Primate Evolution (65 - 5 mya) Emergence of Primates First primates Environment - Cretaceous to Palaeocene Plesiadapiforms or adapids and omomyids? Continental drift (Pangea = Laurasia & Gondwanaland) Success of primates Arboreal theory Visual predation hypothesis (Cartmill 1972, 1992) Early Primates Prosimians (65mya) Monkeys (35mya) Apes (23mya) Hominids (5mya) Early Primates - Traits Common physical primate traits: Dense hair or fur covering Warm-blooded Live young Suckle Infant dependence Common social primate traits: Social life Play Observation and imitation Pecking order Common Primate Traits Continents at the end of the Meszoic Here are the placement of the continents at the end of the Cenozoic and beginning of the Mesozoic, about 65 m.y.a. Classification of Primates The primate order is divided into two suborders The prosimians, or lower primates include the lemurs, lorises, tarsiers, and tree shrews, while the anthropoids, or higher primates include monkeys, apes, and humans Prosimians Prosimians are generally small ranging from species the size of a mouse up to those as large as a house cat They are arboreal, have five digits on each hand and foot with either claws or nails, and are typically omnivorous They have large, forwardly directed eyes specialized for night vision, hence most are nocturnal Tarsiers are prosimian primates Prosimians As their name implies pro means "before," and simian means "ape”. prosimians are the oldest primate lineage, and their fossil record extends back to the Paleocene. During the Eocene prosimians were abundant, diversified, and widespread in North America, Europe, and Asia. • Ring-Tailed Lemurs are also prosimians Early Cenozoic Primates The earliest primates date to the first part of the Cenozoic (65-54 m.y.a.). The Eocene (54-38 m.y.a.) was the epoch of prosimians with at least 60 different genera in two families. The omomyid family lived in North America, Europe, and Asia and may be ancestral to all anthropoids. The adapid family was ancestral to the lemur-loris line. Eocene Prosimian Notharctus, a primitive Eocene prosimian; found in North America. Omomyid An artist’s reconstruction of Shoshonius, a member of the Eocene omomyid family. Anthropoids Anthropoids branched off from the prosimians during the Eocene. Anthropoid eyes are rotated more forward compared to prosimians. Anthropoids have a fully enclosed bony eye socket. Anthropoids have a dry nose separate from the upper lip. Anthropoids have molar cusps. New World Monkey New World Monkeys constitute a superfamily belonging to the suborder Anthropoidea (anthropoids) Old World Monkey Another superfamily of the anthropoids: the Old World Monkeys Great Apes The third superfamily is the great apes, which include gorillas and chimpanzees. Early Anthropoids Undisputed remains of early anthropoids date from 34 million years ago; Fayum area southwest of Cairo, Egypt Found remains from different types of anthropoids, including: Parapithecids (monkey-like) Propliopithecids (ape-like) - some believe that the common ancestor for both Old and New World monkeys belonged to this group Best known of group - Aegyptopithecus which is believed to be after the fashion of the howler monkey One of the Earliest Anthropoids Skull of Aegyptopithecus zeuxis Miocene Anthropoids Miocene Period: 24 to 5.2 million years ago First hominid appeared in Africa where remains have been found dating 5 million years old Early Miocene Period Proconsul found in sites in East Africa Middle Miocene Kenyapithecus 16 to 10 million years ago with molars resembling modern hominoids Late Miocene Apes Movement to Europe and Asia due to warmer weather conditions; migration from Africa Two main groups: Sivapithecus - link to orangutans Dryopithecus Proconsul A skull of Proconsul africanus from