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''Paranthropus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of extinct
hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
which contains two widely accepted
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
: '' P. robustus'' and ''
P. boisei ''Paranthropus boisei'' is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959, and species descrip ...
''. However, the validity of ''Paranthropus'' is contested, and it is sometimes considered to be
synonymous A synonym is a word, morpheme, or phrase that means exactly or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are all ...
with ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
''. They are also referred to as the robust australopithecines. They lived between approximately 2.6 and 1.2 million years ago (mya) from the end of the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...
. ''Paranthropus'' is characterised by
robust Robustness is the property of being strong and healthy in constitution. When it is transposed into a system, it refers to the ability of tolerating perturbations that might affect the system’s functional body. In the same line ''robustness'' ca ...
skulls The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
, with a prominent
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
-like
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
along the midline—which suggest strong chewing muscles—and broad,
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
teeth used for grinding. However, they likely preferred soft food over tough and hard food. ''Paranthropus'' species were generalist feeders, but ''P. robustus'' was likely an
omnivore An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nutr ...
, whereas ''P. boisei'' was likely herbivorous and mainly ate bulbotubers. They were
bipeds Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
. Despite their robust heads, they had comparatively small bodies. Average weight and height are estimated to be at for ''P. robustus'' males, at for ''P. boisei'' males, at for ''P. robustus'' females, and at for ''P. boisei'' females. They were possibly
polygamous Crimes Polygamy (from Late Greek (') "state of marriage to many spouses") is the practice of marriage, marrying multiple spouses. When a man is married to more than one wife at the same time, sociologists call this polygyny. When a woman is ...
and
patrilocal In social anthropology, patrilocal residence or patrilocality, also known as virilocal residence or virilocality, are terms referring to the social system in which a married couple resides with or near the husband's parents. The concept of locat ...
, but there are no modern analogues for australopithecine societies. They are associated with bone tools and contested as the earliest evidence of fire usage. They typically inhabited woodlands, and coexisted with some early human species, namely '' A. africanus'', ''
H. habilis ''Homo habilis'' ("handy man") is an extinct species of archaic human from the Early Pleistocene of East and South Africa about 2.31 million years ago to 1.65 million years ago (mya). Upon species description in 1964, ''H. habilis'' was highly c ...
'' and ''
H. erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor' ...
''. They were preyed upon by the large carnivores of the time, specifically crocodiles, leopards,
sabertoothed cat Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...
s and hyenas.


Taxonomy


Species


''P. robustus''

The genus ''Paranthropus'' was first erected by Scottish-South African
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Robert Broom Robert Broom FRS FRSE (30 November 1866 6 April 1951) was a British- South African doctor and palaeontologist. He qualified as a medical practitioner in 1895 and received his DSc in 1905 from the University of Glasgow. From 1903 to 1910, he ...
in 1938, with the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
'' P. robustus''. "''Paranthropus''" derives from
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...
παρα ''para'' beside or alongside; and άνθρωπος ''ánthropos'' man. The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
, a male braincase, TM 1517, was discovered by schoolboy Gert Terblanche at the
Kromdraai fossil site Kromdraai (means crooked turn in afrikaans) is a fossil-bearing breccia-filled cave located about east of the well-known South African hominid-bearing site of Sterkfontein and about northwest of Johannesburg, South Africa. It is situated withi ...
, about southwest of
Pretoria Pretoria () is South Africa's administrative capital, serving as the seat of the Executive (government), executive branch of government, and as the host to all foreign embassies to South Africa. Pretoria straddles the Apies River and extends ...
, South Africa. By 1988, at least six individuals were unearthed in around the same area, now known as the Cradle of Humankind. In 1948, at
Swartkrans Swartkrans is a fossil-bearing cave designated as a South African National Heritage Site, located about from Johannesburg. It is located in the Cradle of Humankind World Heritage Site and is notable for being extremely rich in archaeological m ...
Cave, in about the same vicinity as Kromdraai, Broom and South African palaeontologist
John Talbot Robinson John Talbot Robinson FRSSAf (10 January 1923 – 12 October 2001) was a distinguished South African hominin paleontologist. His most famous discovery (with Robert Broom) was the nearly complete fossil skull of the hominin species '' Australop ...
described ''P. crassidens'' based on a subadult jaw, SK 6. He believed later ''Paranthropus'' were morphologically distinct from earlier ''Paranthropus'' in the cave—that is, the Swartkrans ''Paranthropus'' were
reproductively isolated The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offsprin ...
from Kromdraai ''Paranthropus'' and the former eventually
speciate Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
d. By 1988, several specimens from Swartkrans had been placed into ''P. crassidens''. However, this has since been synonymised with ''P. robustus'' as the two populations do not seem to be very distinct.


