March 15th – Early Stone Age Important slides: 9, 40 – 42, 45, 47, 52, 55, 57-58, 64, 65 ,67 Moving from Chimps to Foragers (differences) Hunting 20-50% of diet; High mobility (foragers); Fluid Social Groups Chimp Culture- “learned” variations. (e.g. shaking hand over their head when they greet) Chimps use simple tools (leaves, sticks, and nut cracking with hammer and anvil) Complex tool manufacture begins LP/ESA- Oldowan (2.6-1.8mya) and Achuelean *References to “stone age” are specific to Africa; no MP in Asia* RAYMOND DART: Taung baby = first hominid from Africa. Limestone cave. Fossils found in Sterkfontein indicated carnivorous australs WRONG- ignoring the site formation processes (Geological processes, burrowing animals, weathering, scavengers, people’s interactions) Earliest Stone Tools found in Ethiopia- Gona- 2.6 MA Early Olduwan Sites: Small, Stones and bones, Complex processes of accumulation, Hominids and carnivores Glynn Isaac Koobi Fora; Oldowan Industry/Technology; hitting rocks to make flakes and core Typology: Choppers (unifocal/bifocal) cores w/ flakes knocked off. Core forms: (discoids, polyhedrons) choppers w/ multiple flakes knocked off. Flakes: (heavy duty, light duty) Spheroids (stones pounded until they’re smooth and round) – meat tenderizer/come from boredom Majority are flakes. Bone Tools: Digging and Puncturing; fossils, tubers, termites Olduvai Swartkrans --- 2.2 mya Sterkfontain Invariant for 1million years Stone tools too hard to make for chimps Marrow extraction – percussion marks; meat eating – cut marks Cores & Flakes seemingly show right handed dominance Experimental Microwear Analysis (Koobi Fora) 16 % of flakes found were used. Distribution of use: 33 % wood 44 % meat 22% grass or reeds Glynn Isaac-Central Place Foraging hypothesis (site near water) Hunting / Efficient scavenging “chimp” (Bunn and Knoll) mostly legs and jaws Mid-shaft cut marks for de-fleshing Prime-age adult prey Therefore: central place for processing of hunted carcass Skeptics Legs and jaws best-preserved b/c of density mid-shaft cut marks rare for de-fleshing Lots of carnivore marks Which species were in the Olduwan (2.6-1.8mya)? Who was around at the time and who was smartest (based on EQ = H. habilis and A. rudolfensis, bc highest EQ = 3) Manual Adapt.: Power grip/ Precision grip - Chimp not well adapted for precision grip Fire/Cooking Burned patches (oxidation); Burned bones: Swartkrans (fire is natural to any/ all environments) No real evidence but Absence of Evidence ≠ Evidence of Absence Also, site shelters Acheulean & Developed Olduwan – 1.7mya – H. erectus/ergaster Kenya – handaxes, flakes and cores – butchery sites possible The Developed Olduwan: Core – Flake, lots of tools (no hand axes) Movius line – no handaxes above this line, so only Africa, ME and S Europe Olduwan – Acheulian Transition hand axes not everywhere increased cognitive complexity (a little) more variety of forms/ more stereotypical more hunting? Fire? March 22nd – Earliest Homo (H. habilis and rudolfensis) Key Slides: 3-10, 16-17, 20, 26-27, 31-33, 39-40, 42-44, 47-49, 51, 56, 59, 64, 67 4.5-7mya: Sahel./Ard. Gracile austral.; 2-4mya: Gracile robust and H. sapien Transition from austral. to Homo erectus (2mya) *Homo - bigger brain and making stone tools Body mass and brain size increases; Longer legs and shorter arms; Smaller faces and teeth Stone tools 2.6 mya. Foragers/carnivores, food sharing and meat eating (homo) Lifestyles: Early hominid diet + scavenged food Small groups with some food sharing and male provisioning Tool-making History Leakeys – Olduvai Gorge – looking for stone tools (1959) Zinjanthropus boisei – earliest responsible for Oldowan tools? found with many bones and stone tools (1964) Homo habilis (“handy man”) different teeth; much larger brain size; hand bones suggest tool-making *H. habilis = intermediary between austral. and H. erectus – more bipedal, more dextrous and intermediate brain size* Fossil Record 1970s – Koobi Fora. --larger brains; more finds aged around 2mya “OH 62” fragmented skeleton and complete palate – only early Homo with postcranial Palates: shorter and more vertical premaxilia giving rounder, more parabolic tooth row 1) Early Homo from Omo, Ethiopia 2.2 mya 2) AL 666 – Hadar, Ethiopia 2.35mya “early homo” stone tools 3) A. garhi, Bourhi, 2.5 mya 3mya – NO stone tools One species or two? “Garbage can” taxon – lumping together anything. Prob.: too much variability w/in taxon Ie. Diff species or genders? – KNM-ER 1813 and KNM-ER 147 (dimorph.) How do we decide between 1 or 2 species? 