo*, and were previously used to split them as species; if the skull were fragmented and split between 2 localities, they would have been split. The braincase is similar to other *Homo erectus*. Skull 5 shows that small brains and large faces and a prognathic, robust morphology are within the range of the Dmanisi specimens. They were small-brained with large brows and a stature, mass and limb proportions at the lower *Homo sapiens* range.

The Dmanisi morph was smaller than ***Homo erectus***, possibly due to being more primitive or being adapted to a different region. The limbs are comparable to *Homo sapiens* and sometimes to *Australopithecus garhi*. The absolute length of the legs is most similar to *Homo* than australopiths, with the leg length and the metatarsal morphology not as derived as other *Homo erectus*, which may suggest improved walking and running was not a sudden evolution, but continued slowly through the Early-Middle Pleistocene. Humeral torsion is quite low in Dmanisi, indicating a different arm movement and orientation, further suggesting a habitual supination and a more laterally-placed shoulder girdle. They were very capable of extended movements, and low torsion may be a basal trait in *Homo*. The spine is more similar to *Homo sapiens* and early *Homo erectus* than to australopiths. Lumbar lordosis is present in these vertebrae. The facet joints and the spinal flexion range are more similar to *Homo sapiens*, which indicates increased compressive loads, and in-turn, capable running and long-range walking. The feet were more medially oriented, which would have distributed weight more evenly. The feet bear modern morphology. Ian J. Wallace, Brigitte Demes, William L. Jungers, Martin Alvero and Anne Su (2008) state too little is known of these elements to reconstruct the feet for sure, believing a pelvis and more pedal bones are required. A juvenile skeleton from Dmanisi that is not directly associated with adults was announced in 2016.

## Classification

Studies suggest it and *Homo antecessor* split sometime during the Early Pleistocene based on morphological differences, dental morphology, geographic and temporal segregation. *H. erectus* is easy to differentiate from the contemporaneous *Homo habilis* and *Homo rudolfensis* since it has a vastly different morphology. *H. ergaster* was first dismissed by many prominent researchers on the basis that it was simply a synonym of *Homo erectus*. There is no doubt these two species are a cohesive subset of plesiomorphic *Homo*, but some question if they represent a single, widespread species or multiple species radiating. This has led to the creation of the terms *Homo erectus* sensu lato, which sinks *H. ergaster* (or the "African" form), *Homo antecessor* and various *Homo heidelbergensis* in, and *Homo erectus* sensu stricto for Asian fossils (in the wide and strict sense, respectively). However, this is unresolved and likely can not be resolved since the lineage bears little difference aside from geographic segregation and time period. *H. erectus* shares many features with *Homo* which has also blurred the lines of what is and is not distinct, if distinct at all. Intraspecific variation in these species is also incredibly high, which is used by many to support lumping *H. erectus*. David Strait, Frederick Grine and John Fleagle (2015) listed *H. erectus* as one of the 7 widely recognized species of *Homo*. Strait, Grine & Fleagle (2015) recover the following, assuming the African morph is indeed a distinct species as *H. ergaster*. A 2022 study suggests that the occurrence of *H. erectus* at Drimolen on the south tip of Africa is roughly contemporaneous with it's supposed date of evolution and the extinction of *Paranthropus robustus* and other hominids.

    D211 was described by Gabunia and Vekua (1995) as a basal *Homo erectus* based on dental similarities with *Homo erectus*. Günter Bräuer and Michael Shultz (1996) noted basal and derived traits and included the population was derived, despite age. Antonio Rosas and José Bermúdez De Castro (1998) suggested they should be *H. sp. indet. (aff. ergaster)*. Gabunia et al. (2000) described Skulls 1 and 2 as similar to *Homo erectus* and thought many traits suggest they were similar (brow, proportions, post-orbital constriction and cranial vault height and thickness). Thus, the analysis placed them into *H. ex. gr. ergaster*, noting that they may have been forerunners for *Homo erectus* and the ancestor to *Homo sapiens.*

    Vekua et al. (2002) described Skull 3 and its mandible, finding a resemblance to *Homo habilis* in brain size and facial characters consistent with small *Homo erectus*. Gabunia, Vekua, Marie-Antoinette de Lumley and Lordkipanidze (2002) describe D2600 as a jaw with proportions differing from all other Hominins, with a mosaic of characters blended from *Australopithecus, Homo erectus* and early *Homo*. Thus, they coin *H. georgicus* as a descendant to *Homo habilis* or *Homo rudolfensis*, before *Homo erectus* emerges. Sang-Hee Lee (2005) supports that the Dmanisi hominins are the same species, but does not specify which. Lordkipanidze et al. (2006) described Skull 4 and its jaw, noting similarity to all others except, possibly, for D2600. Tehy find it to be ancestral to ***Homo erectus*** and also note the subspecies (most of which are now disused or sunken into ***Homo erectus***). Skulls 1-4 and D2600 were analyzed by G. Philip Rightmire, Lordkipanidze and Vekua (2006) conclude they are the same species, with D2600 as tentative. They find them to be similar to *Homo habilis* (via size, cranial capacity) and *Homo erectus* in far more characters. Thus, they find primitive features to be retentions. Rightmire, Lordkipanidze and Vekua conclude the habline features are plesiomorphic and retentions, with no reason to exclude them from *Homo erectus*. Vekua then supported D2600 as a distinct species, though others were unsure; Vekua preferred it keep the name *H. georgicus*. Matthew M. Skinner, Adam D. Gordon and Nicole J. Collard (2006) compared skulls, finding sexual dimorphism of a greater expected degree, suggesting this: either they were 1 taxon with high sexual dimorphism of uncertain inclusion in *Homo*, or that D2600 should be split. In 2008, a more detailed analysis by Rightmire, Lordkipanidze and Adam Van Arsdale conclude an excessive dimorphism rate comparable to gorillas, as a 1 species. Lordkipanidze, Vekua, María Martinón-Torres, José María Bermúdez de Castro, Aida Gómez-Robles, Ann Mergvelashvili and Leyre Prado (2008) analyzed the teeth of Skulls 2 and 3; they find a mosaic of *Australopithecus*, *Homo erectus* and *Homo habilis*-like traits. D2600 diverges in size and root morphology, though, noting *Homo habilis* does have dental dimorphism. P. James Macaluso Jr. (2010) concludes Skulls 2 and 3 are the same taxon, but D2600 is tentative.

