Evolution of Primates

Homo habilis


Vocabulary for Lithic Industries

Conchoidal Fracture

Flake Terminology

CoreTerminology

Prepared Core Techniques

 

NONHUMAN PRIMATES

The history of the order Primates is documented by a rich fossil record, and our models for interpreting this record continue to improve as new information and new analytical techniques become available. This record can be summarized as a series of adaptive radiations, the first of which occurred in the Mesozoic. Primates were one of the early branches of the mammalian adaptive radiation that developed after the Jurassic. This first primate adaptive radiation produced a diverse array of 'archaic primates' that date from late Cretaceous through Eocene

Archaic primates are a disparate group, usually loosely classified into the Suborder Plesiadapiformes. The earliest purported primate is Purgatorius, retrieved from Cretaceous deposits of Purgatory Hill in Montana and from later Paleocene sites. Known only from teeth, it had a dental formula of:

3 1 4 3
________

3 1 4 3

It is classified as a primate because of the blunt cusps on its teeth, unlike the sharp cusps of Insectivores. Earliest primates of the Paleocene appear to occupy the same insectivore niche as did the Mesozoic mammals. Their fingers terminate in claw-like nails and they appear to have climbed with assistance of claws (as do modern squirrels) rather than by grasping with fingers. Large infraorbital fossae suggest that the face bore large sensory vibrissae. Eyes are relatively small relative to the size of skull, fields of vision are generally directed laterally as well as forward, and the orbit lacks a postorbital bar. Later Paleocene primates have a complex dentition, with mesial incisors often markedly procumbent like those of the modern aye-aye and many extant rodents. Premolar and molar anatomy are variable, suggesting diversification into a variety of dietary niches, particularly an increased component of plant materials. The general design of molars resembles that of more modern primates.


The Archaic Primates



Suborder Plesiadapiformes (Simons and Tattersall, 1972)
Superfamily Paromomyoidea (Simpson, 1940)

Family incertae sedis
Genus Purgatorius (Van Valen and Sloan, 1965) Cretaceous to Paleocene, N. America

Family Microsyopidae (Szalay, 1969a)
Genus Berruvius (Russel, 1964) Paleocene to Eocene, Europe
Genus Micromomys (Szalay, 1973) Paleocene to Eocene, N. America
Genus Navajovius (Matthew and Granger, 1921) Paleocene to Eocene, N. America
Genus Palaechthon (Gidley, 1923) Paleocene, N. America
Genus Palenochtha (Simpson, 1937) Paleocene, N. America
Genus Plesiolestes (Jepsen, 1930b) Paleocene, N. America
Genus Talpohenach (Kay and Cartmill, 1977) Paleocene, N. America
Genus Torrejonia (Gazin, 1968) Paleocene, N. America

Genus Alveojunctus ( ) Eocene, N. America
Genus Arctodontomys ( ) Eocene, N. America
Genus Craseops ( ) Eocene, North America
Genus Microsyops (Leidy, 1972) Eocene, N. America
Genus Niptomomys ( ) Eocene, N. America
Genus Tinimomys (Szalay, 1974b) Eocene, North America
Genus Uintasorex (Brown and Rose, 1976) Eocene, N. America

Family Paromoyidae (Simpson, 1940)
Genus Paromomys (Gidley, 1923) Paleocene, N. America
Genus Ignacius (Matthew and Granger, 1921) Paleocene to Eocene, N. America
Genus Phenacolemur (Matthew, 1915) Paleocene to Eocene, N. America, Europe

Genus Arcius () Eocene, Europe
Genus Elwynella () Eocene, N. America

Family Picrodontidae Simpson, 1937)
Genus Draconodus (Tjomida, 1982) Paleocene, N. America
Genus Picrodus (Douglass, 1908) Paleocene, . America
Genus Zanycteris (Matthew, 1917a) Paleocene, N. America

Superfamily Plesiadapoidea (Trouessart, 1897)
Family Plesiadapidae (Trouessart, 1897)
Genus Chiromyoides (Stehlin, 1916) Paleocene to Eocene, N. America, Europe
Genus Nannodectes Gingerich, 1974) Paleocene, N. America
Genus Platychoerops (Charlesworth, 1854) Eocene, Europe
Genus Plesiadapis (Gervais, 1877) Paleocene to Eocene, N. America, Europe
Genus Pronothodectes (Gidley, 1923) Paleocene, N. America

Family Saxonellidae (Russel, 1964)
Genus Saxonella (Russell, 1964) Paleocene, N. America, Europe

Family Carpolestidae (Simpson, 1935b )
Genus Elphidotarsius (Gidley, 1923) Paleocene, N. America
Genus Carpodaptes (Matthew and Granger, 1921) Paleocene, N. America
Genus Carpolestes (Simpson, 1928) Paleocene to Eocene, N. America

Angiosperms (flowering plants) appear during the Early Cretaceous. Seeds of these earliest flowering plants were small and primarily are dispersed by wind or water. Most Early Cretaceous angiosperms were shrubs and weeds that flourished in unstable environments. Insect pollinators and flowering plants coevolved during the Cretaceous. In the late Cretaceous and Paleocene, small arboreal mammals began to exploit the angiosperms, and a proliferation of flowering plant species appear bringing with them a great variety of resources (flowers, fruits, buds, gums, nectars, leaves, bark, and pollen. These resources, and the insects that utilized them, became the foraging grounds of birds and small arboreal omnivorous mammals. By the late Paleocene, large fruits with stored nutrients signal an adaptation of some angiosperm species to animal dispersed propagation. A mammal or bird is enticed by reward or attractant to transport relatively large seeds with substantial stored nutrients away from the parent plant. By the Eocene, a major shift in angiosperm evolution has occurred and modern looking tropical forests, contain a great variety of animal dispersed fruits (Sussman, 1991). Birds, bats, rodents, and primates become important arboreal seed dispersal agents.

Late Paleocene and the following Eocene saw a second and third primate adaptive radiation. The second adaptive radiation was dispersal of prosimian-like primates, represented by two families, Adapidae and Omomyidae. The third radiation is represented by the earliest anthropoid grade of primate evolution, the Oligopithecinae to which Catopithecus and Proteopithecus belong.

Since some of these Eocene primates achieved a prosimian grade of development, this second radiation is described as the beginnings of the euprimates, the "primates of modern aspect." These early euprimates have nails on their digits rather than claws, and an opposable hallux (first toe) suggests the basic pattern of climbing by grasping. A postorbital bar shields the back of the eye of many species, and fields of vision are oriented more forward. These early euprimates are small omnivores that eat gum and search for nectar, small fruits and insects in the terminal branches of trees. They find food visually (as well as by sound and smell) and manipulate it with their hands. They are an important animal component of the Eocene angiosperm forest.


Eocene Primates



Order Primates (Linnaeus, 1758)
Suborder Prosimii (Illiger, 1811)

Infraorder Tarsiiformes (Gregory, 1915b)

Family Omomyidae (Trouessart, 1879)
Subfamily Anaptomorphinae (Cope, 1883)
Genus Absarokius (Matthew, 1915) Eocene, N. America
Genus Aycrossia () Eocene, N. America
Genus Altanius (Dashzeveg and McKenna, 1977) Eocene, N. America
Genus Anaptomorphus (Cope, 1872) Eocene, N. America
Genus Anemorhysis (Gazin, 1958) Eocene, N. America
Genus Chlororhysis (Gazin, 1958) Eocene, N. America
Genus Gazinius () Eocene, N. America
Genus Lovenia (Simpson, 1940) Eocene, N. America
Genus Mckennamorphus (Szalay, 1976) Eocene, N. America
Genus Steinius () Eocene, N. America
Genus Strigorhysis () Eocene, N. America
Genus Teilhardina (Simpson, 1940) Eocene, France
Genus Tetonius (Matthew, 1915) Eocene, N. America
Genus Trogolemus (Matthew, 1909) Eocene, N. America
Subfamily Omomyinae (Trouessart, 1879)
Genus Arapahovius () Eocene, N. America
Genus Chumashius (Stock, 1933) Eocene, N. America
Genus Dyseolemur (Stock, 1934) Eocene, N. America
Genus Hemiacodon (Marsh, 1872a) Eocene, N. America
Genus Jemezius () Eocene, N. America
Genus Macrotarsius (Clark, 1941) Eocene to Oligocene, N. America
Genus Omomys (Leidy, 1869) Eocene, N. America
Genus Ourayia (Gazin, 1958) Eocene, N. America
Genus Shoshonius (Granger, 1910) Eocene, N. America
Genus Stockia (Gazin, 1958) Eocene, N. America
Genus Uintanius (Matthew, 1915) Eocene, N. America
Genus Utahia (Gazin, 1958) Eocene, N. America
Genus Washakius (Leidy, 1873) Eocene, N. America

Subfamily Microchoerinae (Lydekker, 1887)
Genus Microchoerus (Wood, 1846) Eocene to Oligocene, England, France, Germany
Genus Nannopithex (Stehlin, 1916) Eocene, France, Germany
Genus Necrolemur (Fihol, 1873) Eocene, France
Genus Psleudoloris (Stehlin, 1916) Eocene, France, Spain

Family Omomyidae, incertae sedis
Genus Hoanghonius (Zdansky, 1930) Eocene, China

Infraorder Adapiformes (Szalay and Delson, 1979)
Family Adapidae (Trouessart, 1879)
Subfamily Notharctinae (Trouessart, 1879)
Genus Cantius (Simons, 1962b) Eocene, N. America
Genus Copelemur (Gingerich and Simons, 1977) Eocene, Rocky Mt.
Genus Notharctus (Leidy, 1870) Eocene, Rocky Mt.
Genus Pelycodus Cope, 1875) Eocene, N. Americas and Europe
Genus Smilodectes (Wortman, 1903) Eocene, Rocky Mt.

