Hominines. The Origin of Anatomically Modern Humans (AMH) Pliocene. Hominine Taxonomy. Late Pliocene. Late Pliocene Climates. Arguments and Evidence

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1 Hominines The Origin of Anatomically Modern Humans (AMH) erectus heidelbergensis Modern Homo sapiens neandertalensis Arguments and Evidence ergaster rudolfensis Homo habilis Millions of Years Ago habilis rudolfensis Family: Hominidae Hominine Taxonomy ergaster Subfamily: Homininae Genus: Homo erectus heidelbergensis neanderthalensis sapiens Pliocene This bipedal/climbing adaptation of early australopithecines remained stable through much of the Pliocene, despite the high levels of climatic variability Adaptation includes retention of the Miocene trait of thick enamel in the cheek teeth so that both hard and soft foods were consumable Elaboration of the pattern results of substantial regional and temporal variation and speciation Late Pliocene Around 3 mya the Isthmus of Panama emerged as a permanent separation of the ocean circulation between the Pacific and the Atlantic/Caribbean Several other planetary and continental forces combine so that by around 2½ mya there is a major expansion of the Antarctic ice sheet and major ice rafts are recorded in the Northern Hemisphere There is a critical drop in temperature about this time, along with a nearly doubling of the level of climatic variability Late Pliocene Climates 1

2 Late Pliocene, 2 Savanna habitats spread into previous woodland settings, along with increased climate variability Some hominids appear to have adapted to this change by increasing dental surface for grinding tough, drought resistant plant foods (e.g., Australopithecus aethiopicus, A. boisei, A. robustus) Other hominids adopted toolmaking, transport of resources, and dietary change including increased use of animal resources (e.g., Homo habilis) The survivor of this transition appears by mya, (Homo erectus) with substantially enlarged brains and large erect bodies Plio-Pleistocene Setting There is little average change in temperature between 2.5 mya and today Dramatic changes in amplitude of changes Breakpoint about 600,000 years ago The Ice Ages with periods of extensive glaciation in high latitudes alternating with brief warm periods The beginning of extreme fluctuation coincides with advanced Homo erectus Continuing fluctuation produces Archaic Homo sapiens Plio-Pleistocene Climates Homo neanderthalensis Homo heidelbergensis Homo erectus Homo habilis/rudolfensis Less Ice More Ice Pliocene Lifecycles Australopithecus afarensis and earlier forms appear to have developed much like modern chimps, with a brief adolescent spurt Australopithecus africanus may have had slightly extended infancy and childhood and an increased adolescence Homo habilis adds a childhood phase, and probably increased adolescence Pleistocene Lifecycles, 2 Alternate Lifecycles Homo erectus increases the childhood phase, with a pronounced adolescent spurt Homo heidelbergensis would have increased childhood and a pronounced adolescent spurt Possibility that the extended childhood is beginning to play a role in transmitting communication skills Homo sapiens recently adds an early maturation to start adolescence 2

3 Homo habilis KNM-ER 1470 Fosils named handy man because of the suggestion of evidence of tools found in contemporary deposits Lived between about 2.4 and 1.5 million years ago Similar to australopithecines in many ways Face is primitive, but projects less than in A. africanus Homo habilis, 2 Back teeth are smaller than those of the australopithecines, but still considerably larger than modern humans Average cranial capacity, at 650 cc, is larger than in australopithecines Brain size varies between 500 and 800 cc, overlapping the australopithecines at the low end and H. erectus at the high end Homo habilis, 3 Brain shape is more humanlike Bulge of Broca's area, essential for speech, is visible in one H. habilis brain cast, indicating it may have been capable of rudimentary speech H. habilis was about 127 cm (5'0") tall, and about 45 kg (100 lb) in weight, although females may have been smaller Homo habilis, 4 Habilis has been a controversial species Some scientists have not accepted it, believing that all H. habilis specimens should be assigned to either the australopithecines or Homo erectus Many now believe that Homo habilis combines specimens from two different species Homo habilis--more ape-like (OH 62) Homo rudolfensis--the more human form (KNM-ER 1470) Homo ergaster The early African forms, including the Turkana boy (The KNM-WT 15000) Very modern post-crania, except for conical rib cage Large body size 3