    Lordkipanidze et al. (2006) describe Skull 5 as the same individual as D2600, as the same taxon as the rest with variation similar to *Pan* and *Homo sapiens*. They find orthognathics to be women or subadults and prognathics to be men. They use this to suggest that some hominid species are actually variation of known species, and, in-turn, synonymizing *Homo erectus*. They find it to be from a migration of *Homo erectus* from Africa, using the quadrinomial name *H. erectus ergaster georgicus*. Jeffrey H. Schwartz, Ian Tattersall and Zhang Chi (2014) respond, rejecting that they are the same taxon and suggest their quadrinomial name is invalid as per the ICZN. They also question their methods and how they differentiate species. They find Skull 5 should keep the name *H. georgicus*, stating that rejecting this idea is a "radical proposition to which few would subscribe". The other side responds in 2014, reaffirming that it is 1 species and that if what Schwartz et al. (2014) was true, 4 species would be contemporaneous, that they ignored data deliberately and that the ICZN actual has no regulation over quadrinomial names and it should not be dismissed. Skull 5 was analyzed in 2017 and compared to *Paranthropus boisei, Homo sapiens* and other plesiomorphs, finding that all remains are of one taxon. They question species diversity in *Homo* and do not refer the remains to any known species, noting a "continuum of forms". In this, they mean Skull 5 shares features with *Homo habilis* and 1, the largest brain, with *Homo erectus; **Homo erectus*** and *Homo habilis* are hypothesized to constitute of a single lineage that emerged in Africa and spread into Eurasia. They suggest the Dmanisi hominins are of an anagenetic sequence descended from *Homo habilis* but ancestral to ***Homo erectus***, nearer to the base of the ***Homo erectus*** lineage and separate from *Homo habilis*. The name *Homo dmanisensis* was used once by Otavio Barduzzi Rodrigues da Costa to refer to this species once, but never again.

### Evolution

The presence of "superarchaic" genes within the genome of Neanderthals and Denisovans, shared by a small amount of modern humans in West Africa, suggests that their last common ancestor interbred with a basal species that had been living in Eurasia since at least 1.9 million years ago. They speculate that this population was likely the same as that of Dmanisi. Dmanisi *H. erectus* may have been one of the first species to migrate from Africa. However, other scraps are known but cannot be confidently assigned to the species level. Dmanisi is deposited atop a thick volcanic layer radiometrically dated to 1.85 million years. Little time passed between this event and the Pleistocene sediments deposition. Paleomagnetic analysis determine the sediments are likely 1.77 million years old, deposited during the Upper Matuyama chron. Other animals discovered here reinforce this date. The hominin-bearing layer was dated to 1.81 ± 0.03 million years in 2010. This time in Georgia may have had depleted resources and a refuge of Hominins in this region. The geography would have been favorable to Hominins, who reached it through the Levantine corridor. Georgia may have been used as a sort of "foothold" before expanding outwards. Some tools found in Romania, Balkans and Spain share similar form to the ones at Dmanisi.

One 2011 study suggests West Africa and early *H. erectus* dispersed into East Asia to form the Peking Man and Southeast Asia for the Java Man. By 1.6 million years, *H. erectus* reached Sangiran and Java, with Zhoukoudian 780,000 years ago in a second wave. Sangiran *H. erectus* have bigger and more similar to basal Western *H. erectus* and *Homo habilis* teeth than the derived Zhoukoudian. However, the latter's teeth reduce over time, probably indicating another colonization of Java by the Zhoukoudian *H. erectus* or a closely related population. Due to superficial resemblance to Trinil, the Nacurrie 1 skull (and related specimens such as Kow Swamp and Coobool Creek) was once thought to have such cranial elongation due to interbreeding between the earliest Australians and relict *Homo erectus*. However, this was determined to be due to mother-inflicted cranial modification, rather than the application of bandages as seen in some populations or interbreeding.

## Paleobiology

The vertebrae of the Turkana Boy do not indicate the presence of respiratory muscles required for speech. Bruce Latimer and James Ohman (2001), however, find this individual had skeletal dysplasia and scoliosis. This indicates *H. erectus* may have used a proto-language less advanced than, but having the framework of, apomorphic language. Other individuals have the auditory capacity that could sense human speech.

Lordkipanidze believed that Dmanisi *H. erectus* used aggressive scavenging (throwing small rocks at predators to steal their game). It is also plausible power scavenging was employed in groups for protection, possibly leading to kinship-dependent social cooperation arising.