Subfamily Adapinae (Trouessart, 1879)
Genus Adapis (Cuvier, 1821) Eocene to Oligocene, Europe
Genus Agerinaia (Crusafont-Pairo and Golpe-Posse, 1973) Eocene, Spain
Genus Anchomomys (Stehlin, 1916) Eocene, Europe
Genus Caenopithecus (Rütimeyer, 1862) Eocene, Europe
Genus Cercamonius (Gingerich, 1975c) Eocene, Europe
Genus Cryptadapis () Eocene, Europe
Genus Donrussellia (Szalay, 1976) Eocene, Europe
Genus Europolemur (Weigelt, 1933) Eocene, German Democratic Republic
Genus Huerzeleris (Szalay, 1974a) Eocene, Europe
Genus Indraloris (Lewis, 1933) Miocene, Northern India
Genus Leptadapis (Gervais, 1876) Eocene, Europe
Genus Mahgarita (Wilson and Szalay, 1976) Eocene, Texas
Genus Microadapis (Szalay, 1974a) Eocene, Switzerland
Genus Periconodon (Stehlin, 1916) Eocene, Europe
Genus Pronycticebus (Grandidier, 1904) Eocene, France
Genus Protoadapis (Lemoine, 1878) Eocene, Western Europe

Family Adapidae, incertae sedis

Genus Lushius (Chow, 1961) Eocene, China

Suborder incertae sedis
Genus Amphipithecus (Colbert, 1937) Eocene, Burma

Eocene euprimates in the families Adapidae and Omomyidae achieved a prosimian grade of development, and although their anatomy still contains archaic features, they are classified as prosimians. For example, brain sizes are relatively small compared to those of modern prosimians. This small brain combined with robust facial architecture gives them a primitive appearance. Some fossil Adapis and Smilodectes species appear to have brain sizes about half that of living mammals of comparable body size. Adapid locomotor adaptations are quite variable, suggesting a variety of niches at least as diverse as those of modern lemurs. Omomyids exhibit many characters that seem more anthropoid-like:

1. The brain is larger, similar in size to that of the modern tarsier.

2. The infraorbital foramen is small, suggesting that vibrissae are not present.

3. The orbit has the beginnings of postorbital closure

4. The face is relatively short.

In spite of locomotor and niche diversity among the Eocene euprimates, similarity in their anatomy implies that they are monophyletic, that is descended from a common ancestral stock.

The Eocene Oligopithecinae suggest the earliest anthropoids, the third primate adaptive radiation, bore a mosaic of anthropoid and prosimian characters(Rasmussen and Simons (1992). For example a nearly complete cranium of Catopithecus has anthropoid characters such as complete postorbital closure, metopic fusion, and an intraorbital lacrimal foramen. Its prosimain-like characters include small body size, relatively small brains, emphasis on olfaction (evidenced by relatively broad intraorbital region), and prosimian-like shearing crests of molar teeth. Large brains and bunodont molars were not present in early anthropoids. Other anthropoid traits, such as a fusion of the mandibular symphysis, are present in some Eocene prosimians.

The Paleocene and Eocene also saw development and early adaptive radiation of most modern mammalian groups. There are few large mammals; the predominant group seems to be rodents. Euprimate faunas of major continental regions diverge with the opening of the Atlantic barrier during the Eocene which isolates the Old World of Europe, Asia and Africa from the Americas. This trend continues with further isolation of continents in the southern hemisphere. Africa and Europe continue to be separated by the Tethys Sea. The Eocene ends with a great discontinuity -the majority of the Eocene mammalian genera abruptly become extinct, and with them the last of the archaic primates that had survived into the Eocene. Fluctuations in sea level at the Eocene/Oligocene boundary and later during the Oligocene may have produced temporary land bridges from Northeast Africa across the Arabian peninsula into Asia.

The Oligocene is a period of great interest to those who reconstruct the evolutionary history of anthropoids, but it is poorly documented. Whereas Eocene deposits are relatively accessible, and Eocene primates are plentiful, surface Oligocene deposits are rare. Consequently, our knowledge of Oligocene events is biased by the few available localities for study. The most important Oligocene site for fossil primates is the Fayum Depression of Egypt, in the desert southwest of Cairo. Eroding sandstones of this area provide most of our knowledge of Oligocene primates.

The anthropoids, the third major primate adaptive radiation, of primate evolution, is more numerous in Oligocene Primates. The skull assumes its modern configuration with postorbital closure, a fused mandibular symphysis, and a fused metopic suture. Grasping hands of anthropoids do not bear a grooming claw. Eye-hand coordination seems more important as the hand replaces the face as an organ for tactile exploration. Fingers are elongated. Orbits face forward, with overlapping fields that could form the basis for anthropoid stereoscopic vision.


Oligocene Primates



Order Primates (Linnaeus, 1758)
Suborder Prosimii (Illiger, 1811)

Infraorder Tarsiiformes (Gregory, 1915b)

Family Omomyidae (Trouessart, 1879)
Subfamily Omomyinae (Trouessart, 1879)
Genus Rooneyia (Wilson, 1966) Oligocene, N. America
Subfamily Ekgmowechashalinae (Szalay, 1976)
Genus Edgmowechashala (Macdonald, 1963) Oligocene, N. America
Family Tarsiidae (Gray, 1825)
Genus Afrotarsius (Simons and Brown, 1985) Oligocene, Africa

Suborder Anthropoidea (Mivart, 1864)
Infraorder Platyrrhini (E. Geoffroy, 1812)
Superfamily Ceboidea (Simpson, 1931)

Family Cebidae (Bonaparte, 1831)
Subfamily Cebinae (Bonaparte, 1831)
Genus Dolichocebus (Kraglievich, 1951) Oligocene, Argentina
Subfamily Branisellinae (Hershkovitz, 1977)
Genus Branisella (Hoffstetter, 1969) Oligocene, Bolivia

Family Atelidae (Gray, 1825)
Subfamily Pitheciinae (Mivart, 1865)
Genus Tremacebus (Hershkovitz, 1974a) Oligocene, Argentina

Infraorder Catarrhini E. Geoffroy, 1812)
Superfamily Parapithecoidea (Schlosser, 1911)

Family Parapithecidae (Schlosser, 1911)
Genus Parapithecus (Schlosser, 1910) Oligocene, Egypt
Genus Apidium (Osborn, 1908) Oligocene, Egypt
Genus Qatrania (Simons and Kay, 1983) Oligocene, Egypt
Genus Catopithecus (), Oligocene, Egypt

Superfamily Hominoidea (Gray, 1825)

Family Propliopithecidae (Straus, 1961)
Subfamily Propliopithecinae (Delson and Andrews, 1975)
Genus Propliopithecus (Schlosser, 1916) Oligocene, Egypt
Genus Aegyptopithecus (Simons, 1965) Oligocene, Egypt
Subfamily Oligopithecinae
Genus Catopithecus (Simons, 1989) Oligocene, Egypt
Genus Proteopithecus (Simons, 1989) Oligocene, Egypt
Genus Oligopithecus (Simons, 1962a) Oligocene, Egypt

In the Americas, anthropoids (Platyrrhines) may either have evolved from early euprimate stock independently of parallel events in the Old World or share a common ancestry with an Eocene anthropoid such as Catopithecus prior to the Atlantic Barrier. New World anthropoids differ in some anatomical characteristics. Platyrrhines, New World anthropoids, retain a more primitive dental formula:

2 1 3 3
_________

2 1 3 3

Catarrhines, Old World anthropoids, have lost the second premolar, retaining the third and fourth premolars of their early mammalian progenitors:

2 1 2 3
_______

2 1 2 3

As the neocortex expanded with larger brain sizes, the internal ear remains connected to the outer opening, the external auditory meatus, by the formation of a tunnel. In Platyrrhines, this tube is enclosed by a cartilaginous structure that terminates in an ectotympanic ring that anchors the eardrum. In Catarrhines an ossified ectotympanic tube connects the eardrum to the outside . The earliest known Oligocene anthropoid, Catopithecus, lacks a Catarrhine ectotympanic tube but has a Catarrhine dental formula. Catopithecus probably represents a Catarrhine that is close to the Platyrrhine/Catarrhine separation.

The Miocene and Pliocene witnessed a radiation of anthropoids in both the Old and New World continents. The continents were close to their modern configuration and the early Miocene of Europe and Africa is thought to be a period of mild or relatively warm climate that progressively became drier and cooler. The anthropoid adaptive radiation continues with appearance of Hominoidea and Cercopithecoidea. At first, hominoids are more numerous and more diverse than cercopithecoids. The middle Miocene radiation of hominoids is interpreted as a diversification of frugivores in the relatively moist forests of that time period. Hominoids are easily recognized by characteristic molar anatomy, the dryopithecine pattern, named after Dryopithecus, the first fossil ape to be described (in 1856 be E. Lartet from near Paris). The middle and late Miocene of Africa is thought to have had a drier climate, a change in environment that discouraged frugivores, and is marked by presence of hominoids with increased molar enamel thickness and by an increase in the number of Old World monkey genera. The middle Miocene radiation of cercopithecoids is thought to reflect dietary specialization toward omnivory in the drier forests. True folivores have thinner molar enamel (Kay 1981). Thicker molar enamel facilitated the crushing of hard food objects, and increased tooth wear from hard or abrasive materials in the diet. Molar cusps with sharp shearing crests are presumed to be more effective at chopping leaf-like materials into digestible shreds. On the other hand, sharp delicate cusps, characteristic of arboreal folivores, can be damaged during mastication of hard objects. The more terrestrial cercopithecoids exhibit extraordinary rates of dental wear, but their thicker enamel and bilophodont molar shape produce strong selenes of enamel that are both resistant to damage under high molar loads and effective at shearing tough foodstuffs into digestible pieces. The late Miocene marks the beginnings of the radiation of the Hominidae, the humans. Near the end of the Miocene, a dramatic lowering of sea level occurred, leaving the Mediterranean Sea a land-locked basin that dried up, opening dry-land connections between Africa and Europe. At the time the African continental plate impacted the smaller Arabian plate about 16 million years ago, Arabian fossil faunas are similar to those of North East Africa. The Arabian plate continues to move counterclockwise away from Africa to produce the Gulf of Aden rifts.