4 Homo erectus Out of Africa Appears in China, Java close to 1.8 mya Increased encephalization African/European sites show hand axe technology Relatively stable forms mya Asian sites have chopper tools H. egaster expansion into Asia predated the development of the Acheulean tradition in Africa Homo heidelbergensis Advanced Homo erectus Best documented at Spanish site of Atapuerca Originally named Homo antecessor by Spanish discoverers Transitional form showing a mix of primitive (H. erectus) and advanced (H. sapiens) features Late Pleistocene Happenings The warm peak of the last interglacial (ca. 120, ,000 years ago) suspiciously coincides with the consensus date estimate of several genetic studies for the origin of modern Homo sapiens The last refinements on language and improvements in technology may have occurred and been a substantial cultural advantage in this time of extreme climatic change The final expansion of childhood to allow the perfecting of language skills may have accompanied this change Consensus date of genetic models for appearance of Modern Homo sapiens Recent Climates The Models Recent African Evolution Model (Complete Displacement) African-European sapiens Model (Partial Displacement) Multiregional Evolution Model (Regional Continuity) Recent African Evolution AMH populations first arose in Africa within the last 200,000 years These populations spread from Africa to the Middle East, Asia, and Europe Pre-modern populations in Asia and Europe were completely displaced by the African migrants 4

5 Recent African Evolution--2 Denies transitions from archaic Homo sapiens to AMH anywhere in the world except Africa Most extreme form considers the appearance of AMH a speciation event, complete with reproductive isolation from pre-modern populations Recent African Evolution--3 Proponents on the fossil side: Stringer, Brauer, Andrews Proponents on the genetic side: Cann, Wilson, Stoneking African-European sapiens Transition from archaic Homo sapiens to AMH in South Africa about 100,000 years ago Environmental and climatic conditions cause migration of AMH from South Africa Moving into Eurasia, AMH hybridized with resident archaic populations, eventually replacing them African-European sapiens--2 The disappearance of of archaic humans was due to both hybridization and replacement Involved a gradual and complex process Still emphasizes replacement Proponents: Brauer, Lieberman, Jackson Assimilation Accepts an African origin for modern humans. Emphasizes gene flow, without major population movement. Also invokes changing selection pressures in different areas resulting in directional morphological change. Assimilation--2 Accepts substantial continuity within major regions. Proponents: Smith, Trinkhaus 5

6 Multiregional Evolution Some local populations in Europe, Asia, and Africa continued their indigenous evolutionary development from archaic Homo sapiens to AMH Denies a solely African origin for all AMH Emphasizes the role of genetic continuity over time within major areas: Africa, Europe, Asia, Southeast Asia/Australasia Multiregional Evolution--2 Invokes gene flow between regional populations to account for similar anatomical transitions from archaic to AMH in the different regions Considers archaic Homo sapiens and AMH members of the same species Also considers the regional populations to be a single species through gene flow Proponents: Wolpoff, Smith, Frayer Genetics The Evidence Genetic distance and variability Fossils Africa, Asia, Europe Contemporary Morphology Morphological distance and variability Genetics Initial attempts to assess the phylogenetic relationships of modern human populations suggested an African point of origin Cann and her colleagues collected mitochondrial (mtdna) data from numerous populations, examined base sequence information, and produced phylogenetic trees asserting an African root 6

7 Genetics--2 Genetic variability shows the greatest amount of diversity in sub-saharan African populations High variability suggests that the African populations have been accumulating genetic mutations for the longest time--i.e., they are the oldest living populations Genetics--3 Genetic distance calculated a number of ways is greatest between African populations and other groups High variability and genetic distance results support either the Recent African Evolution or the African-European sapiens models Point to an early origin of African populations Genetics--4 Horai et al. (1995) sequenced all 16,500 bases of mtdna for 3 humans and 4 apes and found the common ancestor for the humans to have lived 143ky + 18ky Four other mtdna studies, one nuclear DNA analysis and one protein polymorphism analysis yield dates from 117ky to 280ky for separation of modern humans (see D Andrade and Morin, 1997) Genetics--4 Dandrade and Morin (1997) find no mtdna lineages in man which are exclusively Asian or European, only exclusive African lineages They conclude that the recent date of modern human separation and the deep African genetic lineage rules out the multiregional model Neandertal DNA A team of U.S. and German researchers has extracted mitochondrial DNA from Neandertal bone showing that the Neandertal DNA sequence falls outside the normal variation of modern humans. These results indicate that Neandertals did not contribute mitochondrial DNA to modern humans Neandertal DNA, 2 Current models hold that Neandertals became extinct only 30,000 years ago and co-existed with modern humans in Europe The team, however, found that Neandertals and modern humans diverged genetically 500,000 to 600,000 years ago Suggests that though they may have lived at the same time, Neandertals did not contribute genetic material to modern humans 7