1) Analogue species to compare amt of variation 2) Probability of attaining same level of variation Conclusion: 2 species – H. rudolfensis and H. habilis: Brain size: r > h Face: r = austral., h = homo-like Teeth: r = austral., h = homo-like (r has larger, flatter postcanines) Early homo postcranials (rarity of associated postcranials and cranials) --Hand and foot from Olduvai Hand with expanded thumb but curved phalanges Foot is modernized; toe is in-line (adducted), arch --H. habilis palate with very small post-crania Big bodies with more homo-like morphology of teeth and skull --Dimorphism like austral. (more than Pan) --Long forelimbs, short hindlimbs Tooth size was larger than humans and brain size was smaller than humans Homo or australopithecus What genus does it belong to? 1) group of species with common ancestry (clade) H. erectus H. habilis H. rudolfensis Australopithecines Bigger brain Distinct browridges Smaller, narrower face and teeth 2) similar adaptive pattern (grade diet/subsistence: H. habilis smaller teeth like H. erectus locomotion: both Rudolf and habilis austral-like brain size: habilis austral-like (less advanced tech. and cognitive abilities) life history pattern: all austral-like Conclusion: Habilis and rudolfensis were more austral like than homo like in overall behavior Might include both in austral. BUT: no solid postcranial data Climate How did climate change influence speciation? Deep sea cores (%age of O18) Over last 6mya: cooler, more seasonal (fluctuations) Causes: continental drift (unpredictable phenomenon) Predictable phenomenon (solar radiation - changes in orbit, tilt, etc. Milankovich cycles) Temperature fluctuation progression toward savannahs and grassland Climate and habitat change evolutionary consequences Forest fragments and becomes extinct Forest fragments and diverges into two new populations of former species ** speciation** April 3rd – Homo erectus Slides 16-20, 28-9, 31, 40-49,56, 58 -65, 67-80 History: H. Erectus unique Bigger brains and bodies and smaller teeth H. Erectus focus on ONE species throughout Africa, Asia, and Europe (H. ergaster was only erectus in Africa) (1890) Pithecanthropus erectus, first H. Erectus discovered Indonesia: Pithecanthropus, China: Sinanthropus pekenensis Ernst Mayer lumps these together into H. erectus (1984) Nariokotome Boy (H. ergaster), Lake Turkana Evolutionary origins/context: Major Transition: Body size increase, Post Canine reduction, EQ increase, more terrestrial o Correlation with cognitive complexity, life history elongation, meat in diet, and more efficient trekking o Erectus – big shift from even habilis and rud. (larger brain/EQ, teeth smaller, bigger body) Pleistocene Dates (Ice Age) – the environment of evolutionary adaptation: Lower: 1.75Mya-780 KA; Middle: 780-125 KA; Upper: 125- 12 KA Earliest 1.9mya Koobi Fora, Kenya and Dmanisi, also, as recently as 800Ka in Algiers Asian Populations (migration from Dmanisi to S and SE Asia): Indonesia Population Specimens: o Sangiran, Trinil, Modgokerto: 1.6MA-470 KA H. Floresensies: The Hobbit 90-13 KA China: Zhoukoudian: 350-500 KA; shovel-shaped teeth as Asian characteristic Narmada, India: MP; Different Cranial Vault Shape Little difference between African and Asian specimens Most likely ancestor for all of these populations: H. Habilis Crania, Dentition and Postcrania: Crania less than 1.2 MA Largest is H. erectus w/ EQ = 3.3 larger EQ is Homo sapiens = 6.3 Female: 630 CC; Male: 810 CC; ID: 1.3 (small) Long, low vault; sagittal keel; prognathism; large face and orbits; browridges (dimorphic), more balanced head o Huge Browridges: Sexual Selection EXTERNAL NASAL MARGIN: Air Humidification, breathing VERY THICK VAULT – systemic robusticity v. veinous cooling(?) Small