Miocene Primates



Order Primates (Linnaeus, 1758)
Suborder Prosimii (Illiger, 1811)
Infraorder Adapiformes (Szalay and Delson, 1979)
Family Adapidae (Trousessart, 1879)
Subfamily Sivaladapinae
Genus Indraloris (Lewis, 1933) Miocene, Asia
Genus Sivaladapis () Miocene, Asia
Genus Sinoadapis () Miocene, Asia

Infraorder Lemuriformes (Gregory, 1915b)

Superfamily Lorisoidea (Gray, 1821)

Family Lorisidae (Gray, 1821)
Subfamily Galaginae (Gray, 1825)
Genus Progalago (Macinnes, 1943) Miocene, East Africa
Genus Komba (Simpson, 1967) Miocene, East Africa
Subfamily Lorisinae (Flower and Lydekker, 1891)
Genus Mioeuoticus (Leakey, 1962) Miocene, East Africa
Genus Nycticeboides () Miocene, Asia

Infraorder Tarsiiformes (Gregory, 1915b)
Family Tarsiidae (Gray, 1825)
Genus Tarsius (Storr, 1789) Miocene to Modern, Asia

Family Omomyidae (Trouessart, 1879)
Subfamily Anaptomorphinae (Cope, 1883)
Genus Ekgmowechashala (Macdonald, 1963)
Miocene, N. America

Suborder Anthropoidea (Mivart, 1864)
Infraorder Platyrrhini (E. Geoffroy, 1812)
Superfamily Ceboidea (Simpson, 1931)

Family Cebidae (Bonaparte, 1831)
Subfamily Cebinae (Bonaparte, 1831)
Genus Neosaimiri (Stirton, 1951) Miocene, Colombia
Subfamily Callitrichinae
Genus Micodon () Miocene, Colombia
Subfamily Aotinae ()
Genus Tremacebus (Hershkovitz, 1974a) Miocene, Argentina
Genus Aotus (Illiger, 1811) Miocene, Colombia
Genus Homunculus (Ameghino, 1891) Miocene, Argentina

Family Atelidae (Gray, 1825)
Subfamily Atelinae (Gray, 1825)
Genus Stirtonia (Hershkovitz, 1970a) Miocene, Colombia
Subfamily Pitheciinae (Mivart, 1865) Miocene, South America
Genus Cebupithecia (Stirton and Savage, 1951) Miocene, Colombia
Genus Mohanamico () Miocene, Colombia

Infraorder Catarrhini E. Geoffroy, 1812)

Superfamily Cercopithecoidea (Gray, 1821)

Family Cercopithecidae (Gray, 1821
Subfamily Cercopithecinae (Gray, 1821)
Genus Parapapio (Jones, 1937) Miocene to Pleistocene, Africa
Genus Macaca (Lacépéde, 1799) Miocene to modern, Europe, N. Africa, Asia

Subfamily Colobinae (Blyth, 1875)
Genus Colobus (Illiger, 1811) Miocene to modern. Africa
Genus Libypithecus (Stromer, 1913) Miocene, Egypt
Genus Microcolobus () Miocene, East Africa
Genus Presbytis (Eschscholtz, 1821) Miocene to modern, Asia
Genus Mesopithecus (Wagner, 1839) Miocene to Pliocene, Europe

Family Victoriapithecidae
Genus Prohylobates (Fourtau, 1918) Miocene, Egypt, Libya
Genus Victoriapithecus (von Koenigswald, 1969) Miocene, Kenya, Uganda

Superfamily Hominoidea (Gray, 1825)

Family Oreopithecidae (Schwalbe, 1915)
Genus Nyanzapithecus (Harrison, 1987) Miocene, East Africa
Genus Oreopithecus (Gervais, 1872) Miocene, Italy

Family Pliopithecidae (Zapfe, 1961a)

Genus Pliopithecus (Gervais, 1849) Miocene, Europe
Genus Laccopithecus (Wu and Pan, 1984) Miocene, China
Genus Dendropithecus (Andrews and Simons, 1977) Miocene, Kenya
Genus Crouzelia (Ginsburg, 1975) Miocene, Europe

Family Proconsulidae ()
Genus Proconsul (Hopwood, 1933a) Miocene, East Africa
Genus Limnopithecus (Hopwood, 1933a) Miocene, Africa
Genus Dendropithecus (Andrews, Pilbeam and Simons, 1976) Miocene, Africa
Genus Simiolus (R.E.F. Leakey and Leakey, 1987) Miocene, Africa
Genus Rangwapithecus (Pilbeam et al., 1977) Miocene, Africa
Genus Micropithecus (Fleagle & Simons, 1978) Miocene, Africa
Genus Dionysopithecus () Miocene, Asia
Genus Platydontopithecus () Miocene, Asia

Family Hylobatidae (Gray, 1870)
Genus Hylobates (Illiger, 1811)
Pleistocene to modern, South East Asia, Southern China

Family Pongidae (Elliot, 1913)

Genus Dryopithecus (Lartet, 1856) Miocene, Africa and Europe
Genus Lufengpithecus () Miocene, Asia
Genus Ouranopithecus () Miocene, Europe
Genus Sivapithecus (Pilgrim, 1910) Miocene, Africa, Europe, Asia
Genus Ramapithecus (Lewis, 1934) Miocene, Africa, Europe, Asia
Genus Gigantopithecus (von Koenigswald, 1935) Miocene to Pleistocene, Asia
Genus Graecopithecus (von Koenigswald, 1972) Miocene, Europe

Family incertae sedis
Genus Turkanapithecus (Leakey and Leakey, 1986a) Miocene, East Africa
Genus Afropithecus (Leakey and Leakey, 1986b) Miocene, East Africa Genus Kenyapithecus (L. Leakey, 1962a) Miocene, East Africa


Pliocene Primates



Order Primates (Linnaeus, 1758)
Suborder Prosimii (Illiger, 1811)
Suborder Anthropoidea (Mivart, 1864)
Infraorder Catarrhini E. Geoffroy, 1812)

Superfamily Cercopithecoidea (Gray, 1821)

Family Cercopithecidae (Gray, 1821
Subfamily Cercopithecinae (Gray, 1821)
Genus Cercopithecus (Linnaeus, 1758) Pliocene to modern, Africa
Genus Papio (Müller, 1773) Pliocene to modern, Africa
Genus Cercocebus (E. Geoffroy, 1812) Pliocene to modern, Africa
Genus Dinopithecus (Broom, 1937) Pliocene, Africa
Genus Gorgopithecus (Broom and Robinson, 1949a) Pliocene, South Africa
Genus Procynocephalus (Schlosser, 1924) Pliocene, Central China, N. India
Genus Paradolichopithecus (Necrasov, Samson, and Radules, 1976) Pliocene, Europe
Genus Theropithecus (I. Geoffroy, 1843)Pliocene to modern, Africa

Subfamily Colobinae (Blyth, 1875)

Genus Cercopithecoides (Mollett, 1947) Pliocene to Pleistocene, Africa
Genus Paracolobus (R. Leakey, 1969)Pliocene, East Africa
Genus Rhinocolobus() Pliocene to Pleistocene, Africa
Genus Dolichopithecus (Deperet, 1889) Pliocene to Miocene, Europe

Superfamily Hominoidea (Gray, 1825)

Family Hominidae (Gray, 1825)
Genus Australopithecus (Dart, 1925) Pliocene to Pleistocene, Africa
Genus Homo (Linnaeus, 1758) Pliocene to modern, Worldwide

Note: Species that continue into the Pleistocene are not repeated in the Pleistocene table.


Pleistocene and subfossil Primates



Order Primates (Linnaeus, 1758)
Suborder Prosimii (Illiger, 1811)
Infraorder Lemuriformes (Gregory, 1915b)

Superfamily Lemuroidea (Gill, 1872)

Family Lemuridae (Gray, 1821)
Genus Varecia (Gray, 1863)
Living and subfossil , Madagascar

Family Megaladapidae (Flower and Lydekker, 1891)
Genus Megaladapis (Major, 1894)
Subfossil , Madagascar

Superfamily Indriodea (Burnett, 1828)

Family Indriidae (Burnett, 1828)
Genus Mesopropithecus (Standing, 1905)
Subfossil , Madagascar

Family Daubentoniidae (Gray, 1863)
Genus Daubentonia (E. Geoffroy, 1795)
Modern and subfossil , Madagascar

Family Archaeolemuridae (Major, 1896)
Genus Archaeolemur (Filhol, 1895)
Subfossil , Madagascar
Genus Hadropithecus (Lorenz, 1899)
Subfossil , Madagascar

Family Palaeopropithecidae (Tattersall, 1973)
Genus Palaeopropithecus (G. Grandidler, 1899)
Subfossil , Madagascar
Genus Archaeoindris (Standing, 1908)
Subfossil , Madagascar


Suborder Anthropoidea (Mivart, 1864)
Infraorder Platyrrhini (E. Geoffroy, 1812)
Superfamily Ceboidea (Simpson, 1931)


Family Atelidae (Gray, 1825)

Subfamily Pitheciinae (Mivart, 1865)
Genus Xenothrix (Williams and Koopman, 1952)
Subfossil, Jamaica
`
Infraorder Catarrhini E. Geoffroy, 1812)

Superfamily Cercopithecoidea (Gray, 1821)

Family Cercopithecidae (Gray, 1821
Subfamily Cercopithecinae (Gray, 1821)

Subfamily Colobinae (Blyth, 1875)
Genus Rhinopithecus
Pleistocene to modern, Asia

Superfamily Hominoidea (Gray, 1825)

Family Hylobatidae (Gray, 1870)
Genus Hylobates (Illiger, 1811)
Pleistocene to modern, South East Asia, Southern China

Family Pongidae (Elliot, 1913)
Genus Pongo (Lacepede, 1799)
Pleistocene to modern, South China to Borneo and Sumatra

The Miocene hominoids are particularly interesting to students of human evolution because they reflect adaptive radiations that immediately preceded that of humans. Four families of Miocene apes are generally recognized: Proconsulidae, Oreopithecidae, Pliopithecidae, and Pongidae. Early Miocene hominoids are best known from East African Proconsulidae, whose species vary widely in size from about 3 kg to over 50 kg. Although teeth and fragmentary skeletal fossils are numerous, a remarkably complete skeleton of one species, Proconsul africanus, was found on Rusinga Island, Kenya. This small female hominoid, weighed about 10 kg (the size of a siamang). Its skull resembles that of a small ape with an expanded brain size; lacks a strong supraorbital torus; and gives the visual impression of a small ape of modest musculature. Locomotor anatomy is that of a generalized arboreal quadruped that engages in under-branch suspension, but does not have the extreme anatomy associated with specialized brachiation. It may not have possessed a tail. Another slightly smaller member of the Family Proconsulidae at about 6 kg, Dendropithecus, has long slender limbs, projecting canine teeth, and sectorial lower third premolars. Micropithecus is even smaller. The largest, P. major, is known only from dental remains, and is a gorilla-sized animal.