8 Neandertal DNA, 3 Since 1991, an interdisciplinary project has focused on the Neandertal-type specimen found in 1856 near Dusseldorf, Germany A sample was removed for DNA analysis The researchers compared the Neandertal sequence with 2,051 human sequences and 59 common chimpanzee sequences (outgroup for comparison) Neandertal DNA, 4 Researchers looked at the Neandertal sequence with respect to 994 human mitochondrial DNA lineages Africans, Europeans, Asians, Native Americans, Australians and Pacific Islanders Results were that the number of base pair differences between the Neandertal sequence and these groups was 27 or 28 for all groups Neandertal DNA, 5 Neandertals inhabited the same geographic region as contemporary Europeans The differences between the Neandertal sequence and modern Europeans is no closer than for other contemporary human populations Neandertal DNA, 6 Phylogenetic tree reconstruction shows the Neandertal sequence branching before the divergence of the modern human mitochondrial DNA lineages, but after the split from chimpanzees The phylogenetic tree shows the first three branches of humans are African Only the fourth branch has non-african sequences The branching pattern indicates that the ancestor of the mitochondrial DNA gene pool of contemporary humans lived in Africa Fossils Earliest widely accepted AMH fossils occur in Africa and the Middle East Asian and European forms appear much later The regional features cited by some authorities as evidence of Multiregional Evolution are now thought to be retentions of robust primitive features from H. erectus or heidelbergensis, and not generally present in modern populations 8

9 Fossils--2 The fossil evidence supports the late presence of Homo erectus in Asia the early presence of AMH in Africa late arrival of AMH in Europe These results are generally inconsistent with the Multiregional Evolution model Contemporary Morphology Cranial measurements taken on a large sample of contemporary populations indicate that most variability occurs within groups African populations show greater variability (roughly three times more than Europeans) than other populations, suggesting greater time depth (Relethford and Harpending 1994) Contemporary morphology--2 Lahr (1995) interprets the variability in cranial morphology between modern populations as evidence of a recent divergence (less than ~100,000 years ago) between modern populations These findings contradict the Multiregional Evolution Model and lend support to an African origin of AMH and hybridization Contemporary morphology--3 Waddle (1994) performed matrix correlation analyses of cranial features of Old World populations to assess consistency with multi-regional vs. singleorigin hypotheses Her results support a single origin for modern humans Not consistent with long-term evolution within regions Interpretation of the Evidence Genetic, fossil, and contemporary morphology lines of evidence tend to support an intermediate model (African- European sapiens, perhaps with more emphasis on gene flow) versus the extreme models (Recent African Evolution and Multiregional Continuity) 9

10 Interpretation of the Evidence--2 Aiello (1993) and Lahr (1995) argue that all of these models are too simplistic to capture the complexity of environmental changes and associated natural selection (cooling and warming associated with major glacial events), fluctuations in population sizes and isolation or gene flow, and behavioral adaptations. Sources Aiello LC (1993) The fossil evidence for modern human origins in Africa: a revised view. Am. Anth., 95: D Andrade R and A Morin (1997) Chimpanzee and human mitochondrial DNA: A principal components and individual-by-site analysis. Am. Anth., 98: Sources--2 Frayer DW, MH Wolpoff, AG Thorne, FH Smith, GG Pope (1993) Theories of modern human origins: the paleontological test. Am. Anth., 95: Horai S, K Hayasaka, R Kondo, K Tsugane, and N Takahata (1995) Recent African origin of modern humans revealed by complete sequences of hominoid mitochondrial DNAs. Proc. Nat. Acad. Sci., 92: Sources--3 Lahr MM (1995) Patterns of modern human diversification: implications for Amerindian origins. Yrbk. Phys. Anthrop., 38: Lieberman L and FLC Jackson (1995) Race and three models of human origin. Am. Anth., 97: Sources--4 Relethford JH (1994) Craniometric variation among modern human populations. Am. J. Phys. Anthrop., 95: Templeton AR (1993) The Eve hypothesis: a genetic critique and reanalysis. Am. Anth., 95:

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