Homo ergaster
Homo ergaster is an extinct species or subspecies of archaic humans who lived in Africa in the Early Pleistocene. Whether H. ergaster constitutes a species of its own or should be subsumed into H. erectus is an ongoing and unresolved dispute within palaeoanthropology. Proponents of synonymisation typically designate H. ergaster as "African Homo erectus" or "Homo erectus ergaster". The name Homo ergaster roughly translates to "working man", a reference to the more advanced tools used by the species in comparison to those of their ancestors. The fossil range of H. ergaster mainly covers the period of 1.7 to 1.4 million years ago, though a broader time range is possible. Though fossils are known from across East and Southern Africa, most H. ergaster fossils have been found along the shores of Lake Turkana in Kenya. There are later African fossils, some younger than 1 million years ago, that indicate long-term anatomical continuity, though it is unclear if they can be formally regarded as H. ergaster specimens. As a chronospecies, H. ergaster may have persisted to as late as 600,000 years ago, when new lineages of Homo arose in Africa.
Those who believe H. ergaster should be subsumed into H. erectus consider there to be too little difference between the two to separate them into distinct species. Proponents of keeping the two species as distinct cite morphological differences between the African fossils and H. erectus fossils from Asia, as well as early Homo evolution being more complex than what is implied by subsuming species such as H. ergaster into H. erectus. Additionally, morphological differences between the specimens commonly seen as constituting H. ergaster might suggest that H. ergaster itself does not represent a cohesive species. Regardless of their most correct classification, H. ergaster exhibit primitive versions of traits later expressed in H. erectus and are thus likely the direct ancestors of later H. erectus populations in Asia. Additionally, H. ergaster is likely ancestral to later hominins in Europe and Africa, such as modern humans and Neanderthals.
Several features distinguish H. ergaster from australopithecines as well as earlier and more basal species of Homo, such as H. habilis. Among these features are their larger body mass, relatively long legs, obligate bipedalism, relatively small jaws and teeth as well as body proportions and inferred lifestyles more similar to modern humans than to earlier and contemporary hominins. With these features in mind, some researchers view H. ergaster as being the earliest true representative of the genus Homo.
H. ergaster lived on the savannah in Africa, a unique environment with challenges that would have resulted in the need for many new and distinct behaviours. Earlier Homo probably used counter-attack tactics, like modern primates, to keep predators away. By the time of H. ergaster, this behaviour had probably resulted in the development of true hunter-gatherer behaviour, a first among primates. H. ergaster was an apex predator. Further behaviours that might first have arisen in H. ergaster include male-female divisions of foraging and true monogamous pair bonds. H. ergaster also marks the appearance of more advanced tools of the Acheulean industry, including the earliest-known hand axes. Though undisputed evidence is missing, H. ergaster might also have been the earliest hominin to master control of fire.
Taxonomy
Research history
The systematics and taxonomy of Homo in the Early to Middle Pleistocene is one of the most disputed areas of palaeoanthropology. In early palaeoanthropology and well into the twentieth century, it was generally assumed that Homo sapiens was the end result of gradual modifications within a single lineage of hominin evolution. As the perceived transitional form between early hominins and modern humans, H. erectus, originally assigned to contain archaic human fossils in Asia, came to encompass a wide range of fossils covering a large span of time. Since the late twentieth century, the diversity within H. erectus has led some to question what exactly defines the species and what it should encompass. Some researchers, such as palaeoanthropologist Ian Tattersall in 2013, have questioned H. erectus since it contains an "unwieldly" number of fossils with "substantially differing morphologies".In the 1970s, palaeoanthropologists Richard Leakey and Alan Walker described a series of hominin fossils from Kenyan fossil localities on the eastern shore of Lake Turkana. The most notable finds were two partial skulls; KNM ER 3733 and KNM ER 3883, found at Koobi Fora. Leakey and Walker assigned these skulls to H. erectus, noting that their brain volumes compared well to the far younger type specimen of H. erectus. Another significant fossil was a fossil mandible recovered at Ileret and described by Leakey with the designation KNM ER 992 in 1972 as "Homo of indeterminate species".