''P. boisei''

In 1959, ''
P. boisei ''Paranthropus boisei'' is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959, and species descrip ...
'' was discovered by
Mary Leakey Mary Douglas Leakey, FBA (née Nicol, 6 February 1913 – 9 December 1996) was a British paleoanthropologist who discovered the first fossilised ''Proconsul A proconsul was an official of ancient Rome who acted on behalf of a consul. A pro ...
at
Olduvai Gorge The Olduvai Gorge or Oldupai Gorge in Tanzania is one of the most important paleoanthropology, paleoanthropological localities in the world; the many sites exposed by the gorge have proven invaluable in furthering understanding of early human ev ...
,
Tanzania Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
(specimen
OH 5 ''Paranthropus boisei'' is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959, and described by h ...
). Her husband
Louis Louis may refer to: * Louis (coin) * Louis (given name), origin and several individuals with this name * Louis (surname) * Louis (singer), Serbian singer * HMS ''Louis'', two ships of the Royal Navy See also Derived or associated terms * Lewis ( ...
named it ''Zinjanthropus boisei'' because he believed it differed greatly from ''Paranthropus'' and ''Australopithecus''. The name derives from " Zinj", an ancient Arabic word for the coast of East Africa, and "boisei", referring to their financial benefactor
Charles Watson Boise Charles Watson Boise (November 9, 1884 – November 15, 1964) was an American-born naturalised British mining engineer. Early life Born in Lakota, North Dakota on November 9, 1884, his family soon moved to Hope where he spent his formative years. ...
. However, this genus was rejected at Mr. Leakey's presentation before the 4th Pan-African Congress on Prehistory, as it was based on a single specimen. The discovery of the
Peninj Mandible The Peninj Mandible(Peninj 1), also called Natron mandible, is the fossilized lower jaw and teeth of an australopithecine specimen, likely that of '' Australopithecus boisei'' or a similar population. It was discovered in West Lake Natron, in Ngor ...
made the Leakey's reclassify their species as ''Australopithecus (Zinjanthropus) boisei'' in 1964, but in 1967, South African palaeoanthropologist
Phillip V. Tobias Phillip Vallentine Tobias (14 October 1925 – 7 June 2012) was a South African palaeoanthropologist and Professor Emeritus at the University of the Witwatersrand in Johannesburg. He was best known for his work at South Africa's hominid fossil ...
subsumed it into ''Australopithecus'' as ''A. boisei''. However, as more specimens were found, the combination ''Paranthropus boisei'' became more popular. It is debated whether the wide range of variation in jaw size indicates simply
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
or a grounds for identifying a new species. It could be explained as
groundmass The matrix or groundmass of a rock is the finer-grained mass of material in which larger grains, crystals, or clasts are embedded. The matrix of an igneous rock consists of finer-grained, often microscopic, crystals in which larger crystals, ca ...
filling in cracks naturally formed after death, inflating the perceived size of the bone. ''P. boisei'' also has a notably wide range of variation in skull anatomy, but these features likely have no taxonomic bearing.


''P. aethiopicus''

In 1968, French palaeontologists
Camille Arambourg Camille Arambourg ( February 3, 1885– November 19, 1969) was a French vertebrate paleontologist. He conducted extensive field work in North Africa. In the 1950s he argued against the prevailing model of Neanderthals as brutish and simian. Du ...
and
Yves Coppens Yves Coppens (9 August 1934 – 22 June 2022) was a French anthropologist. A graduate from the University of Rennes and Sorbonne, he studied ancient hominids and had multiple published works on this topic, and also produced a film. In October 2 ...
described "''Paraustralopithecus aethiopicus''" based on a toothless mandible from the
Shungura Formation The Shungura Formation is a stratigraphic formation located in the Omo river basin in Ethiopia. It dates to the Late Pliocene to Early Pleistocene. Oldowan tools have been found in the formation, suggesting early use of stone tools by hominins. A ...
, Ethiopia (Omo 18). In 1976, American anthropologist
Francis Clark Howell Francis Clark Howell (November 27, 1925 – March 10, 2007), generally known as F. Clark Howell, was an American anthropologist. Born in Kansas City, Missouri, F. Clark Howell grew up in Kansas, where he became interested in natural history. H ...
and Breton anthropologist
Yves Coppens Yves Coppens (9 August 1934 – 22 June 2022) was a French anthropologist. A graduate from the University of Rennes and Sorbonne, he studied ancient hominids and had multiple published works on this topic, and also produced a film. In October 2 ...
reclassified it as ''A. africanus''. In 1986, after the discovery of the skull
KNM WT 17000 KNM-WT 17000 (also known as "The Black Skull") is a fossilized adult skull of the species ''Paranthropus aethiopicus''. It was discovered in West Turkana, Kenya by Alan Walker in 1985. Estimated to be 2.5 million years old, it is an adult with ...
by English anthropologist
Alan Walker Alan Olav Walker (born 24 August 1997) is a British-born Norwegian music producer and DJ primarily known for the critically acclaimed single " Faded" (2015), which was certified platinum in 14 countries. He has also made several songs including ...
and
Richard Leakey Richard Erskine Frere Leakey (19 December 1944 – 2 January 2022) was a Kenyan paleoanthropologist, conservationist and politician. Leakey held a number of official positions in Kenya, mostly in institutions of archaeology and wildlife cons ...
classified it into ''Paranthropus'' as '' P. aethiopicus''. There is debate whether this is synonymous with ''P. boisei'', the main argument for separation being the skull seems less adapted for chewing tough vegetation. In 1989, palaeoartist and zoologist Walter Ferguson reclassified KNM WT 17000 into a new species, ''walkeri'', because he considered the skull's species designation questionable as it comprised the skull whereas the holotype of ''P. aethiopicus'' comprised only the mandible. Ferguson's classification is almost universally ignored, and is considered to be synonymous with ''P. aethiopicus''.