Oreopithecidae are best known from Oreopithecus bambolii, relatively complete crushed fossils found in coal deposits that formed in wet forests of what is now northern Italy about 14 million years ago. Its dentition is somewhat different from other apes, but its locomotor anatomy appears to be the suspensory configuration widely exhibited by hominoids. Curiously, its pelvis has an expanded iliac blade, a character associated with bipedalism in hominids.

The Pliopithecidae are a group of small gibbon-sized apes from Europe and Asia with primitive looking molar teeth. Their skeleton has the form and proportion similar to modern suspensory primates with almost equally long arms and legs. Pliopithecus is a siamang-sized European quadruped that retains some primitive characteristics, including an incompletely ossified ectotympanic tube.

The Pongidae are a quite variable group of apes who, among other features, have upper canine teeth that exhibit less honing function on P3. Dryopithecus is represented by several species of chimpanzee-sized apes. Dryopithecus and Sivapithecus are known primarily from dental fossils, but one late Miocene Sivapithecus face from the Southern foothills of the Himalayas is unusually complete and appears to have an profile reminiscent of that of orangutans. Another fossil ape genus, Sivapithecus, range in body size from 10 kg to 80 kg. It is a particularly important genus because its wide range extends from Africa across Europe to China. Sivapithecus is distinguished from Ouranopithecus by its more robust mandible, larger mesial teeth relative to cheek teeth, a sectorial P3, and a facial outline that resembles that of an orangutan. Ouranopithecus is a poorly known Chinese ape about the size of a chimpanzee that has many characteristics that are later to be found in early Australopithecines; enlarged molar cusps, thickened enamel, and absence of a sectorial facet on the lower third premolar. Gigantopithecus, judging by tooth and mandible fragments, is the largest of all primates, perhaps exceeding 300 Kg.

Afropithecus, Kenyapithecus, and Turkanapithecus are Miocene apes whose phylogenetic relationships are less certain. Afropithecus is about the size of a female gorilla. The profile of its narrow face contrasts sharply with that of Proconsul. Turkanapithecus is a gibbon-sized ape with a face intermediate in forward projection between Proconsul and Afropithecus. Its maxillary molar and premolar teeth exhibit small accessory cusps that distinguish it from other hominoids. Kenyapithecus, known only from incomplete dental materials, exhibits the thickened molar enamel, robust mandible, and large upper premolars that are characteristic of humans and great apes.

The initial human adaptive radiation, begun in the late Miocene, is well established during the Pliocene. Cercopithecoids diverge into the Cercopithecines (the omnivorous monkeys) and the Colobines (leaf-eating monkeys). As Old World monkeys became more diverse and Miocene apes vanished with dryer late Miocene climates, early humans moved into the savanna habitat.

Thus the human lineage, not especially remarkable among the other primate groups, began its history. But our perception of the record of that history is colored by the ideas and insights of its discoverers. In reality, the record of human history was recovered piecemeal and in random sequence. D ata represented in anatomy and context do not speak for themselves but had to be perceived and interpreted according to the scientific idioms prevalent at the time of their discoveries.

PLIOCENE AND EARLY PLEISTOCENE HUMANS


Ardipithecus ramidus. White, Suwa & Asfaw, 1994

Date: 4.4 MYBP

Distribution: Africa

Sites: Aramis

Synonyms: Australopithecus ramidus

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Most of the Ardipithecus ramidus fossils await description, but the nomen was distinguished from A. afarensis primarily on dental characters. The A. ramidus upper canine is shorter and more incisiform (shaped like an incisor) than that of a pongid, but it is still larger than the canine of later hominids. The mandibular pm3 is not sectorial, so this early hominid lacks the pongid honing mechanism that sharpens the canine against the lingual sectorial cusp of pm3. Tooth enamel is thin like the enamel of chimpanzees, not thick as seen in later hominids. Details of the postcranial skeleton are as yet unpublised, but preliminary hints from a partial humerus suggest human-like arms, ruling out quadrupedalism as the habitual mode of locomotion.
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Australopithecus (Dart, 1925)

The early hominid genus, Australopithecus, is characterized by a mosaic of pongid-like and human-like traits. They have the brain size and brain morphology of pongids and the bipedality of humans. Teeth are characterized by thick enamel and large cusps. The cheek teeth (molars and premolars) are notably large. Australopithecus represents an early radiation of bipedal apes, a niche that later was to become a human life-way. The four generally accepted Australopithecine species groups exhibit varying degrees of increasing megadontia (McHenry 1984).
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Australopithecus anamensis

Date: 3.9 to 4.2 MYBP

Distribution: Africa (Figure 13-5 map)

Sites: Kanapoi

Synonyms:

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Scientists working in the Lake Turkana area of Kenya after 1992 were also finding hominid fossils that are close to 4 million years old. Mave Leakey led a team to Kanapoi, where, in deposits dating between 3.9 and 4.2 MYBP, they recovered fragments of a bipedal hominid, Australopithecus anamensis .
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Australopithecus afarensis (Johanson, White, and Coppens, 1978)

Date: 5 to 3 MYBP

Distribution: Africa

Sites: Baringo, Hadar ( AL 162, AL 288, AL 200), Laetoli, Lothagam, Turkana

Synonyms:

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Brain volume in Australopithecus afarensis is relatively small (380-450 cc) and body size ranges from 30 to 60 kg, comparable to pygmy chimps. However, sexual dimorphism is comparable to that of modern gorillas. Dental proportions are somewhat chimp-like with incisors and canines slightly larger than those of modern humans while premolars and molars are slightly larger than those of apes. Maxillary canines are particularly interesting since they are intermediate between the projecting canine of the pongid and the small non-projecting canine seen in modern humans. The form of P3 varies greatly, ranging from a primitive sectorial form (AL 128-23) to the rotated, molarized biscuspid (AL 33w-1) seen in later hominids (Leonard and Hegmon 1987). The small skull exhibits flaring mastoid processes containing large sinuses. Attachments of robust muscles on the skull surface are marked by easily recognized lines and elevated crests. A. afarensis skulls have a powerful musculature for mastication and shoulder stability. Markings for the temporalis muscle (the temporal line) are distinct and indicate that temporal muscles covered most of the parietal surfaces. If the right and left temporal muscles were large enough to meet along the sagittal suture, the suture would be hidden by a bony sagittal crest. The relative strength of the shoulder and neck musculature is reflected in the robusticity of the mastoid process and nuchal crests.

A. afarensis postcranial anatomy, like the skull, is a mosaic of pongid and human traits. The pelvis and lower limbs are more human than ape, and the shoulder and upper limbs are more ape than human. The scapula suggests a shoulder orientation more like that of a climbing ape than a human. The relative length of the thumb is comparable to a modern human hand. Pongid-like characters include distal phalanges in the hand that terminate in relatively slender apical heads, strongly developed flexor sheath ridges on the proximal and middle phalanges, and curved proximal phalanges. Joint surface characters of the hand as well as the foot, are also a mixture of similarities to modern humans and modern pongids (Aiello and Dean, 1990). The form and orientation of the ankle, the emphasis on dorsiflexion, incipient development of a longitudinal arch, and reduction in opposability of the first toe are characteristics of a habitual biped (Latimer et al., 1987; Latimer and Lovejoy, 1990). The pelvis is clearly that of a biped, and the leg and foot, although not completely modern, differ from apes in the direction of bipedal adaptations. Pongid characters of the feet include relatively long and curved phalanges on digits two through five (Aiello and Dean, 1990).

Independent evidence of the presence of a modern striding human-type gait was found in a volcanic ash fall at Laetoli. Some of these remarkable footprints are indisputably human, and record that human bipedality, in A. afarensis or some other hominid, was present in East Africa 3.6 million years ago. MD Leakey (1981, p. 99) proposed that at site G, "All three individuals were walking in step and most likely holding one another since any deviation in course, no matter how slight, is closely followed by all three and the stride length is virtually the same, in spite of differences in the size of the prints." These three hominids, two larger (adults?) and a smaller one (child?) were crossing a flat featureless surface of volcanic ash. Its texture was like fine sand and their feet sank deeply into the soft surface (White and Suwa 1987).