dentition, shovel-shaped incisors: Yuanmou, Nariokotome Postcrania 1) Body Mass between 60 - 70 KG Possible Reasons; Lower quality food, intraspecies competition, predator defense, life history, increased efficiency of transport 2) Much shorter arms; shorter distal limbs (brachial and crural indices) Eco-geography: Climates Also because not swinging from trees Surface area/volume lower in colder climates In tropical regions: Longer body, taller body sizes 3) Relatively smaller digestive tract in H. Erectus Thoracic shape: More cylinder-like 5) Pelvis: more expanded (more efficient locomotion) and bigger joints (loading) KEY POINTS: H. Erectus first seen 1.9 MA and spread quickly into Eurasia Bigger, more modern bodies, larger brains, smaller faces, teeth April 5th – Why Homo erectus? Slides: 6, 7, 12-15,16, 19,20, 24, 32, 42-58, 65, 75,76,86, 87, 91-97, 100,101 Phylogenetic Context: Long, lived species with little change Big change around 2mya Conditions: Before: Ape-like, woodland biped, simple and primitive, little technology After: cognitive abilities, language, hunting , slower IBIs and maturation Signature tools of the LP Oldowan - Sharp-edged flakes = access to meat, brain, marrow, foods, etc.; no projectiles or throwing tools (2.6mya) Acheulean - bifaces (1.7mya) Migration H. Erectus was first hominid species to leave Africa (Dmanisi, Indonesia) Global climate fluctuations habitat fluctuations more open habitats Rapid spread of H. erectus because of the open habitats Interglacials: forests expand, deserts shrink Glacials: Forests shrink, savannahs expand ENERGETICS AND REPRODUCTION H. erectus more like australopiths and chimps (IBI etc.) Higher body mass + energetic costs of reproduction = need for more energy/food Supplemental feeding less of a problem, but teeth are smaller/thinner o Weaning at age 3, very similar to human foragers (chimps = 5yrs) Meat-eating 2.6 MA; 1.8MA = medium to large animals (hunting tech. improves) Early Stone Age: --No distance projectiles; close weaponry, endurance running o Middle of day = peak heat, run at non-optimal speed, drive prey into hypothermia and track o Narrow waist, long legs, arches, Achilles, narrow thorax = benefits o Key adaptations for stabilization: Big gluteus muscles; head stabilization – foramen magnum orientation; long, detached necks, decoupling of head and pectoral girdle (independent motion) o Thermoregulation sweating and loss of fur; Trekkingexternal nose Persistence hunting (+): Minimal risk/cost, minimal technology, High yield Persistence hunting (-): Hot (seasonal?), need water, salt, physical fitness, speed, open habitat, social system, food processing Modern forager pirating: Vulture Run Fend off competitors Running requirements: brain increase, sharing, running, meat-eating, provisioning -- more societal-oriented activities develop with H. erectus April 10th - ARCHAIC HOMO Slides: 4, 5, 9, 10, 12-13, 19, 21, 24, 26, 52, 57-60, 62-63, 72-75, 79-81, 96, 97 (splitters) H. erectus – 2.0 to 0.5mya – Kenya, Georgia, Indonesia Basically bigger brains and faces Low, thick cranial vault; robust postcranial ( > modern humans); large orbits Smallish teeth Big bodies, non–arboreal running Substantial climatic variability within locations – cold and wet, then mild, then cold and dry Late early stone age - handaxes etc. Africa Middle East Asia (India) Indonesia *fossils beginning to look like Neanderthals – nose, chin, cranial vault, flared ilia; wide birth canal. Avg male height 5’7’’; 50% died between 10 and 18, no one older than 40 (like Pan.) Traditional grade view: H. erectus “Archaic” homo H. sapiens Multiregional origins – H. erectus 1.8mya spreads to Europe, Africa, Asia, Australasia ^^^Both of these are no longer used ^^^ More clade-based view: “replacement”/ “African origin” Homo erectus evolves and moves out into other regions; modern humans develop in Africa (only), branch out into other continents and replace A. homo H. sp. indet – LCA between some Homo and us (closer than erectus) lots of variability post 0.5mya – not clear how to sort them (prefer splitters) Lieberman’s preferred splitters cladogram: Homo sapien H. helmei H. rhodesi H. neand. H. heidel Homo erectus Note: H. helmei, H. rhodesiensus, H. neanderthalensis, and H. heidelbergensis are all part of H. species indeterminate Examples of Archaic Homo: 1) Bodo (Ethiopia 500ka), Broken hill (Zambia), Jinnuishan (China) – female skeleton 2) “The Hobbit” - Flores, Indonesia 80 Ka built raft?; a few primitive tools skull 380 cc (smallest brain size of any hominid); 1m tall; tiny body, long arms; body proportions much like Lucy INSULAR DWARFING? (see notes from last lecture) – reversal of size; microcephalic? 