### Life History

A female *H. erectus* pelvis from Gona suggests they were very capable of child birth with a prenatal brain size of 315, 30-50% of an adult's brain size. This falls between chimpanzees and *Homo sapiens*. The Mojokerto child suggests a prenatal brain 72-84% of an adults, which suggests a more apelike trajectory. *H. erectus* has a postnatal growth intermediate of *Pan* and *Homo sapiens*. This suggests altriciality evolved later in human evolution, possibly in the concestor of *Homo neanderthalensis* and Denisovans. This may also suggest a lifespan lower than *Homo sapiens*. The SK 54 cranium was historically allocated to *Paranthropus*, but more recent morphological analysis suggests that it may be early *Homo* comparable with Mojokerto, a *Homo erectus* individual.

### Society

4 sites outside Ileret, Kenya preserves 97 footprints made 1.5 million years ago by a group of ~20 people, probably all being males of some specialized task group (like a foraging party or border patrol. If this is true, this would indicate sexual division of labour, which differentiates hominids from most other similar groups. apomorphic hominids are known to do this, with *Homo erectus* observed doing similar things possibly to defend from large savannah predators, avoiding areas with a high density of predators. Male-male, female-male friendships may have been important in their societies. Mothers did not have the same child-rearing capabilities since their children grew faster. They may have lived in a monogamous society due to less sexual dimorphism, as found in apomorphic apes. It is unknown if *H. erectus* had humanlike sexual dimorphism rates, but their women would have been taller than their ancestor species if true, which may have been caused by reduced pressure or a dietary/female fertility shift. If these are correct, it could indicate a shift in behaviour for the females, which could have been harems hard to maintain.

    *H. erectus* lived in the dangerous savannah, likely forming large groups and warding off predators with tools and counter-attack measures. Apomorphic primates form large male only troops and have effective counter-attack plans. A similar behaviour may be plausible for early *Homo*. Leslie C. Aiello and R.I.M. Dunbar (1993) estimated the group sizes of *Homo habilis* and *Homo rudolfensis*, using neocortex (which correlates to group sizes) size to find a group size of 70-65 individuals. Bipedalism is also energetically cheaper, so groups may have been larger. They find the estimate for *H. erectus* to be 91-116 individuals. Counter-attacks were probably carried on to *H. erectus* and were probably further developed. It was likely the first primate to transition to a social carnivore. Opportunistic hunting likely evolved from this due to competition with other carnivores. This niche swap may have changed a number of things throughout human evolution. Opportunistic hinting, predator defense and confrontational scavenging may have triggered a psychological transition and acquiring prey likely became collaborative. It is speculated labour was divided based on sex, with men practicing social hunting (as apomorphic primates do) and women foraging. Individuals likely shared meat, strengthening bonds with each other. Foragers would have also done the same. This may have caused friendships and monogamous pair bonds. These may have been carried on and amplified through generations.

The Turkana Boy's spinal cord is narrower than *Homo sapiens*, meaning the respiratory muscles were likely not as developed to produce or control speech. Bruce Latimer and James Ohman (2001) diagnose this specimen with skeletal dysplasia and scoliosis, however, noting this specimen should not be used to represent the entire species. Marc Meyer et al. (2006) describe an *H. erectus* from Georgia with vertebrae and spine within the range of *Homo sapiens*. This suggests this individual could talk. They find speech was likely possible in *Homo* very early on, diagnosing the Turkana Boy with possible spinal stenosis, as a candidate for congenital defect. Regula Schiess et al. (2013-2014) conclude no evidence of congenital defects in Turkana Boy. They thus consider it representative of *H. erectus*.

### Diet

    Richard Wrangham (1999) suggested *H. erectus* speciates from *Homo habilis* due to fire use and cooking 2 million years ago, which would explain the doubling of brain size in 500,000 years and the appearance of the typical human body. Cooking speeds nutrient absorption due to making proteins easier to digest, which destroys pathogens and increases the carrying capacity of their environment; this further allows their population to grow, causing selective pressure for sociality, which requires the greater brain meat consumption allows for. However, the usage of fire does not correlate with significant technological advancement in the fossil record, and cooking was probably not commonplace until after 400,000 years ago. *H. erectus* may have been the first hunter/gatherers based on their new meat dependence, emphasizing teamwork, labour division and food sharing. Sites mostly preserve big game, like elephants, rhinoceros, hippopotamuses and board, having many leftovers after a kill, which could have employed preservation methods such as drying to store food, if they used most of their prey. They probably required large game meat, with their disappearance in Levant correlated with the extinction of *Palaeoloxodon*. Their diets are location-dependent; the Geher Benot Ya'agov *H. erectus* ate 55 types of fruits, vegetables, seeds, nuts and tubers, using fire to roast plants, but also eating amphibians, reptiles, birds, deer and elephants and invertebrates; FwJJ20 lakeside East Turkana *H. erectus* ate bovids, hippos, rhinos, turtles, crocodiles and catfish, with larger animals likely scavenged and the turtles and fish likely caught live. A 2022 study finds that sampling error has caused many to think that *H. erectus* relied solely on meat, instead finding equal consumption of roasted tubers and foraging to sustain caloric requirements. Some Afro-European sites targeted specific species, such as Olorgesaile, with 50-60 individual *Theropithecus oswaldi*. This increase of meat usage would suggest development in group hunting strategies in the Middle Pleistocene. *H. erectus* were not as dentally versatile as their ancestors, processing a narrow range of food, with tools used to process tough food and leading to a strong jaw to be unnecessary. This may have increased a flexible diet, but does not equate to a varied diet. This may have allowed *H. erectus* to live in many environments beyond Africa. Trinil populations likely drove *Megalochelys* extinct due to overhunting.