the Kenya National Museum. Proposed appearance of Proconsul africanus. Kenyapithecus Fossil jaw bones from Equatorius, probably ancestral to Kenyapithecus africanus and K. wickeri. Sivapithecus Sivapithecus belongs to the ramapithecid genera along with Gigantopithecus. Sivapithecus is now believed to be ancestral to the modern orangutan. A Sivapithecus skull. Gigantopithecus Gigantopithecus is the largest primate that ever lived, some standing over 10 feet tall and weighing 600 pounds. Since it died out around 250,000 years ago, it coexisted with Homo erectus. Some people believe it is still alive today as the yeti and bigfoot. A reconstruction of Gigantopithecus by Russel Ciochon and Bill Muns. Dryopithecus Dryopithecus lived in Europe during the middle and late Miocene. This group probably includes the common ancestor of the lesser apes (gibbons and siamangs) and the great apes. Dryopithecus has the Y-5 arrangement of molar cusps typical of Dryopithecus and of hominoids. Oreopithecus Oreopithecus bambolii lived between 9-7 m.y.a and spent much of its time standing upright and shuffling short distances. Its big toe splayed out 90 degrees from the other toes. Oreo is Greek for “mountain” Early Hominids Chronology of Hominid Evolution The Pleistocene (2 m.y.a. to 10,000 B.P.) is the epoch of human life. Lower Pleistocene (2 to 1 m.y.a.): Australopithecus and early Homo Middle Pleistocene (1 m.y.a. to 130,000 B.P.): Homo erectus and archaic Homo sapiens Upper Pleistocene (130,000 to 10,000 B.P.): modern Homo sapiens Hominid Evolution Homo habilis (2.0 – 1.6mya) H. rudolfensis (2.4-1.6mya) H. erectus (1.9-27kyBP) H. heidelbergensis (800-100kyBP) H. neanderthalensis (300-30kyBP) H. sapiens (130kyBP – present) Scale: Millions of Years BP The Varied Australopithecines There are two major hominid genera: Australopithecus and Homo. However, in 1992 Berhane Asfaw and Tim D. White discovered substantial remains considered to be from hominids ancestral to the australopithecines; these remains have been called Ardipithecus ramidus (thus establishing a third hominid genus) and dated a 4.4 m.y.a. A more recent (1995, by Maeve Leakey and Alan Walker) discovery has been named Australopithecus anamensis and been dated at 4.2 m.y.a. Evolution of Bipedalism Tool use and bipedalism (Darwin/Washburn) Energy efficiency and bipedalism (Isbell/Young) Radiator theory (Falk) Body temperature and bipedalism (Wheeler) Habitat variability and bipedalism (Potts) Reproduction and bipedalism (Lovejoy) Canine reduction and bipedalism (Jolly) Skeletons Comparison of human and chimpanzee skeletons. Pelves A comparison of human and chimpanzee pelves. Dentition Comparison of dentition in ape, human, and A. afarensis palates. Crania Comparison New World Monkey Great Ape Hominids show a trend toward a large and internally reorganized brain. Human An increase in brain size and organization is apparent when comparing the brains of the new world monkey, the great ape, and the human. Sahelanthropus tchadensis Discovered in Chad in 2002 and dated at nearly 7 million years, this skull is presently the oldest known hominid The Species of Australopithecus Ardipithecus ramidus 4.4 - ? A. anamensis 4.2 - 3.9 A. afarensis 4.2 - 2.5 A. bahrelghazali 3.5 - 3.0 A. africanus 3.5 - 2.5 P. aethiopicus 2.7 - 2.3 A. garhi 2.5 - ? P. boisei 2.3 - 1.3 P. robustus 2.0 - 1.0 million years ago Phylogenetic Tree Phylogenetic tree for African apes and hominids. Ardipithecus ramidus Hominid who walked bipedally 4.4 mya Discovered in 1992 by Tim White in Aramis, Ethiopia (as yet largely unpublished) Distinct enough to be a new species? ape-like dentition bipedal locomotion overall hominid-like skeleton small cheek teeth with thin enamel and large canines arm bones are hominid-like foramen magnum indicates bipedalism