3) Dmanisi (1.8mya) H. erectus/H. ergaster “The Earliest Europeans” Gran Dolina, Spain 800-900ka - H. antecessor – juvenile Cut marks on bones and defleshed – cannibalism? Sima de los Huesos – 300-400KA; cave complex – 2000 human specimens – 24 individuals – burials? Mauer, Germany – H. heidelbergensis – a lot of mandibles April 12th – DMANISI* Slides:1, 5, 10, 11,9 (about erectus); 13, 14, 18, 22-24, 29-30, 33-38, 40, 42 (Dmanisi) *note: this lecture is NOT on the test, but the information is probably still important Significance of Dmanisi -SE Georgia earliest traces of human occupation outside Africa -Animal bones (tooth and cut marks), stone artifacts, five hominids (skulls and postcranial) -medieval settlement (1200-1300ad) reveals early Pleistocene animal bones -stone tools derived from volcanistic material (simple, crude toolds and cores) -Masavera Basalt (1.85mya) ash and sediment accumulation after lava cooled -hominid metatarsal discovered 2001 (1991) mandibular proportions (thick) and tooth crown dimensions (small) resembles H. erectus Similarities between Dmanisi and earlier Homo and/or Austral. Small cranial capacity; Postorbital constriction; Supramastoid cresting; Transverse torus absent; Mandibular fossa proportions as in earlier Homo; Midfacial contour projecting; Hard palate shallow rather than arched D3444 – completely toothless – older individual (possible male) survived for several years in an impaired state *see slides 33-35* **consider all specimens as single population – intragroup variation in early Homo Neanderthals – April 17th Slides: 12, 13, 18, 22-39, 41, 42, 44, 45 Why care? Top of skull found in 1856 by J. Fuhlrott in Feldhofer Cave – very controversial First fossil found in Belgium in 1829 Busk - skull – 1848 - Gibraltar, revived interest in 1863 William King said Neanderthals were new species in 1864 Evolutionary context Acted like a mirror to paleoanthropologists - they're like us but not Extreme views: they're stupid v. they're just as smart as us o Debate about neanderthals has often been about people's psychological extremes Definition Restricted in time & space: MP – 200ka to 27ka in Europe and Middle East First time we see them during an interglacial (warm) period in Europe o Glacier covering northern Europe, further south was tundra (barren) Replacement view of human origins – Neanderthals developed from H. erectus that left Africa and was then replaced by H. sapiens Lumpers and splitters clade view (see archaic homo for cladogram) Evolved in europe, but did not give rise to modern humans Anatomy: SKULL: Shared derived features from archaic homo and heidelbergensis: long, low, thick vault, long mandibular ramus, small mastoid, no chin, big browridge, big orbits, no canine fossa, convex face (resistance to bending), occipital bun Unique derived features (apomorphies): occipital bun, small mastoid process, retromolar space, big external nose (humidifying the air), midfacial projection (resistance to twisting) BRAIN: Higher EQ than modern humans, Cro-Magnon even bigger; larger brains relative to body size, but body size less than ours TEETH Shovel-shaped incisors (primitive), Taurodont roots (derived) Ear: posterior semicircular canal (diagnostic feature different from modern humans) POSTCRANIA More barrel-chested More flared pelvis Longer pubic bone Shorter distal limbs (cold) Robusticity (declines very recently) Key Fossil Sites Atapuerca - Sima de los Huesos (northern Spain), Boxgrove, England Homo sp. Indet. From Europe = H. antecessor, H. heidelbergensis Heidelbergensis in Greece and Spain show derived features from Nean. Multiple dispersals from