Although incompletely represented, the thorax of A. afarensis is reconstructed to resemble Homo. The first rib, for example has a single articulation (with the first thoracic vertebra), unlike that of extant pongids, whose first rib articulates with both C7 and T1. The clavicle has a human configuration. The position of the shoulder joint and the cross-section shape of the chest appear human rather than pongid (Ohman 1990).
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Australopithecus africanus (Dart, 1925)

Date: 3 to 2 MYBP

Distribution: Africa

Sites: Makapansgat (MLD 1, MLD 37/38), Olduvai Bed I, Sterkfontein (Sts 5), Taung, Turkana (Omo)

Synonyms: Plesianthropus transvaalensis (Broom 1938); Australopithecus prometheus (Dart, 1949)

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Australopithecus africanus has a slightly larger brain volume (430-520 cc). Temporal muscles are similar in size to that of A. afarensis, but a larger parietal surface requires no posterior or nuchal crests to accommodate them. Maxillary premolars are enlarged and molariform. In general the molar occlusal surface is greatly enlarged and premolars are expanded to function as additional molars. Canine and incisors are relatively small, and P3 is not sectorial. The mandible is larger and more robust than in A. afarensis. The palate is less prognathic and the position of the cheek teeth in relation to the zygomatics and muscles of mastication are shifted to increase mechanical effectiveness. Lateral borders of the nasal aperture are reinforced to form anterior pillars of bone that sustain additional forces during mastication.(return to outline)

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Australopithecus robustus (Broom, 1938)

Date: 2 to 1.5 MYBP

Distribution: Africa

Sites: Kromdraai, Swartkrans (SK 46, SK 48,)

Synonyms: Paranthropus robustus (Broom, 1938); Paranthropus crassidens (Broom, 1949)

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Australopithecus robustus continues a specialization toward enlarging the molar surfaces by enhancing the cheek teeth. The mandible is more robust and there is an extensive addition of bone mass in the body of the mandible. The anterior portion of the temporal muscle is emphasized and enlarged, with its added fibers meeting in the sagittal plane where a crest forms that does not continue to the occipital area. Other muscles of mastication are enhanced and the zygomatic processes are massive and flaring. The face is shortened by movement of the palate and cheek teeth even farther behind the zygomatics. This produces maxillae with zygomatic processes positioned forward in a "dished" face, that is one with a concave profile. Brain size is similar to A. africanus.
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Australopithecus boisei (L. Leakey, Tobias, and Napier, 1964)

Date: 2.8 to 1.4 MYBP

Distribution: Africa

Sites: Olduvai Gorge (OH 5), Peninj, Lake Turkana (KNM-ER 406, KNM-ER 732, KNM-WT 17,000)

Synonyms: Paraustralopithecus aethiopicus (Arambourg and Coppens), 1968; Homo aethiopicus (Olson, 1985)

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Australopithecus boisei represents the extreme of massive molarization of cheek teeth, specialized for some type of mastication that utilized large, flat, crushing surfaces. The massive mandible has even greater bone mass in the mandibular body than does that of A. robustus. The large anterior portion of the temporal muscle produces a large sagittal crest which, although it does not extend to the occipital region, is located near the back of the sagittal region, resulting in long anterior temporal muscle fibers. If strength and action of a muscle are governed by the length and cross-section size of the muscle mass, this appears to be a maximization of anterior temporal muscle strength. A remarkable specimen from West Turkana in Kenya identified as specimen KNM-WT 17,000, and nicknamed the "Black Skull", is intermediate in character with robusticity and crests similar to those of A. boisei, a long face like A. africanus, and a cranial base like A. afarensis. Some anatomists have proposed that KNM-WT 17000 merits a separate species name as representative of a lineage intermediate in anatomy between A. africanus and A. boisei. Cranial volumes remain small (410 to 530 cm3).

Later Australopithecus species (A. africanus, A. robustus and A. boisei) represent lineages that specialized in a dietary niche that changed the masticatory apparatus, emphasizing crushing molars, extended the molar anatomy field mesially to incorporate premolars, and reconfigured the face and skull to maximize molar crushing strength. Whatever this niche might have been, it does not appear to be the niche that gave rise to the lineage antecedent to modern humans. A. afarensis is primitive enough to be near the stem of the radiation of bipedal hominids, and perhaps close to the beginnings of our modern human lineage. If so, A. afarensis may have given rise to at least two daughter hominid lineages (Australopithecus and Homo) who, for a time, shared the African landscape. Whatever else, A. afarensis is the only known representative of the initial radiation of bipedal hominoids.
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Homo (Linnaeus, 1758)

The genus Homo, humanity, originating during this Pliocene radiation of bipedal apes, does not exhibit the expansion of cheek teeth and facial specializations for powerful mastication seen in later and more specialized Australopithecines. Instead, Homo, retaining a more primitive chimpanzee-like face and dentition, exhibits reduced projection of canine, non-sectorial P3, and expanded brain size. Changes in these two anatomical complexes, canine teeth and brain volume, transform cranial and facial anatomy from pongid to human. Since early humans do not share the expanded cheek teeth, they represent a different niche, and probably a different lineage from Australopithecus. The lineage Homo is a continuum of forms that grade into one another and disperse widely over the world. Consequently delineations between chronospecies and geographic races are problematic. Even though the human fossil record is remarkably complete, enthusiastic scholars find the fuel for intense debates in the gaps of hominid history, and our curiosity demands more complete knowledge of the events and processes that shaped human biology.
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Homo habilis (L. Leakey, Tobias, and Napier, 1964)

Date: 2.4 to 1.5 MYBP

Distribution: Africa (

Sites: Laetolii, Olduvai (OH 13, OH 24, OH62); Omo; Sterkfontein Extension (Stw 53); Turkana (KNM-ER 1470, KNM-ER 1590, KNM-ER 1813)

Synonyms: Telanthropus capensis (Robinson, 1953)

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Homo habilis is a small hominid characterized by comparatively small and narrow cheek teeth, a narrow mandible, a larger cranial capacity (600 to 800 cm3), and a bipedal locomotor anatomy. Hands are still robust with relatively long arms, reminiscent of their suspensory ancestry. Legs are short and feet are surprisingly modern.

H. habilis overlaps in time and location with A. boisei and A. robustus. Sites associated with these hominids contain stone tools attributed to the Oldowan Industrial Complex.

Oldowan tools are characterized by a lack of standardization, that is, they are fabricated for function with relatively little stereotyping in form. The manual skill required in their manufacture is rudimentary and easily mastered by a modern human with a few minutes practice. Part of this artifact assemblage consists of objects with naturally useful properties; oval stones for hammers or naturally sharp edges for chopping. Such artifacts are recognized as "utilized" artifacts when they exhibit signs of damage or wear through usage. A manuport is an object that appears to have been transported from some distant source but bears no evidence of modification. The term "artifact" also includes objects thought to be formed by human manufacture, either as tools or as by-products. The term "tool" or "implement" is reserved for artifacts whose trimming scars or polish suggests modification for usage. Flakes trimmed from a core to make a tool are "debitage" if they exhibit no scars of utilization. Flakes that bear evidence of use (damage scars, polishing) are classified as "utilized flakes." Common Oldowan tools include burins, choppers, discoids, polyhedrons, scrapers, and spheroids. Utilized artifacts include hammer-stones, pebbles, and flakes. Manuports are common. Flake debitage marks sites of tool manufacture. Several types of Oldowan activity sites are known, three of which are described here.

1. One type is represented by the KBS site at Koobi Fora, briefly occupied, dated to about 1.8 MYBP, and was sealed and protected by an ash fall. In an area about 15 m x 15 m archaeologists found about 500 artifacts, mostly debitage flakes from manufacture of stone tools. Bone fragments of crocodile, gazelle, hippo, pig, and porcupine were present. The site is a single horizon, suggesting a single occupation by hominids who manufactured tools and brought parts of animal carcasses to the site. Perhaps this was a brief home base for a band or foraging party. About 1 km away are the remains of a hippopotamus in an ash-filled stream channel. Mixed with the hippo carcass are about a hundred artifacts, mostly flakes. Although there is no way of knowing whether the hominids killed the hippo, they appear to have partially butchered the carcass.

2. Site DK 1, in the lower levels of Bed I at Olduvai Gorge, is an occupation site with evidence of longer usage. It lies just above the basalt layer that underlies Olduvai Gorge and is overlain by a volcanic ash dated at 1.75 MYBP. The most surprising feature at DK 1 is a loosely-piled circle of basalt stones, mostly from 10 to 25 cm in diameter. They appear to have been carried from an exposure of the underlying basalt layer and piled to form a crude circle 3 to 4 m in diameter and from 15 to 20 cm high at slightly over 0.5 m intervals around the circle. The area immediately outside the circle is clear, but there is a scatter of stones from the piles starting less than meter outside the circle. The apparent living area inside and outside the circle is littered with Oldowan tools and fragments of animal bones, suggesting that the structure might have been a temporary shelter, perhaps skin or grass supported by upright branches anchored in part by piles of stones. The prey species, if the animal bone litter represents midden material, includes animals as large as elephants.

3. Another Olduvai Gorge living floor is known from the Zinjanthropus site, FLK. Here, a large area littered with artifacts and fragmentary bone is surrounded by a relatively barren zone, beyond which artifacts and debris again increases. This pattern is consistent with the hypothesis that the central occupation area was enclosed, at least to the South and East, by a thorn fence. Artifacts and bone fragments were not dropped in the part of the site occupied by the fence or windbreak.

Oldowan artifacts are associated with numerous occupation sites and a surprising number of kill/butchering sites. The largest of the butchered animals is the elephant, Deinotherium. Although one can not exclude the possibility that the butchery sites are opportunistic scavenging of accidentally trapped animals, the repeated discovery of large animals that died and were butchered under identical circumstances lends credence to the idea that Oldowan hominids may have deliberately driven large animals into the mud and killed them. The artifact types of the Oldowan Industry becomes more diverse in with passage of time.
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Vocabulary for Lithic Industries

The Director of the Danish National Museum, Christian Jurgensen Thomsen (1788-1865) organized the antiquities on display in 1819 into three ages: the Age of Stone, the Age of Bronze, and the Age of Iron. Thompsen's Stone Age was divided by sir John Lubbock (1834-1913) into two units, the Paleolithic Age (chipped stones) and the Neolithic Age (polished stones) (Lubbock, 1865; Prehistoric Times). Edward Lartet subdivided the Paleolithic on the basis of fauna into three divisions - Lower Paleolithic (Hippopotamus Age), Middle Paleolithic (Cave Bear and Mammoth Age), and Upper Paleolithic (Reindeer Age). The term Mesolithic was added to denote an intermediate period before the Neolithic. Paleolithic generally refers to lithic industries before the beginning of the Holocene (ca. 10,000 YBP). This vocabulary is Eurocentric; that is, traditional usage of the terms described lithic industries in Europe, and do not apply to other regions as accurately. In Africa, prehistorians adopted the terminology Early Stone Age (ESA), Middle Stone Age (MSA), and Late Stone Age (LSA) to refer to industries generally analogous to divisions of the Paleolithic in Europe. J.G.D. Clark (1967) proposed a vocabulary that is less Eurocentric and more readily adapted to different regions:

TERM DESCRIPTION
Mode I Emphasis is on simple flakes, cores, and choppers (Oldowan, Clactonian). Flake tools are unstandardized in form.