Others

In 1963, while in the Congo, French ethnographer Charles Cordier assigned the name "P. congensis" to a super-strong, monstrous
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its sister g ...
-man cryptid called "Kikomba", "Apamándi", "Abanaánji", "Zuluzúgu", or "Tshingómbe" by various native tribes which he heard stories about. In 2015, Ethiopian palaeoanthropologist
Yohannes Haile-Selassie Yohannes Haile-Selassie Ambaye (born 23 February 1961) is an Ethiopian paleoanthropologist. An authority on pre-''Homo sapiens'' hominids, he particularly focuses his attention on the East African Rift and Middle Awash valleys.Mangels, John (20 ...
and colleagues described the 3.5–3.2 Ma '' A. deyiremeda'' based on three jawbones from the
Afar Region The Afar Region (; aa, Qafar Rakaakayak; am, አፋር ክልል), formerly known as Region 2, is a regional state in northeastern Ethiopia and the homeland of the Afar people. Its capital is the planned city of Semera, which lies on the paved ...
, Ethiopia. They noted that, though it shares many similarities with ''Paranthropus'', it may not have been closely related because it lacked enlarged molars which characterize the genus. Nonetheless, in 2018, independent researcher Johan Nygren recommended moving it to ''Paranthropus'' based on dental and presumed dietary similarity.


Validity

In 1951, American anthropologists
Sherwood Washburn Sherwood Larned Washburn ( – ), nicknamed "Sherry", was an American physical anthropologist, and "a legend in the field." He was pioneer in the field of primatology, opening it to the study of primates in their natural habitats. His research ...
and Bruce D. Patterson were the first to suggest that ''Paranthropus'' should be considered a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...
of ''Australopithecus'' as the former was only known from fragmentary remains at the time, and dental differences were too minute to serve as justification. In face of calls for subsumation, Leakey and Robinson continued defending its validity. Various other authors were still unsure until more complete remains were found. ''Paranthropus'' is sometimes classified as a
subgenus In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
of ''Australopithecus''. There is currently no clear consensus on the validity of ''Paranthropus''. The argument rests upon whether the genus is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
—is composed of a common ancestor and all of its descendants—and the argument against monophyly (that the genus is
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
) says that ''P. robustus'' and ''P. boisei'' evolved similar gorilla-like heads independently of each other by coincidence (
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
), as chewing adaptations in hominins evolve very rapidly and multiple times at various points in the family tree (
homoplasy Homoplasy, in biology and phylogenetics, is the term used to describe a Phenotypic trait, feature that has been gained or lost independently in separate lineages over the course of evolution. This is different from Homology (biology), homology, w ...
). In 1999, a chimp-like
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
forearm bone was assigned to ''P. boisei'', the first discovered ulna of the species, which was markedly different from ''P. robustus'' ulnae, which could suggest paraphyly.


Evolution

''P. aethiopicus'' is the earliest member of the genus, with the oldest remains, from the Ethiopian
Omo Kibish Formation The Omo Kibish Formation or simply Kibish Formation is a geological formation in the Lower Omo Valley of southwestern Ethiopia. It is named after the nearby Omo River and is subdivided into four members known as Members I-IV. The members are numbe ...
, dated to 2.6 mya at the end of the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Malema Malema is a town in the district of Malema. It is situated within the province of Nampula in Mozambique. The district borders the districts of: Ribaue, Lalaua, Alto Molocue, Gurue, Cuamba, and others. The district is dependent on agricultur ...
, Malawi. ''P. boisei'' changed remarkably little over its nearly one-million-year existence. ''Paranthropus'' had spread into South Africa by 2 mya with the earliest ''P. robustus'' remains. It is sometimes suggested that ''Paranthropus'' and ''Homo'' are
sister taxa In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
, both evolving from ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
''. This may have occurred during a drying trend 2.8–2.5 mya in the
Great Rift Valley The Great Rift Valley is a series of contiguous geographic trenches, approximately in total length, that runs from Lebanon in Asia to Mozambique in Southeast Africa. While the name continues in some usages, it is rarely used in geology as it i ...
, which caused the retreat of woodland environments in favor of open savanna, with forests growing only along rivers and lakes. ''Homo'' evolved in the former, and ''Paranthropus'' in the latter
riparian A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the terrestrial biomes of the Earth. Plant habitats and communities along the river margins and banks ar ...
environment. However, the classifications of ''Australopithecus'' species is problematic.
Evolutionary tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
according to a 2019 study:


Description


Skull

''Paranthropus'' had a massively built, tall and flat skull, with a prominent
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
-like
sagittal crest A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are exceptiona ...
along the midline which anchored massive
temporalis muscle In anatomy, the temporalis muscle, also known as the temporal muscle, is one of the muscles of mastication (chewing). It is a broad, fan-shaped convergent muscle on each side of the head that fills the temporal fossa, superior to the zygomati ...
s used in chewing. Like other australopithecines, ''Paranthropus'' exhibited sexual dimorphism, with males notably larger than females. They had large
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
with a relatively thick
tooth enamel Tooth enamel is one of the four major Tissue (biology), tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the Crown (tooth), crown. The ...
coating (
post-canine megadontia Post-canine megadontia is a relative enlargement of the molars and premolars compared to the size of the incisors and canines. This phenomenon is seen in some early hominid ancestors such as ''Paranthropus aethiopicus.'' Archaeological evidence ...
), and comparatively small
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s (similar in size to modern
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
s), possibly adaptations to processing abrasive foods. The teeth of ''P. aethiopicus'' developed faster than those of ''P. boisei''. ''Paranthropus'' had adaptations to the skull to resist large bite loads while feeding, namely the expansive
squamosal suture The squamosal suture, or squamous suture, arches backward from the pterion and connects the temporal squama with the lower border of the parietal bone: this suture is continuous behind with the short, nearly horizontal parietomastoid suture, whic ...
s. The notably thick
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sepa ...
was once thought to have been an adaptation to resist a high bite force, but is better explained as a byproduct of facial lengthening and nasal anatomy. In ''P. boisei'', the jaw hinge was adapted to grinding food side-to-side (rather than up-and-down in modern humans), which is better at processing the
starch Starch or amylum is a polymeric carbohydrate consisting of numerous glucose units joined by glycosidic bonds. This polysaccharide is produced by most green plants for energy storage. Worldwide, it is the most common carbohydrate in human diets ...
y abrasive foods that likely made up the bulk of its diet. ''P. robustus'' may have chewed in a front-to-back direction instead, and had less exaggerated (less
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
) anatomical features than ''P. boisei'' as it perhaps did not require them with this kind of chewing strategy. This may have also allowed ''P. robustus'' to better process tougher foods. The braincase volume averaged about , comparable to gracile australopithecines, but smaller than ''Homo''. Modern human brain volume averages for men and for women.


Limbs and locomotion

Unlike ''P. robustus'', the forearms of ''P. boisei'' were heavily built, which might suggest habitual
suspensory behaviour Suspensory behaviour is a form of arboreal locomotion or a feeding behavior that involves hanging or suspension of the body below or among tree branches. This behavior enables faster travel while reducing path lengths to cover more ground when trave ...
as in
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s and
gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India ...
s. A ''P. boisei''
shoulder blade The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either ...
indicates long
infraspinatus muscle In human anatomy, the infraspinatus muscle is a thick triangular muscle, which occupies the chief part of the infraspinatous fossa.'' Gray's Anatomy'', see infobox. As one of the four muscles of the rotator cuff, the main function of the infraspi ...
s, which is also associated with suspensory behavior. A ''P. aethiopicus'' ulna, on the other hand, shows more similarities to ''Homo'' than ''P. boisei''. ''Paranthropus'' were
biped Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' a ...
s, and their hips, legs and feet resemble ''A. afarensis'' and modern humans. The pelvis is similar to ''A. afarensis'', but the hip joints are smaller in ''P. robustus''. The physical similarity implies a similar walking gait. Their modern-humanlike big toe indicates a modern-humanlike foot posture and range of motion, but the more distal ankle joint would have inhibited the modern human toe-off gait cycle. By 1.8 mya, ''Paranthropus'' and ''
H. habilis ''Homo habilis'' ("handy man") is an extinct species of archaic human from the Early Pleistocene of East and South Africa about 2.31 million years ago to 1.65 million years ago (mya). Upon species description in 1964, ''H. habilis'' was highly c ...
'' may have achieved about the same grade of bipedality.