Africa (accretionary model) Immigration to Europe then 3 periods of extinction; during interglacial periods (1ma, 500ka, 300ka) Neanderthals are a European species Few early fossil location in Italy, Croatia, France Golden Age: 55-75 Ka Terminal Phase: <50-27 Ka o Youngest Neanderthal found at southern tip of Spain 27ka Neands and modern humans (older than neand) found in Middle East (Tabun) There are no neanderthals in Asia or Africa Life as a Neanderthal Life History o 4 year-old molar eruption and brain size comparable to a modern human of 5-6 years; grew up like modern humans o Tooth wear starts about 3 years o Enamel defects peak at 3.5 years o Hypoplaisas (undergrowth) in teeth suggest that weaning begins about 3-3.5 years o Growth differences before age 2, very different trajectories from modern humans though rate of growth is similar Brain size by taxon Language: Fossil record - ability to produce speech Hypoglossal nerve – ability to make vowel sounds Miscellaneous: Levallois Core Anterior dental-wear indicates eating large-herd animals (reindeer etc.); ambush hunting High resistance to bending and twisting Cannibalism, busted up skulls Replaced by modern humans, little interbreeding Origin of H. Sapiens –Fossil Evidence April 19th Slides: 3, 6-10, 14, 18, 21, 25, 28, 34-35, 44-46, 71, 77, 86-90, 104 Paleontological Climatic context of Pleistocene o Every 100K years, cycle of cold and warm periods, habitats moving o 100-200K, key events occurred for our species' evolution Major changes from old view (multiregional) to Replacement model o Genetic evidence o Dating technology o Impact of cladistics (shared derived characters - cladograms) Replacement View (now dominant) o Homo erectus gave rise to Homo Sapiens: Erectus leaves Africa 1.8mya and H. sapiens leave Africa 200Ka replacing Neanderthals What makes 'anatomically modern humans' modern Cranium (Archaic Homo from Broken Hill vs. Cro Magnon) o Globular braincase o Steep forehead o Small, retracted face o Divided browridge o Small orbits o Canine fossa o Chin – resorptive field at the top, depository at bottom of mandible Primitive features - not derived (cranium) o Big brain o Vertical face o Variable browridge Postcranium (derived features from neanderthalensis) o Shallower thorax o Lass flared pelvis o Shorter pubic bone o Longer distal limbs *Robusticity* Distribution (time and space) Predictions: multiregional vs. recent African origin o Oldest fossils support african origin: S. African and Ethiopia (~200Ka) o Regional features recent (except Africa) o First modern Homo sites (Ethiopia- Omo, Herto) Behavioral Modernity in Africa – bone points, pigments, blades, microliths MP of Levant (Turkey etc.) key site = Tabun, Neanderthals 60-80Ka and modern humans 80-130Ka – means modern humans there before Neanderthals – see next lecture for explanation; also Levallois core – MP modern humans and neanderthals Homo in Asia, Europe, Africa H. erectus leaves Africa 1.8mya, into Asia spread btwn 1mya and 100ka. H. sapiens reaches Asia 40-60ka o Asia - Zhoukoudian (250-500 Ka); Shandingdong (60ka) o Europe – H. heidelbergensis 900-200Ka, Neand. 200-27Ka o Australia 40-13Ka Modern humans were tall and thin b/c evolved in warm climate, Africa Cultural capability in MP – Levallois core, spear points, scrapers Developmental bases of human origins (Evo-Devo) evolutionary developmental biology o Developmental shifts to transition to new species o Major shifts in skull architecture? Globular vault Facial retraction o Morphometric analysis: where does growth differ? o Problem of integration – more than 1 gene contributes to bone growth Temporal lobe - part of the brain that got bigger in human evolution (speech) 3 major shifts: 1)Relative facial shortening in H. sapiens, 2) longer anterior cranial base, 3)flexed cranial base Regional continuity What would intermediates/hybrids look like? o Gradual event: intermediates should be present o Mosaic idea - independent heritable traits o Problem: most traits are polygenic (integration problem) Regional features are recent - similarity and overlap in skull shape April 24th - Origins of Modern Humans: Genetic Evidence