### Technology

*H. erectus* invented the Acheulean tool industry, being the first to make lithic flakes larger than 10 centimeters and hand axes; the latter were likely multi-purpose, used for slicing meat, wood or plants. Thomas Wynn (1979) argues these technologies require operational intelligence, as they are more complex than the Oldowan industry, finding no significant intelligence disparity between *H. erectus* and *Homo sapiens*, with increasing intelligence likely not greatly affecting cultural evolution. Juveniles, however, who have a faster growth, show a lower intelligence. Some sites, such as Melka Kunture, Olorgesailie, Isimila and Kalambo Falls show a mass accumulation of hand axes, likely to do generation occupation. 1.76 million years ago from West Turkana shows the first of the Acheulean industry, but Oldowan lithics are also present, with both technologies coexisting for some time.

    *H. erectus* used the first fire, but the exact date is unknown because campfires preserve poorly in the fossil record. FxJj20 Koobi Fora and GnJi Chemoigut age as far back as 1.5 million years ago, with Wonderwerk Cave at 1.7 million years. They would have transported fire to caves and maintained them for some time, likely when the opportunity arose. The firekeepers would have had to have known slow-burning materials, such as feces. After 400-300,000 years, fire becomes much more present in the record, which may be explained by advanced keeping techniques after time; this may also suggest *H. erectus* opportunistically used fire. Fire starting was invented and lost multiples times over different populations, with the earliest evidence of this as Gesher Benot Ya'agov, 700,000 years old. This fire is recorded in several layers close to water, which indicates an unnatural fire. Fire light may have increased walking hours, where the additional ones were probably used for socialization. Fire usage is linking to the origin of language. Fire also made sleeping on the ground possible since the flames would deter predators.
The migration into Europe was also probably linked to fire, since it was very cold; usage here after 400-300,000 years ago is absent. If early European *H. erectus* lacked fire, it is unknown how they remained warm, avoided predators and prepared food. Lower light in these areas correlates to less natural fires. It is possible they may have knew how to maintain fires in certain areas, preparing food distanced from home, which would lead to a spread out range of fire use and hominin activity in the archaeological record. *H. erectus* may have only migrated north during the warmer interglacials, where fire, food storage and clothing was not required, with their dispersal patterns suggesting they stayed in the warmer lower-to-middle latitudes.

    A 366 x 427 × 30 centimeter (12 × 14 × 1 foot) circle constructed of volcanic rock were found in 1962 in Olduvai Gorge. Rocks piled 15-23 centimeters 6-9 inches) high in 61-76 centimeter (2-2.5 foot) intervals were suggested by Mary Leakey to support poles that supported a windbreak or hut, since apomorphic tribes construct similar walls to build temporary shelter, using branches as poles and grasses and animal hide as a screen. This is 1.75 million years old, being the oldest architecture. European dwellings following the Holstein Interglacial are claimed in Bilzingsleben, Terra Amata, Fermanville and Saint-German-des-Vaux. The oldest dwelling and campfire are recorded in 700,000-year-old Přezletice, with a base 3 × 4 meters (9.8 × 13.1 feet) at the exterior and 3 × 2 meters (9.8 × 6.6 feet) at the interior; probably being a firm surface hut with a vaulted roof constructed from thick branches or thin poles, supported by a foundation constructed of large rocks and earth and was likely a winter base camp. 1.6 million-year-old Wonderwerk Cave is the first evidence of cave habitation, with a global sporadic until ~600,000 years.

It is unknown when clothing first arose, having an earliest estimate of 3 million years (due to a lack of body hair). Head and body lice, the latter only inhabiting human clothing, diverged 170,000 years ago, meaning clothes were well in use at this point, even before encountering colder climates. Animal hide is thought to have been used first as clothing, with the oldest hide scrapers at 780,000 years; this does not directly indicate clothing. It is postulated that early hominins like *H. erectus* had access to boabab trees and weaverbirds, to which they would have lived around frequently and may have allowed early taxa to weave simple baskets and use this technology to expand Out of Africa.

Archeulean industry on islands unconnected to the Pleistocene mainland may represent seafaring *H. erectus* as early as 1 million years ago in Indonesia, arriving to the islands of Flores, Timor and Roti, which would have made crossing the Lombok Strait required before 800,000 years at the least. They may have been the first seafaring Europeans, also crossing at the Strait of Gibraltar. A 2021 genetic study on 3 isolated *H. erectus* populations found in interbreeding with *Homo sapiens*. If they did sea fare, this represents great planning abilities, which would have taken months. The population of Flores may have became *Homo floresiensis*, with another population possibly becoming *Homo luzonensis*.