Height and weight

In comparison to the large, robust head, the body was rather small. Average weight for ''P. robustus'' may have been for males and for females; and for ''P. boisei'' for males and for females. At Swartkrans Cave Members 1 and 2, about 35% of the ''P. robustus'' individuals are estimated to have weighed , 22% about , and the remaining 43% bigger than the former but less than . At Member 3, all individuals were about . Female weight was about the same in contemporaneous ''H. erectus'', but male ''H. erectus'' were on average heavier than ''P. robustus'' males. ''P. robustus'' sites are oddly dominated by small adults, which could be explained as heightened predation or mortality of the larger males of a group. The largest-known ''Paranthropus'' individual was estimated at . According to a 1991 study, based on
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
length and using the dimensions of modern humans, male and female ''P. robustus'' are estimated to have stood on average , respectively, and ''P. boisei'' . However, the latter estimates are problematic as there were no positively identified male ''P. boisei'' femurs at the time. In 2013, a 1.34 Ma male ''P. boisei'' partial skeleton was estimated to be at least and .


Pathology

''Paranthropus'' seems to have had notably high rates of pitting enamel hypoplasia (PEH), where
tooth enamel Tooth enamel is one of the four major Tissue (biology), tissues that make up the tooth in humans and many other animals, including some species of fish. It makes up the normally visible part of the tooth, covering the Crown (tooth), crown. The ...
formation is spotty instead of mostly uniform. In ''P. robustus'', about 47% of
baby teeth Deciduous teeth or primary teeth, also informally known as baby teeth, milk teeth, or temporary teeth,Illustrated Dental Embryology, Histology, and Anatomy, Bath-Balogh and Fehrenbach, Elsevier, 2011, page 255 are the first set of teeth in the ...
and 14% of adult teeth were affected, in comparison to about 6.7% and 4.3%, respectively, in any other tested hominin species. The condition of these holes covering the entire tooth is consistent with the modern human ailment
amelogenesis imperfecta Amelogenesis imperfecta (AI) is a congenital disorder which presents with a rare abnormal formation of the enamel or external layer of the crown of teeth, unrelated to any systemic or generalized conditions. Enamel is composed mostly of mineral, ...
. However, since circular holes in enamel coverage are uniform in size, only present on the
molar teeth The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone t ...
, and have the same severity across individuals, the PEH may have been a genetic condition. It is possible that the coding-DNA concerned with thickening enamel also left them more vulnerable to PEH. There have been 10 identified cases of
cavities Cavity may refer to: Biology and healthcare *Body cavity, a fluid-filled space in many animals where organs typically develop **Gastrovascular cavity, the primary organ of digestion and circulation in cnidarians and flatworms *Dental cavity or too ...
in ''P. robustus'', indicating a rate similar to modern humans. A molar from
Drimolen The Drimolen Palaeocave System consists of a series of terminal Pliocene to early Pleistocene hominin-bearing palaeocave fills located around north of Johannesburg, South Africa, and about north of Sterkfontein in the UNESCO World Heritage Site ...
, South Africa, showed a cavity on the
tooth root Dental anatomy is a field of anatomy dedicated to the study of human tooth structures. The development, appearance, and classification of teeth fall within its purview. (The function of teeth as they contact one another falls elsewhere, under de ...
, a rare occurrence in fossil great apes. In order for cavity-creating bacteria to reach this area, the individual would have had to have also presented either alveolar resportion, which is commonly associated with
gum disease Periodontal disease, also known as gum disease, is a set of inflammatory conditions affecting the tissues surrounding the teeth. In its early stage, called gingivitis, the gums become swollen and red and may bleed. It is considered the main caus ...
; or super-eruption of teeth which occurs when teeth become worn down and have to erupt a bit more in order to maintain a proper bite, and this exposed the root. The latter is most likely, and the exposed root seems to have caused
hypercementosis Hypercementosis is an idiopathic, non-neoplastic condition characterized by the excessive buildup of normal cementum (calcified tissue) on the roots of one or more teeth. A thicker layer of cementum can give the tooth an enlarged appearance, which ...
to anchor the tooth in place. The cavity seems to have been healing, which may have been caused by a change in diet or mouth microbiome, or the loss of the adjacent molar.