Mode II Direct percussion techniques produce very large flakes and artifacts have standardized forms (Acheulean).

Mode III Emphasis is on flakes produced from prepared cores (Mousterian, Stillbay).

Mode IV Emphasis is on blades and burins.

Mode V` Emphasis is on microliths.

Mode VI Includes ground (polished) stone tools and pottery. A great variety of lithics are present, but emphasis is often on blades. Grinding stones for preparing flour from cereals are often


This vocabulary can be combined with that of European archaeology:


Paleolithic (Old Stone Age) -
Early (Lower) Paleolithic
Mode I and Mode II lithics.
Analogous to Early Stone Age in Africa.

Middle Paleolithic
Mode III lithics.
Generally analogous to Middle Stone Age in Africa. Although Mode IV technology is rare in Africa, it is known from both MSA and LSA African assemblages.

Late (Upper) Paleolithic
Mode IV lithics.
Generally analogous to Late Stone Age in Africa.

Mesolithic
Mode V lithic technology.
Neolithic
Mode VI technology.

This terminology does not work well for all geographic regions. For example, the Paleolithic of Eastern Asia lacks a typical Mode II technology. The Eastern boundary of Mode II lithic distribution across Asia and Europe is called Movius' Line (after Movius).
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Conchoidal Fracture
Very smooth grained (glass-like) stone materials have consistent fracture characteristics and readily produce very sharp edges. If sufficient force is applied near the edge of a stone to fracture it (Figure 38.__ A.1), its components separate in a conchoidal fracture pattern. The parent block is called a core, and the portion removed is a flake. The point at which force is applied, the poi nt of percussion, is usually recognizable by a small scar of damaged crystalline matrix. The fracture surface on the core is the flake release surface and the analogous surface on the flake is the ventral surface. These surfaces have consistent features that allow us to reconstruct each application of force. The flake release surface tends to expand cone-like away from the point of percussion in a plane parallel to the surface of the core. The release surface curves outward (dorsally) away from the point of percussion, thinning the flake distally and producing a noticeable proximal bulge, a bulb of percussion. This bulb, also known as the conchoid, is the area of maximum convexity. The surface of the flake bearing the point of percussion is called the striking platform or talon. The release surface of both core and flake exhibit ripples formed as the fracture forces oscillated in its passage through the stone's crystalline matrix. These ripples are visible in the stone as arcs whose focal point is the point of percussion. Characteristics of a flake are determined by the nature of the fracture force and the shape of the core.
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Flake Terminology
The line of pressure that produced the flake is indicated by the focal point and orientation of the concentric ripples in the release surface. For measurement, The main flake axis is the presumed line of pressure. An end-struck flake (13) is elongated along the main axis (length). A side-struck flake is elongated at right angles to the main axis (14). If the main axis is oblique to the axis of symmetry, the flake is corner-struck (16).

When flaking serves to modify the form (outline) of the implement, it is called trimming. The term retouch is applied with flaking modifies the edge without altering form. The edge may be retouched bifacially (38), unifacially (39), or with alternate retouch in which no two adjacent flakes are from the same direction ( ). Implements with unifacial edges which are prepared from different surfaces are called alternately unifacial or retouche altere (41).
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Core Terminology
The elementary or clacton technique involves the removal of ad hoc flakes by heavy blows. The core usually has an edge angle of less than 90 o and the flakes produced have wide angles. Note that the angle of the core and the angle of the flake, measured on the release surfaces are supplementary. The clacton type core has flakes removed from a number of directions (19) and may be bipolar (20) or uniconical (21) in cross section. If flaking is continued, the core shape approaches a spheroid and is called a polyhedral core. Such polyhedral cores which have their flake scars largely removed by battering are called spheroids. Cores can be almost any shape, but if there are less than 5 primary flake scars, they are considered casual cores.
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Prepared Core Techniques
The phrase prepared core technique refers to a standardization of cores to result in a predictable form of flake and to increase the efficiency of flake manufacture.

The levallois core (or tortoise core) is made from a split pebble (24) whose dorsal surface is prepared by a series of radial flakes (25). A levallois core may have as many 19 to 20 primary dorsal flake scars. The edge of the core is flaked to produce a suitable angle (26) and small flakes may be removed to form a striking platform (27). A powerful blow removes most of the dorsal surface of the core as a single levallois flake (28). The core now has a slightly concave dorsal surface (29) and the dorsal surface of the large levallois flake shows radial flake scars whose origin (point of percussion) has been left on the core (30). The levallois flake often has a platform with minute facets, radial or multidirectional flake scars on the dorsal surface without origins, and a blank flake release surface. The dorsal surface levallois core is returned to its original shape by radial trimming flakes. This procedure removes the depression that marks the flake release scar of the levallois flake and restores the radial symmetry of the core. After a few trimming flakes to prepare the striking platform, another levallois flake is removed. The processes of trimming and flaking continues until the core is too small to produce satisfactory levallois flakes.

The Mousterian core (or disc core) begins with a split pebble whose dorsal surface is circular (33). It is trimmed with bold (short thick) radial flakes to produce a spoke-like pattern on an angular polyhedral core. Mousterian cores typically have 5 to 7 primary dorsal flake scars. The blow to produce flakes is delivered to intersection of flake scars (33a) in order to produce Mousterian flakes (34) that are ribbed and pointed. Note that the dorsal surface of the flakes will bear flake scars that are convergent. The core is rotated and radially flaked in this manner until it is too small to continue. This procedure can produce many side-struck flakes, and over-striking results in a levallois-like flake.

Cores that are intermediate between Levallois and Mousterian are described as Levallois-Mousterian.

The blade core is flaked to produce parallel ridges (35,36). This allows the removal of long flakes with parallel margins (36a). A blade is an elongate flake with parallel margins, a tendency for the primary flake scars to be parallel, and a length that is two times its breadth.
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PLEISTOCENE HUMANS

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Homo erectus (Dubois, 1892)

Date: 1.8 to 0.2 MYBP

Distribution: Africa, Europe, Asia

Sites: Chenchiawo; Hexian; Mauer; Modjokerto; Olduvai [O.H.9; O.H. 12]; Omo; Sangiran [#4; 17]; Salé; Ternifine; Tiblitz, Trinil; Turkana [East: KNM-ER 1808, 3733; 3883; West: KNM-WT 15000]; Zhoukoutian (Lower cave)

Synonyms: Pithecanthropus erectus (Dubois 1893); Hylobates giganteus (Bumuller, 1899); Sinanthropus pekinensis (Black and Zdansky, 1927); Homo modjokertensis (von Koenigswald, 1936); Atlanthropus mauritanicus (Arambourg, 1954); Homo leakeyi (Heberer, 1963); Tchadanthropus uxoris (Coppens, 1965); Homo ergaster (Groves and Mazak, 1975)

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Homo erectus is somewhat larger than H. habilis. For example the slight, over one meter tall OH62 (H. habilis) contrasts markedly with the two meter tall KNM-WT 15000 (H. erectus). The larger bodied H. erectus has a substantially increased cranial capacity. Brain volume ranges from 800 to 1200 cc, with a mean of about 900 cc. The general features of the skull are somewhat more robust than those of H. habilis. Cranial bones are slightly thicker and there is a large supraorbital torus. The thickened roof of the calvaria sometimes produces an elevation (a sagittal keel) down the mid-line of the top of the skull vault. The occipital bone is strongly angled at the nuchal area and has a broad occipital torus. Maximum breadth of the skull vault is near the base, usually between the ear apertures. In spite of a larger and robust face, cheek teeth (molars are premolars) are relatively small, and the mandible has no chin. Alveolar prognathism is pronounced. The relatively small endocranial volume relative to the size of the supraorbital torus and face produces a marked postorbital constriction, and usually a supratorial sulcus. No articular tubercle is associated with the mandibular fossa.

The essentially modern postcranial skeleton is remarkable primarily in its pelvic anatomy. Though the ilium of the os coxa has a buttress of cortical bone that rises vertically from the acetabulum toward the iliac crest, the general anatomy and function of the hip joint are similar to that of contemporary humans. Several early hominid coxae are known -KNM-ER 3228; Olduvai Hominid 28; Arago XLIV (Rose 1984). Femora are relatively straight with the position of minimum circumference relatively low on the shaft when compared to modern femora.

Rightmire (1986) estimates the average body weight of H. erectus to be about 48 kg. Thus, a brain to body ratio suggests a brain volume averaged about 87% of that of a modern human of comparable size (Rightmire, 1990).

One of the most important H. erectus fossils is KNM-WT 15000, a nearly complete skeleton of a 12 year old male from a horizon about 1.6 MYBP at Nariokotome, West of Lake Turkana. Another extraordinary fragmentary postcranial skeleton (KNM-ER 1808) shows extensive inflammation of the periosteum, the fibrous membrane that surrounds bone. Widespread abnormal bone deposition has occurred on the periosteum, a general and non-specific tissue reaction. A popular, but not conclusive, interpretation was that KNM-ER 1808 suffered from hypervitaminosis of vitamin A (Walker, Leakey & Walker, 1985).

H. erectus dispersed and differentiated in geographical races across Africa and Asia. There are numerous archaeological sites that could be H. erectus in Europe, but no unquestionable H. erectus fossils are currently known.

H. erectus is associated with the Acheulean Industrial Complex except in Asia East of India, where the technology is more Oldowan-like. Lithic industries like the Oldowan and Acheulean, which emphasize choppers, simple flake tools, handaxes and/or cleavers, are grouped under the term Early Stone Age (ESA), or the generally equivalent term, Lower Paleolithic. ESA sites comprise the earliest extensive evidences of human activities. Since stone tools readily survive in habitation or activity sites, there is a rich archaeological record during H. erectus time periods. The important H. erectus site at Zhoukoudian in northern China contains evidence of fires. Numerous H. erectus archaeological sites contain animal bones, suggesting habitual hunting. A common animal represented in East African H. erectus sites is an extinct relative of the gelada baboon, but H. erectus butchered game as large as hippo and elephant in numerous localities.