*H. erectus* is known to have expressed themselves, possibly being the first artists. This is seen in *Pseudodon* shell DUB1006, which bears engravings possibly referable to *H. erectus*. They also collected red pigments such as ochre, which are found in lumps at some sites. Engraved lines on an ox rib associated with Archeulean lithics were found, having similar meander designs to human cave art. The same is found in 3 ostrich eggshell beads. The Venus of Tan-Tan and the Venus of Berekhat Ram are thought to have been produced by *H. erectus* from rocks weathered to look similar to humans, engraved to further match. The latter has trace pigments on the front side, possibly being coloured. Marek Kohn and Steven Mithen (1999) suggest the mass-produced hand axes that accumulate in some sites show no wear, and may actually be display tools the males would use too try and find a partner, discarded afterwards. However, the reported lack of wear may be due to lack of use-wear studies, with only a few sites having these accumulations. This suggests they were probably generational and not for sexual selection[*citation needed*].

10,000 Oldowan tools are known from Dmanisi, and the fossil-bearing Kvemo Orozmani site near Dmanisi bears Oldowan as well.

Socotra: Valery Zhukov discovered the Oldowan tools from Socotra in 2008 near Hadibo, publishing a small Russian book in Moscow in 2014 to share the results of his finds and presenting ~100 photographs of the technology. In 2009, an expedition worked to discover more Oldowan near wasi Tharditror and Raquf village. Special attention was given to the former and the rock terraces between the sites. They discovered thousands of artifacts 2-3 kilometers northwest of Hamer Kahyuf hill in "two ancient workshops". This industry lacks cores and having mostly choppers of one and both sides. They found large scrabbles at the site as well. At Raquf, they present the same typological line, which allowed for linkage. Near Aihaft, one more site was found containing artifacts most similar to Hadibu valley. This suggests the presence of *H. erectus*-like hominins on Socotra abusing the raw materials and water sources the land bridge-connected island provided. They did not report any Microlithic sites.

### Paleopathology

The Turkana Boy has juvenile spinal disc herniation that caused scoliosis since it was juvenile. This would typically cause lower back pain and sciatica, which probably restricted this individual from walking, bending and other activities. This individual survived into adolescence probably due to group healthcare. The 1000-700,000 year old Java Man specimen has an osteocyte on the femur, probably Paget's disease of bone and osteopetrosis, which causes skeletal fluorosis due to eating food contaminated with volcanic ash filled with fluorine. This specimen was found in strata filled with ash, and probably died within a few days to weeks, as do livestock who do the same.

    Skull 4 is an individual who has no remaining teeth but one, who lived a long time after losing their teeth. Though it would have technically been possible to live by themselves, they were likely aided by group members. They may have eaten animal brains or marrow to survive. D2280 bears 4 ovular lesions on the frontal, left parietal and occipitals, which are indicative of trauma, cysts, metastatic cancer and infectious disease. Additionally, a large, shallow depression with slightly elevated boundaries and being concave so that the diploë are lacking or absent sits on the left parietal; this is very likely to be a result of a trauma injury. Two other depressions on the left side of the frontal and occipital may have arisen due to blunt force tauma, which makes intrapersonal violence plausible. The location and structure of the 4th lesion suggests a diagnosis of treponemal disease, which makes lesions on the cranium and especially the frontal common. Perimortem perforations on the occipital were caused by large carnivorous predators or scavenging birds. This mosaic of pathology makes this individual the first instance of trauma in *Homo*.

## Paleoecology

Dubois' collected was transferred to Naturalis (Leiden, the Netherlands) between 1895 and 1900. Trinil 2, the first skullcap discovered and the holotype, has been dated using the Trinil Fauna and the nearby geological horizon to about 700 kya by Ralph von Koenigswald. Though this is generally agreed upon, a group of Dutch paleontologists used more than 20,000 animal fossils from the Dubois collection in the 1980s to date the layer Trinil is known from. This layer if known as the Trinil H. K. Fauna (H. K.=Haupt Knochenschicht), being the main fossilferous layer, finding an age of 900,000-1,000,000 years old. A paper from 2014 finds a maximum of 0.54±0.10 million years and a minimum of 0.43±0.05 million years using Ar-Ar and luminescence dating from human-predated shells at Trinil. Fossils such as Sangiran 2 and Mojokerto may provide an even earlier date, and the systematics on this are still underway. Fossils at Trinil are considered the holotype. They were found scattered in an alluvial deposit and detractors had doubted if they were of a single taxon or individual. Rudolf Virchow (1895) argued that the femur was of a gibbon, with Dubois having difficulty defending his arguments since excavation was not extended and the stratigraphy of the finds were not provided in enough detail. The femur is very modern in appearance, which was thought to have possibly been a reworked fossil. Based on anatomical and archeological evidence, they likely fed on meat like other *Homo erectus*, and may have been a rare species. They used shells to cut meat. Their inferred dispersal times coincides with the extinction of *Megalochelys*, likely due to overhunting, since this animal would have been slow and could have been stored for some time. With being about 1.8 million years old, it is likely that these Javanese populations were some of first first Out-of-Africa dispersals. They likely lived in ever-wet forests that resembled a savannah, but was hydromorphic, Plants from Trinil include grass, ferns, *Ficus* and *Indigofera*, which are typical of lowland rainforests.