Palaeobiology


Diet

It was once thought ''P. boisei'' cracked open nuts with its powerful teeth, giving OH 5 the nickname "Nutcracker Man". However, like gorillas, ''Paranthropus'' likely preferred soft foods, but would consume tough or hard food during leaner times, and the powerful jaws were used only in the latter situation. In ''P. boisei'', thick enamel was more likely used to resist abrasive gritty particles rather than to minimize chipping while eating hard foods. In fact, there is a distinct lack of tooth fractures which would have resulted from such activity. ''Paranthropus'' were generalist feeders, but diet seems to have ranged dramatically with location. The South African ''P. robustus'' appears to have been an omnivore, with a diet similar to contemporaneous ''Homo'' and nearly identical to the later '' H. ergaster'', and subsisted on mainly C4 savanna plants and C3 forest plants, which could indicate either seasonal shifts in diet or seasonal migration from forest to savanna. In leaner times it may have fallen back on brittle food. It likely also consumed seeds and possibly
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s or
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
s. A high cavity rate could indicate
honey Honey is a sweet and viscous substance made by several bees, the best-known of which are honey bees. Honey is made and stored to nourish bee colonies. Bees produce honey by gathering and then refining the sugary secretions of plants (primar ...
consumption. The East African ''P. boisei'', on the other hand, seems to have been largely herbivorous and fed on C4 plants. Its powerful jaws allowed it to consume a wide variety of different plants, though it may have largely preferred nutrient-rich bulbotubers as these are known to thrive in the well-watered woodlands it is thought to have inhabited. Feeding on these, ''P. boisei'' may have been able to meet its daily caloric requirements of approximately 9,700 kJ after about 6 hours of foraging. Juvenile ''P. robustus'' may have relied more on tubers than adults, given the elevated levels of
strontium Strontium is the chemical element with the symbol Sr and atomic number 38. An alkaline earth metal, strontium is a soft silver-white yellowish metallic element that is highly chemically reactive. The metal forms a dark oxide layer when it is ex ...
compared to adults in teeth from Swartkrans Cave, which, in the area, was most likely sourced from tubers.
Dentin Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
exposure on juvenile teeth could indicate early weaning, or a more abrasive diet than adults which wore away the
cementum Cementum is a specialized calcified substance covering the root of a tooth. The cementum is the part of the periodontium that attaches the teeth to the alveolar bone by anchoring the periodontal ligament.Illustrated Dental Embryology, Histology, a ...
and enamel coatings, or both. It is also possible juveniles were less capable of removing grit from dug-up food rather than purposefully seeking out more abrasive foods.


Technology

Bone tools dating between 2.3 and 0.6 mya have been found in abundance in Swartkrans, Kromdraai and
Drimolen The Drimolen Palaeocave System consists of a series of terminal Pliocene to early Pleistocene hominin-bearing palaeocave fills located around north of Johannesburg, South Africa, and about north of Sterkfontein in the UNESCO World Heritage Site ...
caves, and are often associated with ''P. robustus''. Though ''Homo'' is also known from these caves, their remains are comparatively scarce to ''Paranthropus'', making ''Homo''-attribution unlikely. The tools also cooccur with ''Homo''-associated
Oldawan The Oldowan (or Mode I) was a widespread stone tool archaeological industry (style) in prehistory. These early tools were simple, usually made with one or a few flakes chipped off with another stone. Oldowan tools were used during the Low ...
and possibly
Acheulian Acheulean (; also Acheulian and Mode II), from the French ''acheuléen'' after the type site of Saint-Acheul, is an archaeological industry of stone tool manufacture characterized by the distinctive oval and pear-shaped "hand axes" associated ...
stone tool industries. The bone tools were typically sourced from the shaft of
long bone The long bones are those that are longer than they are wide. They are one of five types of bones: long, Short bone, short, Flat bone, flat, Irregular bone, irregular and Sesamoid bone, sesamoid. Long bones, especially the femur and tibia, are subj ...
s from medium- to large-sized mammals, but tools made sourced from
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s,
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
s and horn cores have also been found. Bone tools have also been found at Oldawan Gorge and directly associated with ''P. boisei'', the youngest dating to 1.34 mya, though a great proportion of other bone tools from here have ambiguous attribution. Stone tools from Kromdraai could possibly be attributed to ''P. robustus'', as no ''Homo'' have been found there yet. The bone tools were not manufactured or purposefully shaped for a task. However, since the bones display no weathering (and were not scavenged randomly), and there is a preference displayed for certain bones, raw materials were likely specifically hand-picked. This could indicate a similar cognitive ability to contemporary Stone Age ''Homo''. Bone tools may have been used to cut or process vegetation, or dig up
tuber Tubers are a type of enlarged structure used as storage organs for nutrients in some plants. They are used for the plant's perennation (survival of the winter or dry months), to provide energy and nutrients for regrowth during the next growing ...
s or
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
s, The form of ''P. robustus'' incisors appear to be intermediate between ''H. erectus'' and modern humans, which could indicate less food processing done by the teeth due to preparation with simple tools. Burnt bones were also associated with the inhabitants of Swartkrans, which could indicate some of the earliest fire usage. However, these bones were found in Member 3, where ''Paranthropus'' remains are rarer than ''H. erectus'', and it is also possible the bones were burned in a wildfire and washed into the cave as it is known the bones were not burned onsite.