Early Acheulean and later Oldowan Industries were contemporaneous in Africa. Indeed, it is possible that some of the later Oldowan tools were scavenged from discarded early Acheulean implements. The Acheulean Industrial Complex is distinguished from the Oldowan by an ability to strike large flakes, a standardization of artifact form, and by a high level of manual skill. Large flakes struck from large cores are retouched to form effective tools, the final form of which bears little resemblance to the original stone.. A characteristic implement is the pear-shaped, bifacially flaked hand-axe. Another common tool is a similar biface modified by a tranchet flake to form a cleaver.

The skills required to manufacture these implements are comparable to playing a piano in that one can learn how to read music in a brief study session, but many months or years of practice are needed to acquire the manual skills to perform the music. A modern human can learn what to do to replicate Acheulean technology in a few hours, but months or years of practice are required to perfect the skill exhibited in artifact manufacture by H. erectus. Manual skills require a substantial portion of the cerebrum be devoted to hand coordination, specifically the expanded and specialized motor cortex (Figure 13- ).

Much of the motor cortex is allocated to the control of muscles that support uniquely human skills: the toe and ankle for bipedality; large hand area with independent control of digits for manual skills; mouth for speech. Part of the process of becoming human can be understood in terms of cumulative changes in the motor cortex. These changes, at least those related to speech, manual skills, and bipedality, must represent at least part of the expansion in brain volume seen in the genus Homo.

The Acheulean Industrial Complex, remarkably uniform and persistent, was the most sophisticated stone tool industry on earth for more than a million years. Some African Acheulean assemblages are so similar to those in Europe and Asia that scientists have joked that Acheulean tool types were genetically coded into the early humans. One of the later Acheulean innovations in tool manufacture is the levallois technique, an important achievement in lithic technology. In this technique, an elongated core is trimmed radially. Side trimming flakes correct any deviations in the shape of the core, and a base or striking platform is carefully prepared. When the core is properly shaped, a large flake (consisting of most of the dorsal surface of the core) is removed by a powerful blow to the platform, producing a levallois flake , shape and size properties of which that permit its ready use as an implement or its retouch into an artifact with a minimum of effort. This is a "mass production" method of generating useful flakes. The core is then retrimmed to appropriate shape and another large flake is removed. This process is repeated until the core is too small to produce suitable flakes. Levallois flakes are easily recognized by the multidirectional flake release scars on their dorsal surface.
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Homo sapiens (Linnaeus, 1758)

It is normal for scientists to use conjecture to fill gaps in the paleontological data, and to speculate about relationships between fossils. Unfortunately, many of the best dated archaeological sites have produced only fragmentary fossils or worse, only artifacts. However, great interest in a history of our own species presents some special problems in scientific interpretation. Perhaps because more detail is expected than is available in the archaeological record, distinctions between hypothesis and observation are often overlooked. Generally, the nature of these errors are to oversimplify, especially to incorporate inadequate population concepts into models. Morphological differences in the Pleistocene humans appear to be extremely variable within a population, as well as between populations (polytypic). The skeletal nature of fossils encourages scientists to place great importance on morphology and to use morphotypes to define lineage groups but there is no infallible guide to distinguish between interdemic (within population) from intra-deme (between populations) variation. A conservative approach would be to list the full range of anatomies and specify - under the rationale that descendants have ancestors- that archaic H. sapiens was variable and that some, perhaps all of these forms are antecedent to modern humans. The following model arbitrarily divides archaic H. sapiens into three subspecies morphotypes, with the caveat that it is a model of convenience and certainly oversimplifies the history of human populations.
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Homo sapiens soloensis (Oppenoorth, 1932)

Date: ca 400,000 to 100,000 YBP

Distribution: Africa, Asia, Europe

Sites Arago {#21]; Dali; Laetoli [L. H. 18]; Mapa; Ndutu; Ngandong [#7]; Omo [Omo 2]; Steinheim; ?Swanscombe

Synonyms: Homo sapiens steinheimensis (Berckhemer 1936), Homo primigenius asiaticus (Weidenreich, 1932 nec Linnaeus, 1758)

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Early or archaic Homo sapiens differs from H. erectus primarily in its larger cranial volume. Since there are geographic populations of H. erectus whose anatomy already expresses an evolutionary trend toward a large skull size, the boundary between chronospecies (H. sapiens and H. erectus) is confounded by morphological overlap in characters. Generally early H. sapiens samples have larger mean brain volumes than those of H. erectus in the same geographic region. Early H. sapiens lack chins and retain a large face and robust cranium reminiscent of H. erectus. Maximum breadth of the cranial vault is near the cranial base, usually low on the temporal bones. In other features, they exhibit a regional diversity even more varied than H. erectus.

Fossils that represent the transition from H. erectus to H. sapiens are lumped in this discussion into a single archaic subspecies, H. sapiens soloensis. Because its wide distribution is be accompanied by regional variation (surely regional races if not speciation), this classification is one of convenience in the absence of effective methods to reconstruct the population biology of fossil forms.
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Homo sapiens rhodesiensis (Woodward, 1921)

Date: ca 0.4 to 0.1 MYBP

Distribution: Africa, Europe

Sites: Bodo, Elandsfontien; Kabwe; Petralona; ?Singa

Synonyms: Cyphanthropus rhodesiensis (Pycraft, 1928), Homo kanamensis (L. Leakey, 1935), Palaeoanthropus njarensis (Reck and Kohl-Larsen, 1936), Africanthropus njarensis (Weinert, 1938)

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A divergent anatomy, i.e. pneumatised supraorbital torii, pneumatised mastoid processes, and massive faces, suggests a separate archaic human lineage. It is possible that this group does not represent a separate taxon from H. sapiens soloensis, but are merely its extremes of size variation or sexual dimorphism. Lithic associations are tenuous, but are either Acheulean or Middle Stone Age.

The Bodo cranium was reconstructed from about 100 fragments found in a 25 m2 area in a context that suggests Middle Pleistocene age - between 0.5 and 0.2 MYBP (Conroy 1978; Kalb et al., 1980; 1984). The immediate area around the Bodo discovery contains abundant Acheulean tools and numerous hippopotamus remains. Some of the hippo skeletons are associated with stone tools and appear to be butcher sites. Cut marks on the Bodo skull indicate intentional postmortem defleshing (White, 1986), a behavior associated among modern peoples with "trophy" preparation and, more commonly, funerary practices.
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Homo sapiens neanderthalensis (King, 1864)

Date: 135,000 to 29,000 YBP

Distribution: Circum-Mediterranean

Sites: Amud, Combe-Grenal, Devil's Tower, Ehringsdorf, Forbes' Quarry, Jebel Irhoud, Krapina, La Chapelle-aux-Saints, La Ferrassie, Le Moustier, La Quina, Monte Circeo, Neanderthal, Saccopastore, Shanidar, Tabun, Teshik-Tash

Synonyms: Protanthropus atavus (Haeckel, 1895); Homo europaeus primigenius (Wisler, 1898); Paleanthropus krapiniensis (Sergi, 1911); Homo primigenius (Schwalbe, 1903); Homo antiquus (Adloff, 1908);Homo transprimigenius mousteriensis (Forrer, 1908), Homo neanderthalensis (Bonarelli, 1909);Homo priscus (Krause, 1909); Palaeanthropus europaeus (Sergi, 1910); Homo calpicus (Keith, 1911);Archanthropus (Arldt, 1915); Anthropus neanderthalensis (Boyd-Dawkins, 1926);Metanthropus (Sollas, 1933); Pithecanthropus neanderthalensis (Sklerj, 1937)

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The term Neanderthal, H. sapiens neanderthalensis King, 1864, refers to a geographic population of early humans that occupied the Mediterranean area between 130,000 YBP and 35,000 YBP. As a group they are markedly different from most other humans, perhaps enough so that it may be biologically more appropriate to treat them as a separate species. Even though many Neanderthal traits are present in modern humans, the constellation of characters together in each individual sets Neanderthals apart from other humans.

Neanderthal cranial capacity ranges from 1,245 to 1,740 cc with an average near 1,520 cc. The skull has an unusual shape (Figure 13- ). Maximum cranial width is low on the temporal bones, producing an oval but long, low, large skull, the base of which is relatively flat. There is a profoundly robust and protruding occipital bone (a character called an occipital bun). Large frontal sinuses are restricted to the prominent supraorbital torus that forms a double arch above the orbits. Though mastoid processes are small (relative to the robusticity and size of the cranium), a powerful musculature is suggested in this region by mastoid and juxtamastoid crests. The face is prognathous and large. The maxilla is inflated and reinforced mesially to form an unusually large buttress for the mesial teeth. However, Neanderthal dentition is not robust when compared to that of earlier Homo. Molars tend toward fused roots and often have an large pulp cavity (taurodontism), but the most unusual dental character appears to reflect specialized use of the incisors and canines. These mesial teeth are usually much more worn than cheek teeth, perhaps from clamping objects with them, or some other special activity. The entire dentition is shifted slightly forward in the face, leaving a retromolar space on the mandible behind M3. Individuals from one site, Krapina, have artificial grooves along the cemento-enamel junction of the mesial teeth, suggesting the habitual use of toothpicks to probe the interproximal dental spaces (Frayer and Russell 1987). A buttressed maxilla and a robust interorbital anatomy suggest specialized use of mesial teeth unlike other populations of Homo.

The postcranial skeleton indicates a powerful, barrel-chested physique with sturdy shoulders and arms. The shoulder joint had slightly less mobility of dorsoventral arm movements in a horizontal plane (Churchill and Trinkaus 1990) -the type of throwing movements used by modern humans. Shafts of radius and ulna are somewhat bowed, but not greater than those of heavily muscled modern humans. Unusually robust cervical spines and ligament attachment areas to the vertebrae, scapula, and long bones indicate exceptionally powerful shoulders and upper arms. Stout fingers terminate in more robust phalanges than modern hands. Both total thumb length and distal thumb phalanx length are longer than in our thumbs, suggesting a more powerful whole hand grip but a weaker precision grip, i.e., holding an object between thumb and fingers. Generally Neanderthal body build is comparable to that of modern humans from cold climates (Allen's Rule); powerful hands, relatively short limbs and an average male stature of about 166 cm (5' 4").