All of Dmanisi except for Skull 5 and its mandible (who are somewhat younger) are contemporaneous. They all lived near a lake shore that was formed through the damming of the Maashavera and Pinazauri rivers. This was a temperate environment that was relatively humid and forested (gallery and woodland, grasslands, bush lands, tree savannahs and rocky terrain with shrubs). Cold winters are also known to have taken place. This climate is probably comparable to a mediterranean climate. Most a=faunal remains suggest a predominatly forest-steppe ecosystem with some parts being full steppe and forest. Forests covered mountain highlands and ground along river channels with flat river valleys covered by steppe vegetation. Forests were probably the most widespread at Dmanisi since deer comprise 80% of all remains. Hackberries are found in association with Hominins at Dmanisi, so they likely fed on them. Pikas, lizards, hamsters, tortoises, hares, jackals, deer, large cats, bears, horses, rhinos, flightless birds, hyaenas, gazelles, antelopes, bison and giraffes are known from Dmanisi. The Dmanisi hominins may have accumulated due to carnivore activity. Beetroots, berries and fruit are known here. Most fossils are known from the 4th layer, including the hominins. Layers 2 and 3 are less fossilferous, with almost no carnivores and no rodents or reptiles. This may be preservation bias or paleoecological changes based around aridisation during the Early Pleistocene.

## Subspecies

### Reassigned Subspecies

* *H. e. georgicus*
* *H. e. soloensis* (=late *H. erectus*?, =*H. soloensis*)
* *H. e. tautavelensis* (=*Homo heidelbergensis*/Neanderthal)
* *H. e. yuanmouensis* (=Yuanmou)
* *H. e. bilzingslebenensis*
* *H. e. lantianensis* (=Lantian)
* *"H. e." "wushanensis"*
* *H. e. mapaensi*
* *"H. e." palaeojavanicus*
* *H. e. narmadensis* (=*H. sapiens*?, =Denisovan?)
* *H. e. ngandongensis*
* *"H. e. naledi"*
* *H. e. cepranensis*
* *"H. e. antecessor*"
* *H. e. heidelbergensis*
* *H. e. reilingensis*
* *H. e. mauritanicus*
* *H. e. sapiens*
* *H. e. capensis*
* *H. e. gongwanglingensis*
* *H. e. chenchiawoensis*
* *H. e. hexianensis*
* *H. e. habilis*
* *H. e. wadjakensis* (=Wajak)
* *H. e. hungaricus*
* *H. e. aesernensis*
* *H. e. ergaster*
* *H. e. olduvaiensis*
* *H. e. leakeyi*
* *H. e. pekinensis*
* *H. e. nankinensis*

## Synonyms

* *H. louisleakeyi*
	+ *H. leakeyi* (rejected)
* *H. olduvaiensis*
* *Telanthropus*
* *H. kenyaensis*
* *H. okotensis*
* *H. mauritanicus*
* *H. ergaster*
* *H. lantianensis*
* *H, yuanmouensis*
* *H. dmanisensis*
* *Atlanthropus*
* *Homo (Proanthropus) louisleakeyi*

* *Anthropopithecus erectus*
* *Pithecanthropus erectus*
* *Pithecanthropus modjokertensis*
* *Pithecanthropus robustus*
* ***Pithecanthropus erectus europaeus* (=Sarstedt)**
* *Homo-proscopinus*
* *Hylobates giganteus*
* *Hylobates gigas*
* *H. modjokertensis*
* *H. trinilis* (=Trinil)
* *H. trinilensis* (=Trinil)
* *H. e. javanensis* (=Trinil)
* *H. e. dubius* (=Sangiran 5, 9)
* *H. e. robustus* (=Sangiran 4)
* *H. e. yunxianensis* (=Yunxian)
* *H. e. erectus* (=Trinil)
* *H. neanderthalensis (Asianicus) pekinensis*
* *H. neanderthalensis (Asianicus) javanensis*
* *Australopithecus africanus modkojertensis*

## Notable Specimens

* KNM-WT 15000 "Turkana/Noriokotome Boy": Once interpreted as a young man (hence the name) but actually of an undetermined sex, the Turkana Boy is a near-complete fossil found on the west shore of Lake Turkana. It was discovered by Kamoya Kimeu (1984) and described Leakey, Walker, Frank Brown and John Harris (1985) as a *Homo erectus*. However, it is traditionally recognized as *H. erectus* for those who recognize it as distinct.
* DNH 134 "Simon": A 2.04-million-year-old juvenile *H. erectus* skull consisting of the top and back of the braincase, not the sides. It was recovered from Drimolen Cave and named after Simon Mokobone. It is one of the oldest *H. erectus*.

* Kozarnika: A molar from Kozarnika cave in Bulgaria that is from an early *Homo* species. It is similar in many ways to *H. erectus*, including in age. Additionally, a 8-centimeter-long bovid fragment recovered from the cave and dated to around the same time as the tooth bears 27 marks on it's edge. This could have been made while butchering prey, but since the strikes hit the bone, they were interpreted as symbolism made by the *H. erectus* that lived in this cave. These would have been some of the first populations to leave Africa.
* UB-10749: A 6-12 year-old child, known from a single lower lumbar vertebral body discovered in 'Ubeidiya, Israel. It is one of the oldest hominins from the Levant region. It was described in 2022 and assigned to *Homo *sp.**, but it likely belongs to an "erectine" species such as *H. erectus* based on correspondence and morphology that most aligns. The ossification of the vertebrae is delayed and the individual was very tall: the largest estimates give a height of 8'1, 9'2 and estimated juvenile height being 5', but it is most likely that this individual was 6'4 if adulthood had been reached. It supports two distinct Out-of-Africa dispersals, lithic tradition variants and highlights Eurasian Pleistocene niches.
* AMNH coll.: Various unidentified specimens listed as belonging to *Homo erectus* in the American Museum of Natural History. What they are comprised of is not listed.
	+ FM 1322 A: A specimen listed as from Cheyenne River, South Dakota. However, this species is unknown from the Americas and may be mistaken.
	+ FM 8032: An unidentified specimen with no location listed.