Social structure

Given the marked anatomical and physical differences with modern great apes, there may be no modern analogue for australopithecine societies, so comparisons drawn with modern primates will not be entirely accurate. ''Paranthropus'' had pronounced
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
, with males notably larger than females, which is commonly correlated with a male-dominated
polygamous Crimes Polygamy (from Late Greek (') "state of marriage to many spouses") is the practice of marriage, marrying multiple spouses. When a man is married to more than one wife at the same time, sociologists call this polygyny. When a woman is ...
society. ''P. robustus'' may have had a harem society similar to modern forest-dwelling
silverback gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
s, where one male has exclusive breeding rights to a group of females, as male-female size disparity is comparable to gorillas (based on facial dimensions), and younger males were less robust than older males (delayed maturity is also exhibited in gorillas). However, if ''P. robustus'' preferred a savanna habitat, a multi-male society would have been more productive to better defend the troop from predators in the more exposed environment, much like savanna
baboon Baboons are primates comprising the genus ''Papio'', one of the 23 genera of Old World monkeys. There are six species of baboon: the hamadryas baboon, the Guinea baboon, the olive baboon, the yellow baboon, the Kinda baboon and the chacma ba ...
s. Further, among primates, delayed maturity is also exhibited in the
rhesus monkey The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally b ...
which has a multi-male society, and may not be an accurate indicator of social structure. A 2011 strontium isotope study of ''P. robustus'' teeth from the
dolomite Dolomite may refer to: *Dolomite (mineral), a carbonate mineral *Dolomite (rock), also known as dolostone, a sedimentary carbonate rock *Dolomite, Alabama, United States, an unincorporated community *Dolomite, California, United States, an unincor ...
Sterkfontein Valley found that, like other
hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
s, but unlike other great apes, ''P. robustus'' females were more likely to leave their place of birth (
patrilocal In social anthropology, patrilocal residence or patrilocality, also known as virilocal residence or virilocality, are terms referring to the social system in which a married couple resides with or near the husband's parents. The concept of locat ...
). This also discounts the plausibility of a harem society, which would have resulted in a
matrilocal In social anthropology, matrilocal residence or matrilocality (also uxorilocal residence or uxorilocality) is the societal system in which a married couple resides with or near the wife's parents. Thus, the female offspring of a mother remain l ...
society due to heightened male–male competition. Males did not seem to have ventured very far from the valley, which could either indicate small home ranges, or that they preferred dolomitic landscapes due to perhaps cave abundance or factors related to vegetation growth.


Life history

Dental development seems to have followed about the same timeframe as it does in modern humans and most other hominins, but, since ''Paranthropus'' molars are markedly larger, rate of
tooth eruption Tooth eruption is a process in tooth development in which the teeth enter the mouth and become visible. It is currently believed that the periodontal ligament plays an important role in tooth eruption. The first human teeth to appear, the decidu ...
would have been accelerated. Their life history may have mirrored that of gorillas as they have the same brain volume, which (depending on the subspecies) reach physical maturity from 12–18 years and have birthing intervals of 40–70 months.


Palaeoecology


Habitat

It is generally thought that ''Paranthropus'' preferred to inhabit wooded, riverine landscapes. The teeth of ''Paranthropus'', ''H. habilis'' and ''
H. erectus ''Homo erectus'' (; meaning "upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as '' H. heidelbergensis'' and '' H. antecessor' ...
'' are all known from various overlapping beds in East Africa, such as at Olduvai Gorge and the
Turkana Basin An '' Acacia'' tree in the Kokiselei river, northern Kenya The greater Turkana Basin in East Africa (mainly northwestern Kenya and southern Ethiopia, smaller parts of eastern Uganda and southeastern South Sudan) determines a large endorheic ba ...
. ''P. robustus'' and ''H. erectus'' also appear to have coexisted. ''P. boisei'', known from the Great Rift Valley, may have typically inhabited wetlands along lakes and rivers, wooded or arid
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It m ...
s, and semiarid woodlands, though their presence in the savanna-dominated Malawian Chiwondo Beds implies they could tolerate a range of habitats. During the Pleistocene, there seems to have been coastal and montane forests in Eastern Africa. More expansive river valleys—namely the
Omo River The Omo River (also called Omo-Bottego) in southern Ethiopia is the largest Ethiopian river outside the Nile, Nile Basin. Its course is entirely contained within the boundaries of Ethiopia, and it empties into Lake Turkana on the border with Keny ...
Valley—may have served as important refuges for forest-dwelling creatures. Being cut off from the forests of Central Africa by a savanna corridor, these East African forests would have promoted high rates of
endemism Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
, especially during times of climatic volatility. The Cradle of Humankind, the only area ''P. robustus'' is known from, was mainly dominated by the
springbok The springbok (''Antidorcas marsupialis'') is a medium-sized antelope found mainly in south and southwest Africa. The sole member of the genus ''Antidorcas'', this bovid was first described by the German zoologist Eberhard August Wilhelm v ...
''Antidorcas recki'', but other antelope,
giraffes The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, ''Giraffa camelopardalis ...
and
elephants Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae and ...
were also seemingly abundant megafauna. Other known primates are early ''Homo'', the
hamadryas baboon The hamadryas baboon (''Papio hamadryas'' ) is a species of baboon within the Old World monkey family. It is the northernmost of all the baboons, being native to the Horn of Africa and the southwestern region of the Arabian Peninsula. These re ...
, and the extinct
colobine The Colobinae or leaf-eating monkeys are a subfamily of the Old World monkey family that includes 61 species in 11 genera, including the black-and-white colobus, the large-nosed proboscis monkey, and the gray langurs. Some classifications spli ...
monkey '' Cercopithecoides williamsi''.