Australopithecines: Robust or Gracile? Two species of australopithecines: robust and gracile Most palaeoanthropologists classify robust species as Paranthropus and gracile species as Australopithecus, although both species are australopithecines. Gracile A. anamensis, A. afarensis, A. bahrelghazali, A. africanus, A. garhi smaller dentition, lighter musculature Robust P. aethiopicus, P. boisei, P. robustus larger teeth, massive jaws, sagittal crest Australopithecines Skulls of Robust (left) and Gracile (right) Australopithecines. Australopithecus anamensis 4.2 - 3.9 mya average weight - 110 pounds primitive bipedalism, possibly climbing found primarily in Kenya discovered in 1995 Australopithecus afarensis 4.2 mya, with oldest definite specimen placed at 3.8 mya apelike features (long arms, prognathic face, toothrow, brain capacity) pelvis, leg, feet, and foramen magnum all indicate bipedalism first discovered by Don Johanson in 1974 and called “Lucy” thought to be the “missing link” until A. anamensis was discovered 20 years later Australopithecus afarensis Left: Trail of footprints of A. afarensis made in volcanic ash, discovered by Mary Leakey at Laetoli. Right: Closeup of footprint at Laetoli Landscape with A. afarensis Re-creation of a Pliocene landscape showing members of Australopithecus afarensis gathering and eating various fruits and seeds. Australopithecus bahrelghazali 3.5 - 3.0 mya discovered by Michel Brunet in Bahr el Ghazal, Chad in 1995 assumed bipedalism (few post-cranial remains) A. africanus 3.5 - 2.5 mya 3.8 - 4.5 feet tall, 55130 lbs ape-like tibia, grasping big toes wide pelvis, parabolic tooth row primitive bipedalism first found by Raymond Dart in Taung, South Africa in 1925 Australopithecus garhi possibly the direct ancestor of early Homo larger molars than afarensis, but not as large as Paranthropus lacks enlarged brain of early Homo toolmaker and butcher 2.5 mya Paranthropus aethiopicus 2.7 to 2.3 mya earliest robust australopithecine, but least well known larger dentition, cheek bones, dish-shaped faces, sagittal crests resembles afarensis but with increases in dental apparatus size assumed bipedalism first discovered in Omo, Ethiopia, 1967-1974 The Black Skull - P. aethiopicus The “black skull”, dated to 2.5 m.y.a., was discovered by Alan Walker in 1985 near Lake Turkana. Paranthropus boisei 2.3 - 1.3 mya; 4.1 - 4.5 feet tall, 75-175 lbs discovered by Louis Leakey at Olduvai Gorge in Tanzania in 1959 originally named Zinjanthropus dished face, sagittal crest parabolic toothrow, wide pelvis primitive bipedalism Paranthropus robustus 2.0 - 1.0 mya; 3.6 - 4.3 feet tall; 70-175 lbs discovered by Robert Broom in 1938 at Kromdraai, South Africa, who created the name Paranthropus dished face, sagittal crest parabolic toothrow, human-like big toes, wide pelvis, no diastema for canines bipedalism (more primitive than modern human walking) Map of Australopithecine Finds Map of Australopithecus sites in Africa, with a focus on the East African rift valley and limestone caves of South Africa. Facts about Australopithecines Species Dates (m.y.a.) Known Dist. Important Sites Body Weight (Mid-sex) Brain Size (Mid-sex) (cm3) Homo sapiens sapiens modern world-wide Beijing, New York, London, Nairobi 60kg/132lbs 1,350 Pan troglodytes modern central Africa Gombe, Mahale 42kg/93 lbs 390 Australopithecus boisei 2.6? to 1.2 E. Africa 39 kg/86 lbs 490 Australopithecus robustus 2.6? to 2.0? S. Africa Olduvai, East Turkana Kromdraai Swartkrans 37 kg/81 lbs 540 Facts about Australopithecines Species Dates (m.y.a.) Known Dist. Important Sites Body Weight (Mid-sex) Brain Size (Mid-sex) (cm3) Australopithecus africanus 3.0 to 2.5? S. Africa Taung, 36 kg/79 lbs Sterkfontein Australopithecus afarensis 3.8 to 3.0 E. Africa Laetoli, Hadar 35 kg/77 