* OH 9 "Chellean Man": First named "*H. leakeyi*" but reestablished as "*H. louisleakeyi*" due to a conditional proposition, this specimen is often called *H. erectus* or *Homo ergaster*; it was also called "*H. e. olduvaiensis*" by Philip Tobias, but this has not been used. The frontal squama is fformer and the the supratoral shelf is broad and not as steep. No midline keeling is present, unlike *H. erectus*. The occipital bun is 18.5 millimeters thick, with the skull having the largest cranial capacity, which indicates a development in cognitive abilities. The robust brow and skull thickness confirm the hypothesis that *H. erectus* continued to change over time. Oldowan tools found near are assumed to be from this individual.
* "Chapala Man" or "arcada": The holotype was recovered from an unknown locality, likely when a drought revealed the beds of Lake Chapala from 1947 to 1956. Archeologists have previously focused on early Indigenous settlements, but Chapala has little been investigated, as it bears fossils from a time when early humans lived in the area. Previously, remains of 12 horses with their skull smashed between the eyes is thought to be evidence of hominin activity. Additionally, swamp deer teeth found with engravings and strung to create necklaces have been found in the same area. The horses were 10,000-20,000 years old, and the swamp deer about 20,000 years old. However, at some point, a fossil brow ridge and incomplete mandible appeared in the collections of Frederico Solorzano. When he took to a conference to discuss *Homo* in the Americas, specialists were skeptical of the find. When Solorzano examined the fossil against anatomically-derived humans, the material did not match, but against *Homo erectus* they bore a resemblance. He compared them with the Tautavel Man. However, the species died out many thousands of years before *Homo* is believed to cross into the Americas, and no evidence supports otherwise. Because of the Chapala Man, archeologists now propose fauna to have crossed into the Americas from Australia and Asia, rather than the previous Bering Strait hypothesis. Mineralized darkened spots on the holotype lead scientists to believe the Chapala Man was quite old, possibly related to *H. sapiens* and *H. erectus*. Some sources place the Chapala man as basal *Homo sapiens*, but the likely story is that Solorzano planted *Homo erectus* remains in a bone pile at Chapala and "discovered" it since little stratigraphic information is known and it was found sitting on the lake's shore rather than deposited.
* Trinil 2 "Java Man": The basis material of "*H. e. erectus*", which is disused, and the holotype of the species. It was discovered by Dubois, who believed it was a giant gibbon and an intermediary form of ape and humans. In 2022, Java asked for the Trinil fossil to be repatriated from Naturalis as well as other artifacts from different eras; including the Dubois collection of which includes the Trinil skullcap. A spokesperson at Naturalis originally expressed concern for proper housing in Indonesia, which caught criticism. Naturalis heeded order and repatriated the fossil.
* Mojokerto or Perning 1 "Mojokerto child": A skullcap found in February 1936 by an Indonesian paleontologist named Andojo, Tjokrohandojo or Andoyo with a team led by Ralph von Koenigswald. Andojo thought it was an orangutan, with von Koenigswald naming it *Pithecanthropus modjokertensis*, mispelling "Mojokerto". Eugène Dubois then contacted him, suggesting a shift to *Homo*, which he did. Now, it is considered a *Homo erectus*. It first had an enigmatic date and was claimed un-dateable. However, it was found to have an earliest date of 1.49 million years.
* BOU-VP-2/66 "Daka calvaria/skull" or "Daka": The Daka skull was discovered from the Dakanihylo Member of the Bouri Formation, Middle Awash region, Ethiopia by Henry Gilbert in 1997. This area is rich in hominin fossils from 0.5 million years ago to over 6 million years ago. It bears resemblance to Asian skulls. It represents either a very late ***Homo erectus*** or a very early *Homo rhodesiensis*. The specimen has a cranial capacity of 995, an occipital (10.6 millimeters) and temporal (10.2 millimeters) thickness, a calvarial height above g-l (51 millimeters), g-op (73 millimeters) and n-op (80 millimeters), an endocranial capacity of 986 and strong flexion. Daka is slightly more damaged on the right and a relatively large space is found on the posterior of the mastoid near the Asterion, an extensively damaged. The temporals are not as inflated as apes and the temporal squamae are about 5.7 millimeters thick, centrally. The posterior temporal is about 10.6 millimeters thick adjacent to the occipitomastoid.

## Gallery

ER3733 face reconstruction vector.pngFacial reconstruction of ER 3733.
**Credit: Clumsystiggy.**Buia face reconstruction.pngFacial reconstruction of Buia.
**Credit: Clumsystiggy.**
## References

* https://www.sciencedirect.com/science/article/abs/pii/S0047248421001391
* https://pubmed.ncbi.nlm.nih.gov/19149203/
* https://en.wikipedia.org/wiki/Homo\_erectus
* https://en.wikipedia.org/wiki/Homo\_erectus
* https://en.wikipedia.org/wiki/Turkana\_Boy
* https://popular-archaeology.com/article/the-remarkable-skulls-of-drimolen/
* https://twitter.com/Ozarchaeomaglab/status/1448854679593488387
* https://www.nhm.ac.uk/discover/news/2022/january/human-ancestor-homo-erectus-probably-wasnt-carnivore-we-thought.html
* https://www.science.org/doi/abs/10.1126/science.aaw7293
* https://books.google.ca/books?id=440TmWXToLAC&dq=%22proanthropus%22&source=gbs\_navlinks\_s