In 1975, palaeoanthropologists Colin Groves and Vratislav Mazák designated KNM ER 992 as the holotype specimen of a distinct species, which they dubbed Homo ergaster. The name roughly translates to "working man" or "workman". Groves and Mazák also included many of the Koobi Fora fossils, such as KNM ER 803 in their designation of the species, but did not provide any comparison with the Asian fossil record of H. erectus in their diagnosis, inadvertently causing some of the later taxonomic confusion in regards to the species.
A nearly complete fossil, interpreted as a young male, was discovered at the western shore of Lake Turkana in 1984 by Kenyan archaeologist Kamoya Kimeu. The fossils were described by Leakey and Walker, alongside paleoanthropologists Frank Brown and John Harris, in 1985 as KNM WT 15000. They interpreted the fossil, consisting of a nearly complete skeleton, as representing H. erectus. Turkana Boy was the first discovered comprehensively preserved specimen of H. ergaster/''erectus found and constitutes an important fossil in establishing the differences and similarities between early Homo and modern humans. Turkana Boy was placed in H. ergaster by paleoanthropologist Bernard Wood in 1992, and is today, alongside other fossils in Africa previously designated as H. erectus, commonly seen as a representative of H. ergaster by those who support H. ergaster'' as a distinct species.
Classification
H. ergaster is easily distinguished from earlier and more basal species of Homo, notably H. habilis and H. rudolfensis, by a number of features that align them, and their inferred lifestyle, more closely to modern humans than to earlier and contemporary hominins. As compared to their relatives, H. ergaster had body proportions more similar to later members of the genus Homo, notably relatively long legs which would have made them obligately bipedal. The teeth and jaws of H. ergaster are also relatively smaller than those of H. habilis and H. rudolfensis, indicating a major change in diet. In 1999, palaeoanthropologists Bernard Wood and Mark Collard argued that the conventional criteria for assigning species to the genus Homo were flawed and that early and basal species, such as H. habilis and H. rudolfensis, might appropriately be reclassified as ancestral australopithecines. In their view, the true earliest representative of Homo was H. ergaster.Since its description as a separate species in 1975, the classification of the fossils referred to H. ergaster has been in dispute. H. ergaster was immediately dismissed by Leakey and Walker and many influential researchers, such as palaeoanthropologist G. Philip Rightmire, who wrote an extensive treatise on H. erectus in 1990, continued to prefer a more inclusive and comprehensive H. erectus. Overall, there is no doubt that the group of fossils composing H. erectus and H. ergaster represent the fossils of a more or less cohesive subset of closely related archaic humans. The question is instead whether these fossils represent a radiation of different species or the radiation of a single, highly variable and diverse, species over the course of almost two million years. This long-running debate remains unresolved, with researchers typically using the terms H. erectus s.s. to refer to H. erectus fossils in Asia and the term H. erectus s.l. to refer to fossils of other species that may or may not be included in H. erectus, such as H. ergaster, H. antecessor and H. heidelbergensis.
For obvious reasons, H. ergaster shares many features with H. erectus, such as large forward-projecting jaws, large brow ridges and a receding forehead. Many of the features of H. ergaster are clearly more primitive versions of features later expressed in H. erectus, which somewhat obscures the differences between the two. There are subtle, potentially significant, differences between the East African and East Asian fossils. Among these are the somewhat higher-domed and thinner-walled skulls of H. ergaster, and the even more massive brow ridges and faces of Asian H. erectus.
The question is made more difficult since it regards how much intraspecific variation can be exhibited in a single species before it needs to be split into more, a question that in and of itself does not have a clear-cut answer. A 2008 analysis by anthropologist Karen L. Baab, examining fossils of various H. erectus subspecies, and including fossils attributed to H. ergaster, found that the intraspecific variation within H. erectus was greater than expected for a single species when compared to modern humans and chimpanzees, but fell well within the variation expected for a species when compared to gorillas, and even well within the range expected for a single subspecies when compared to orangutans. Baab concluded that H. erectus s.l. was either a single but variable species, several subspecies divided by time and geography or several geographically dispersed but closely related species. In 2015, paleoanthropologists David Strait, Frederick Grine and John Fleagle listed H. ergaster as one of the seven "widely recognized" species of Homo, alongside H. habilis, H. rudolfensis, H. erectus, H. heidelbergensis, H. neanderthalensis and H. sapiens, noting that other species, such as H. floresiensis and H. antecessor, were less widely recognised or more poorly known.