lbs 430 Australopithecus anamensis 4.2 E. Africa Kanapoi Insufficient data no published skulls Ardipithecus ramidus 4.4 E. Africa Aramis Insufficient data no published skulls 490 Homo habilis 612 cc brain 2.3 - 1.6 mya first toolmaker prognathic face, brow ridge probable meat-eater possibly arboreal discovered in 1960 by Leakeys no speech Artist’s representation of a Homo habilis band as it might have existed two million years ago. Skull of Homo erectus A reconstruction of the skull of Homo erectus, a widely distributed species whose remains have been found in Africa, Europe, India, China, and Indonesia. Homo erectus 1891 - Eugene Dubois discovers H. erectus in Java Dubois calls it Pithecanthropus erectus initially, also dubbed “Java Man” finds in China called Sinanthropus dates from 1.9 mya to 27,000 years B.P. 994 cc brain size (compare to 612 for H. habilis) Acheulean tool industry Photograph of Nariokotome boy, an early Homo erectus found near Lake Turkana, Kenya. Neanderthal Skull Reconstructed Neanderthal skull The Neanderthals were characterized by prominent heavy brow ridges and weak chin Homo neanderthalensis discovered in the Neander Valley (Tal) near Dusseldorf, 1856 massive brain--about 1,400cc on average large torso, short limbs, broad nasal passages later remains show decrease in robustness of the front teeth and face, suggesting use of tools replaced teeth retained occipital torus, some mid-facial prognathism The skull of the classic Neandertal found in 1908 at La Chapelle-auxSaints. Homo sapiens Archaic – 100,000 to 35,000 years BP Sometimes called Homo sapiens and Homo sapiens neanderthalensis Modern – 35,000 years BP to present Anatomically modern Sometimes called Homo sapiens sapiens Cro-Magnon Man Cro-Magnon humans 35,000 years B.P. in western Europe to 17,000 years B.P. 1,600 cc cranial capacity Name comes from a hotel in France Not a different species, just old Homo sapiens from Europe Artist’s reconstruction of a Cro-Magnon man Archaic H. sapiens Culture Art Traces of art found in beads, carvings, and paintings Cave paintings in Spain and southern France showed a marked degree of skill Female figurines 27,000 to 22,000 years B.P. Called “venuses,” these figurines depicted women with large breasts and broad hips • Perhaps it was an example of an ideal type, or perhaps an expression of a desire for fertility Archaic H. sapiens Culture Cave paintings from 20,000 years ago at Vallon-Pontd’Arc in southern France (left) and from Lascaux, in southwest France Cave paintings Mostly animals on bare walls Subjects were animals favored for their meat and skins Human figures were rarely drawn due to taboos and fears that it would somehow harm others Upper Palaeolithic – Hotbed of Culture 40 – 10k yBP Shelters 15,000 yBP Ukraine Some made with mammoth bones Wood, leather working; carpentry Tools From cores to blades Specialization Composite tools Bow and arrow Domestication of dogs Gathering rather than hunting became the mainstay of human economies. Top: Straw Hut Left: Mammoth bone hut Bottom: Tool progression Modern Homo Sapiens Regional-Continuity Model (Milford Wolpoff, UMich) Humans evolved more or less simultaneously across the entire Old World from several ancestral populations. Rapid-Replacement Model (Chris Stringer, NHM London) Humans evolved only once--in Africa from H. heidelbergensis ancestors--and then migrated throughout the Old World, replacing their archaic predecessors. Also called the “Out of Africa” and “Killer Ape” hypothesis. Social Organization Hunter-gatherer analogy Small group, low population density, nomadism, kinship groups Migration North America was the last colonized by hominids. Beringia (land bridge) between Russia and Alaska Asian origin of Native Americans 30,000 to 12,000 years B.P. was first migration