1. ↑ https://en.wikipedia.org/wiki/Java\_Man
2. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 https://en.wikipedia.org/wiki/Dmanisi\_hominins
3. ↑ https://www.reuters.com/lifestyle/oddly-enough/georgian-archaeologists-find-18-million-year-old-human-tooth-2022-09-09/
4. ↑ https://en.wikipedia.org/wiki/Java\_Man
5. ↑ https://www.youtube.com/watch?v=CazsHKnxmHQ&list=WL&index=2&ab\_channel=UniversityofCaliforniaTelevision%28UCTV%29
6. ↑ https://en.wikipedia.org/wiki/Dmanisi\_hominins
7. ↑ https://en.wikipedia.org/wiki/Dmanisi\_hominins
8. ↑ https://www.marilia.unesp.br/Home/Pos-Graduacao/Filosofia/Dissertacoes/costa\_obr\_me\_mar.pdf
9. ↑ https://www.inverse.com/science/super-archaics-humans-who-may-have-mated-with-your-ancestors
10. ↑ https://www.earth.com/news/human-ancestor-neanderthals-denisovans/
11. ↑ https://www.sciencenews.org/article/earliest-known-hominid-interbreeding-occurred-700000-years-ago
12. ↑ https://en.wikipedia.org/wiki/Homo\_erectus
13. ↑ https://pubmed.ncbi.nlm.nih.gov/20633924/
14. ↑ https://www.science.org/doi/10.1126/sciadv.ade7165#.ZFKqUVoNsgU.twitter
15. ↑ https://en.wikipedia.org/wiki/Olorgesailie
16. ↑ https://en.wikipedia.org/wiki/Homo\_erectus
17. ↑ https://www.researchgate.net/publication/358797310\_Terra\_Amata\_Does\_the\_Oldest\_Paleolithic\_Building\_Site\_Indicate\_the\_Use\_of\_Advanced\_Basket\_Weaving\_Technology
18. ↑ https://en.wikipedia.org/wiki/Homo\_erectus
19. ↑ https://ro.uow.edu.au/cgi/viewcontent.cgi?article=6441&context=smhpapers
20. ↑ https://paleoanthropology.org/ojs/index.php/paleo/article/view/1022/985
21. ↑ https://www.ivran.ru/issledovaniya-kamennogo-veka-sokotry
22. ↑ http://old.ivran.ru/expeds/76 (link discontinued; archived text here)
23. ↑ https://www.sciencedirect.com/science/article/abs/pii/S0047248422000409
24. ↑ https://en.wikipedia.org/wiki/Java\_Man
25. ↑ https://australian.museum/learn/science/human-evolution/homo-erectus/
26. ↑ https://ccschmitt.github.io/Arago%2021%20(Tautavel%20Man).html
27. ↑ http://www.china.org.cn/e-gudai/1.htm
28. ↑ https://www.researchgate.net/publication/222412926\_The\_earliest\_settlement\_of\_Germany\_Is\_there\_anything\_out\_there
29. ↑ https://www.britannica.com/topic/Lantian-man
30. ↑ https://studylib.net/doc/7690604/europanthropus-heidelbergensis
31. ↑ https://www.researchgate.net/publication/222412926\_The\_earliest\_settlement\_of\_Germany\_Is\_there\_anything\_out\_there
32. ↑ https://onlinelibrary.wiley.com/doi/10.1002/ajpa.24330
33. ↑ http://news.bbc.co.uk/2/hi/science/nature/3512470.stm
34. ↑ https://nutcrackerman.com/2022/02/06/homininos-corpulentos-corredor-levantino/
35. ↑ https://arstechnica.com/science/2022/02/1-5-million-year-old-vertebra-hints-at-a-story-of-early-human-migration/
36. ↑ https://www.nature.com/articles/s41598-022-05712-y
37. ↑ http://research.amnh.org/paleontology/search.php?action=detail&specimen\_id=167340
38. ↑ http://research.amnh.org/paleontology/search.php?action=detail&specimen\_id=112409
39. ↑ https://en.wikipedia.org/wiki/Olduvai\_Hominid\_9
40. ↑ http://patagoniamonsters.blogspot.com/2019/12/more-on-erectus-calvaria-from-chapala.html
41. ↑ http://patagoniamonsters.blogspot.com/2019/12/chapala-homo-erectus-brow-ridge.html
42. ↑ https://www.latimes.com/archives/la-xpm-2004-oct-03-adfg-bones3-story.html
43. ↑ http://patagoniamonsters.blogspot.com/2011/01/homo-erectus-in-america-evidence.html
44. ↑ https://en.wikipedia.org/wiki/Java\_Man
45. ↑ https://www.dutchnews.nl/news/2022/10/indonesia-wants-java-man-back-committee-to-investigate-claims/
46. ↑ https://www.nytimes.com/2022/11/09/arts/design/naturalis-museum-java-man-indonesia.html
47. ↑ https://en.wikipedia.org/wiki/Mojokerto\_child
48. ↑ https://en.wikipedia.org/wiki/Daka\_skull
49. ↑ https://www.sciencedirect.com/science/article/abs/pii/S0047248415002754?via%3Dihub