Slides: 13-17, 18-19, 20-24, 25, 27, 29, 31, 36, 37, 43, 44, 48, 55, 59, 60, 67, 70, 79, 85, 88-91 Molecular clock – DNA mutates at a certain rate can look at differences and determine how far back they split (relatedness) using non-recombinant DNA (Ie: how long ago Mitochondrial Eve left) - DNA A’s bind to T’s and C’s to G’s; 23 human chromosomes, #23 is sex chr. - Central Dogma – DNA RNA proteins - Codon = 3 base pair long coding for an amino acid, building block of proteins - Most of genome unexpressed (junk DNA) - 2 types of DNA in cells: nuclear and mitochondrial (mtDNA) Nuclear Mitochondrial Y-chromosome Inheritance pattern Biparental Maternal Paternal Recombination Yes No No Mutation rate Slower Faster slower - 5 types of mutations – point, inversion, translocation, duplication, deletion - Haplotypes –alleles (gene variations) correlated with each other in diff. populations w/ identifiable groups of base pairs; look at how many alleles correlate in different parts of the world; All haplotypes exist in Africa, but only a few of them traveled to different parts of the world (Ie: one in Asia, one in Europe but they are both in Africa) – see slides 43 and 44 o Can create a tree based on these haplotypes, mutations around them o Use parsimony (simplest rship possible to form cladograms) o DNA hybridization – more energy to break up the bonds; can determine relatedness, more related when more energy needed - Genetic evidence can be used for dating (molecular clock) - Multiregional model: erectus left Africa, went to other continents and evolved into H. sapiens via DNA crossover - Recent Africa Origin Model (AKA “Out of Africa”) – erectus left Africa, went to other continents, (inter)glacials temporarily blocks migration, sapiens evolved from erectus in Africa, spread out over the earth, erectus lineages die out, confirmed by little interbreeding - Hybridization Model – same as out of Africa, except suggests erectus-sapien interbreeding, erectus-sapien hybrids created and genes survive: no evidence - Population bottleneck – small erectus population changing into H. sapiens, so limited variation; speciation – all genetic variation in world is in Africa - Humans have low variation; variation highest in Africa (bottlenecking) - Fst = relative amount of between group variation relative to mean o Low Fst (0) means most variation within populations o High Fst (1) means most variation between populations - Searching for a mitochondrial Eve through cutting mtDNA, 290k – 140kya - Tree – no racial grouping, racial clusters all spread out - Coalescence time: upper time limit for pop. div. (LCA); collapses if bottleneck – varying levels of genetic variation, less variation reduces divergence window - Modern mtDNA is very different than Neanderthal mtDNA - DNA primers (cutters) only cut Neanderthal DNA, not human DNA = evidence for profound genetic differences - Date of human/Neanderthal split = 370kya (acc. to nuclear DNA) o LCA population size = 3,000 o Origin of humans probably slightly older than LCA of humans and Neand. = meaning that humans were in existence before Neanderthals o No interbreeding Conclusions: 1) Humans recently rated (150Ka); 2) Origin of species a little older than LCA; 3) Multiregional model incompatible with genetic data; 4) no evidence for hybrid. April 26th – Archaeology of Modern Humans Slides: 3-5, 6, 8-10, 14, 16-20, 22-33, 50, 54, 55, 57-60, 62, 63, 66, 72,78 - bias in archaeological record – some behaviors/environments are better preserved - MP – Levallois Cores, little variability in time/space, no art - UP – blade cores (twice as long as wide), lots of variability (cultures), art present - 2 theories: UP developed in Europe vs. UP imported from Africa (evolution v. revolution) o Prob w/“evolution” theory – Neanderthals used UP tools, Chatelperrionian o Confusing data from Middle East (Ie: Tabun) and all made MP tools o No correlation btwn archaeological and fossil records; none necessary tho - Bar-Yosef’s Explanation (accepted) – based on Tabun o Stage 1 – (>100Ka) Neands and MidE. sapiens from Africa – MP tools o Stage 2 – (60-80Ka) Europe - glaciers push Neands to MidE, pushed out H. sapiens