Predators

The left foot of a ''P. boisei'' specimen (though perhaps actually belonging to ''H. habilis'') from Olduvai Gorge seems to have been bitten off by a crocodile, possibly ''
Crocodylus anthropophagus ''Crocodylus anthropophagus'' is an extinct species of crocodile from the Pleistocene of Tanzania. It lived 1.84 million years ago. It was a large-sized predator reaching a length of . Etymology ''Crocodylus anthropophagus'' was first named by ...
'', and another's leg shows evidence of leopard predation. Other likely Olduvan predators of great apes include the
hunting hyena ''Chasmaporthetes'', also known as hunting or running hyena, is an extinct genus of hyenas distributed in Eurasia, North America, and Africa during the Pliocene-Pleistocene epochs, living from 4.9 million to 780,000 years ago, existing for about ...
''Chasmaporthetes nitidula'', and the
sabertoothed cat Machairodontinae is an extinct subfamily of carnivoran mammals of the family Felidae (true cats). They were found in Asia, Africa, North America, South America, and Europe from the Miocene to the Pleistocene, living from about 16 million until ...
s ''
Dinofelis ''Dinofelis'' is a genus of extinct sabre-toothed cats belonging to the tribe Metailurini or possibly Smilodontini. They were widespread in Europe, Asia, Africa and North America at least 5 million to about 1.2 million years ago (Early Pliocene t ...
'' and ''
Megantereon ''Megantereon'' was a genus of prehistoric machairodontine saber-toothed cat that lived in North America, Eurasia, and Africa. It may have been the ancestor of ''Smilodon''. Taxonomy Fossil fragments have been found in Africa, Eurasia, and No ...
''. The carnivore assemblage at the Cradle of Humankind comprises the two sabertooths, and the hyena '' Lycyaenops silberbergi''. Male ''P. robustus'' appear to have had a higher mortality rate than females. It is possible that males were more likely to be kicked out of a group, and these lone males had a higher risk of predation.


Extinction

It was once thought that ''Paranthropus'' had become a specialist feeder, and were inferior to the more adaptable tool-producing ''Homo'', leading to their extinction, but this has been called into question. However, smaller brain size may have been a factor in their extinction along with gracile australopithecines. ''P. boisei'' may have died out due to an arid trend starting 1.45 mya, causing the retreat of woodlands, and more competition with savanna baboons and ''Homo'' for alternative food resources. South African ''Paranthropus'' appear to have outlasted their East African counterparts. The youngest record of ''P. boisei'' comes from Konso, Ethiopia about 1.4 mya; however, there are no East African sites dated between 1.4 and 1 mya, so it may have persisted until 1 mya. ''P. robustus'', on the other hand, was recorded in
Swartkrans Swartkrans is a fossil-bearing cave designated as a South African National Heritage Site, located about from Johannesburg. It is located in the Cradle of Humankind World Heritage Site and is notable for being extremely rich in archaeological m ...
until Member 3 dated to 1–0.6 mya (the
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...
), though more likely the younger side of the estimate.


See also

* ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Australopi ...
'' * ''
Ardipithecus ''Ardipithecus'' is a genus of an extinct hominine that lived during the Late Miocene and Early Pliocene epochs in the Afar Depression, Ethiopia. Originally described as one of the earliest ancestors of humans after they diverged from the chimp ...
'' * ''
Graecopithecus ''Graecopithecus'' is an extinct species of hominid that lived in southeast Europe during the late Miocene around 7.2 million years ago. Originally identified by a single lower jaw bone bearing a molar tooth found in Pyrgos Vasilissis, Athens, ...
'' * ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
'' * ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial cranium, nicknamed ''T ...
''


References


Further reading

* *


External links


Reconstructions of ''P. boisei''
by
John Gurche John Gurche is an American artist known for his paintings, sculptures, and sketches of prehistoric life, especially dinosaurs and early humans. Gurche is currently an Artist in Residence at the Museum of the Earth in Ithaca, New York. Gurche st ...
*
Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. In 2021, with 7 ...
(August 2016). {{Taxonbar, from=Q111463 Prehistoric primate genera Pliocene primates Pleistocene primates Pleistocene extinctions Cenozoic mammals of Africa Pleistocene genus extinctions Fossil taxa described in 1938