In contrast, the Neanderthal pelvis differs notably from the average modern pelvis (Rak and Arensburg, 1987). Acetabula are located more posteriorly (toward the sacrum), and coxae are reorganized to support the biomechanics of this rearrangement of the hip joint. The ischium is shorter and the subpubic angle of the male Neanderthal of 110o is more obtuse than the average 60o of a modern human male pelvis. The altered biomechanics of the Neanderthal pelvis includes a distal shift in the internal obturator groove. These changes constrict the dimensions of the pelvic aperture, the vital passageway constricting the birth canal. An adequate opening is maintained by increasing the length of the pubic rami.

The longer pubic ramus was known from fragmentary coxae, and there had been some speculation that the Neanderthal pelvis had a relatively larger pelvic inlet than other modern H. sapiens. The possibility of a larger pelvic inlet stimulated speculation that Neanderthals were larger at birth and perhaps had a longer gestation period than modern humans. However, discovery of an intact male coxae and sacrum with the Kebara 2 skeleton demonstrated that is spite of its altered configuration, the Neanderthal pelvic inlet is similar in size to modern humans (Rak and Arensburg 1987). Differences in pelvic anatomy exhibited by Neanderthals appear to be related to biomechanics of posture, and perhaps to sexual dimorphism.

There has been speculation that Neanderthals exhibited a soft palate and larynx anatomy that differed from modern humans in a manner that precluded or made speech difficult. Proponents of this model expected the Neanderthal hyoid bone to be different from modern anatomy in form and location. Subsequent discovery of a Neanderthal hyoid at Kabara that is identical to modern humans falsified that model and raises the possibility that Neanderthals were anatomically capable of fully modern speech sounds (Arensburg et al., 1990).

Neanderthals are relatively well known, considering their limited temporal and geographical distribution, because they sometimes buried their dead in limestone caves. These burials, protected from scavengers in soil conditions conducive to bone preservation, provide a better sample of Neanderthal anatomy than is available for other early hominids. Neanderthal artifacts are usually considered part of the Mousterian Industrial Complex, a series of artifact assemblages that show variation in geography and time. Similar artifact assemblages in Africa are known to be associated with non-Neanderthals, while some French Neanderthals used Chatelperronian artifacts, a tool assemblage usually associated with more modern looking people.

The Mousterian Industrial Complex employed the Mousterian prepared core technique in which an oval core was carefully trimmed by removing shaping flakes in a radial pattern across its working surface. Next, a series of modest sized flakes was removed in a radial pattern, producing a number of generally pointed flakes with convergent flake release scars on their dorsal surfaces. These "Mousterian" flakes were then either used "as is" or retouched into artifacts. Additional shaping flakes may be removed to recreate the desired core outline, and then another series of flakes are removed in a radial pattern. This process was repeated until the core becomes unsuitable, usually due to its diminishing size. Like the levallois technique, this procedure is a mass produces useful standardized flakes. Mousterian flakes tend to have convergent flake scars on their dorsal surface and do not typically have the multidirectional dorsal flake scars seen on levallois flakes, although a slightly more powerful blow during flake removal will produce a levallois-like flake from a Mousterian core.

Flakes are retouched to make backed blades, burins, denticulates, points, side scrapers, and occasional small handaxes. Though styles and ratios between artifact types vary with geographic region and activity, the Mousterian is generally similar everywhere it is recognized. Although bone and engraved or incised objects are often found in Mousterian sites, bone tools are rare. Lithic industries like the Mousterian, which emphasize small flakes produced with a prepared core technology, are termed Middle Stone Age (MSA) or Middle Paleolithic. A major departure from the ESA is the rarity of large bifacial tools like hand axes or cleavers.
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Homo sapiens sapiens (Linnaeus, 1758)

Date: ca. 100,000 YBP to Holocene

Distribution: Africa, Asia, Europe

Sites: ?Border Cave, Combe-Capelle, Cro-Magnon, Fish Hoek, Grimaldi Jebel Qafzeh [#6], Mugharet es Skhul [Skhul 5], Omo (Omo 1), Predmosti; Zhoukoutian Upper Cave,

Synonyms: Homo sapiens fissilis (Gorjanovic-Kramberger, 1905); Homo grimaldii (Lapouge, 1906); Notanthropus eurafricanus recens (Sergi, 1911); Notanthropus eurafricanus archaius (Sergi, 1911); Homo mediterraneus fossilis (Behm, 1915); Homo capensis (Broom, 1917); Homo sapiens cromagnonensis (Gregory, 1921); Homo larterti (Pycraft, 1935); Palaeanthropus palestinus (McCown and Keith, 1932 in Weidenreich 1932b).

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Homo sapiens sapiens contemporary with Neanderthals are documented in Africa and across southern Asia. They coexist in some areas, especially Western Asia (Mt. Carmel, Qafzeh). Though these H. sapiens sapiens have robust faces, their skull anatomy is clearly modern. Although the face is still robust, it lacks the specialized Neanderthal buttress for large mesial teeth. The reduced bone mass of less robust H. sapiens mandibles is offset by external chin development that serves to reinforce and prevent dislocation of the mandibulary symphysis during mastication. Cranial capacities of natomically modern humans range from 1000 cc to 2000 cc, but generally are above 1,200 cc. The calvarium is also changed transversely. Maximum width of the calvaria is now high upon the parietals where a parietal eminence marks a ballooning of the middle and posterior cranial fossae. This change in the shape of the endocranial space suggests that neurological changes are occurring in the human lineage that are not adequately measured by estimates of endocranial volume.

The supraorbital torus has three separate components; 1) a medial glabella torus; 2) an orbital torus immediately above and central to the orbit; and 3) a lateral torus, the superciliary arch. Though any combination or absence is possible, it is rare for all three components to be represented to form a continuous supraorbital torus. The face, especially the maxilla, tends to be reduced in robusticity. A consequence of this reduction is the presence of depressions (canine fossa) below the maxillary portion of the orbit. The essentially modern postcranial skeleton encompasses the range of size, variability, and robusticity seen in living humans. A proximal styloid process on the third metacarpal adds stability to the joint in the middle of the hand (Marzke and Marzke 1987) and digit proportions suggest thumb opposability and precision grip.

The general anatomical trend is toward declining skeletal and dental robusticity as culture begins to take its modern role in its interactions with human biology. These more modern humans populate Africa, Europe, and Asia, replacing, and perhaps interbreeding with, older human populations in those areas. The ancient history of our species is probably the same pattern of trade, colonization, interbreeding, warfare, genocide, recolonization, and trade that characterizes recent human history. Though geographic isolation as well as the cumulative effects of mutation, genetic drift, and differing selective pressures permitted formation of regional varieties of humans, the early races of fossil Homo sapiens sapiens probably have little if any relevance to modern variation. This most recent human radiation resulted in dispersal over the Old World of completely modern phenotypes, with reduced faces, parietal eminencies, and chins. The beginnings of this radiation are uncertain, but this phenotype repopulated the Old World, ventured across the Pacific, and into the Americas. The progressive increase in current world population is an continuation of that expansion of our modern genotype.

The lithic technology generally associated with early modern H. sapiens sapiens, the Late Stone Age (LSA), or Upper Paleolithic, is characterized by the appearance of a blade technology (a blade is a long flake with parallel sides) and an emphasis on the working of bone, wood, ivory, antler, and other hard materials. Innovative use of wood or bone handles allowed production of cutting tools whose working edges were formed by microliths, tiny stone flakes imbedded into the implement. Superb Late Stone Age artists, exhibited their talents in drawings, sculpture, engravings, and paintings. Beads and non-utilitarian objects adorned clothing. One of the most interesting LSA artifacts is a piece of antler, dated to about 30,000 YBP from France, that depicts lunar phases of the moon (Marshack, 1972). Red ochre was widely collected, and may have been used for body painting. Burial of the dead becomes commonplace. The corpse was likely dressed in beaded clothing and other ornamentation. Tools and food (represented by animal bones) may be placed with the body. Red ochre was likely to be dusted over the corpse. The burial practice of dusting with red ochre extends from France to the upper caves at Zhoukoutian during the LSA.

A more important characteristic of these technologies is that they change rapidly. Styles vary over short distances and across short time frames. The explosive rate of innovation that characterizes modern humanity is evident everywhere.

Modern humans moved into coastal Australia about 55,000 YBP, and slowly penetrated into less accessible central areas. It is unlikely that one would ever find direct evidence of the very first humans to reach the Americas, but archaeological sites are numerous after 13,000 YBP. Asia, long populated by modern humans, was a source of repeated migrations across the narrow Bering Straits into the New World. The straits do not pose an impassable barrier to modern Arctic peoples since they fill with winter ice and are easily navigated by a boat during summer. During the last North American glaciation, this region would have been above sea level and accessible through an ice-free corridor about 50,000 to 40,000 YBP and again between 27,000 and 8,000 YBP. Early Americans spread rapidly across the continents, reaching Maine by 11,000 YBP and the tip of South America by 9,500 BC. As the numbers of archaeological sites increase in the Americas, many Pleistocene animals (including giant bison, ground sloth, horse, mammoth, and mastodon) become extinct. Perhaps human hunters reduced the density of easily hunted species rapidly enough that native predators like saber tooth cats took the last of the great prey and then became extinct themselves (Martin, 1982). The same drama occurred earlier in Europe as animals like the mammoth and woolly rhinoceros vanished, along with predators (other than humans) that specialized in hunting them. The great plains of North America are littered with bison or mammoth kill sites. Spears are thought to be thrown using an atlatl or spear thrower . The projectile point is often a skillfully fashioned pointed blade. The Pleistocene closes with a warming of world climate, a retreat of glacial ice, a massive extinction of megafauna, and an increasing effect of human activities on in world ecosystems.
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15 Aug 2004
Department of Department of Anthropology, College of Liberal Arts , UT Austin
Comments to cbramblett@mail.utexas.edu