o Stage 3 – (60-45Ka) Neands. back in Europe w/ MP tools; Sapiens bacl in MidE w/ UP tools from Africa o Stage 4 – (45Ka) sapiens to Europe w/ UP tools displace last neanders *explains layers of Neanderthals and h. sapiens and why Neand were found in Spain* - MP (in Africa = MSA): o prepared core; core-flake technology; Levallois technique - make a core with a flatedge, flake off and retouch o 1st evidence of controlled fire hearths; 250Ka o NO MP IN ASIA - UP (in Africa = LSA) o art/symbolism/intentional burials o Blade core, microliths and Levallois from MP o Aurignacian, Gravettian, Solutrean, Magdalenian o Atlatl, bows and arrows, more complex diets, diversity of food sources o Population density higher in UP, open air sites, elaborate graves Explaining the UP; Cognitive revolution hypothesis (people got smarter suddenly) or African evolution hypothesis FIRST HOMO SAPIENS IN AFRICA THE SAME TIME AS FIRST NEANDERTHAL - Gradual transition to UP in Africa 1st migration out of Africa around 100kya during MP times 2nd migration from Africa 50kya, brought UP tech with them Displacement after 45k but gradual; 30K wiped out - What happened when neanders encountered humans? o Humans hunted/killed neanders o Humans better suited for environment o Neanders interbred with humans (no evidence) - Time of rapid climate change Transition from interglacial to glacial = open path to Europe to closed path - Humans weaned earlier in UP than in MP - Chatelperronian – UP industry from 35-30k ya Lieberman Neands were outcompeted and went extinct Derived characteristics Homo erectus Cranial Increased cranial capacity Big, vertical face; external noses Smaller teeth Long, low thick cranial vault Large browridges and orbits (sexually dimorphic) Sagittal keel Occipital torus Postcranial Tall 5 feet, 10 inches More sexual dimorphism Barrel-shaped thorax Rel. short arms Tall, narrow waist Rel. narrow hips Rel long legs Rel long tibia Rel large lower body joints More robust bones *Archaic homo = larger Homo erectus; bigger face and brain size (lecture17, slide63) UNIQUE - larger brain size; earliest hominid to have a brain as large as (relative to body mass) modern humans long, low cranial vault large orbits browridges separate SHARED - external nose knee joints --> supporting large body mass --> long distance trekking/hunting general robusticity in bones thick ramus in mandible --> resistance to twisting/bending small teeth Neanderthals (Diagnostic in Bold) Skull Occipital bun Small mastoid Retromolar space Big nose Midfacial projection (resistance to twisting) Convex face (resistance to bending) Taurodont roots Posterior semicircular canal Body Shape Barrel-chested (more than erectus???) Flared pelvis Longer pubic bone Shorter distal limbs Robust bones Homo sapiens Cranium Globular braincase, Steep forehead, Small, retracted face, Divided browridge, Small orbits, Canine fossa, Chin Postcranium Shallower thorax Less flared pelvis Shorter pubic bone Longer distal limbs Lab Summaries Lab 1: Biomechanics Primer Shape/size of bones, joints, muscles indicates natural selection and applied forces Bone muscles systems - input force (Fi) = force being applied (pull of tendons, gravity, external load) - output force (Fo) = force derived from motion (teeth, feet, digits) Lever Arms and Torques - Feeding and locomotion systems levers - Joints usually the center of rotation (COR) or pivot - In-Lever arm (Li) = goes from Fi to COR - Out-Lever arm (Lo) = goes from COR to Fo - Force x lever = turning force/moment/torque o FiLi > FoLo – lever rotates toward Li o FiLi < FoLo – lever rotates toward Lo o FiLi = FoLo – no motion (equilibrium) st - 1 class lever systems – forces on opp. sides of COR (extensor, triceps), 2nd class – COR, out, in, 3rd class – COR, in, out (flexor muscles – biceps) Mechanical Advantage, V, and range of motion - increase out-force by increasing length of in-lever or decreasing out-lever - Mechanical advantage = Li/Lo o Higher mech adv means less in-force to produce equal out force - Velocity – Lo/Li determined by distance from pivot and rate of turning o Vi = VoLi/Lo and vice versa - tradeoff between force and velocity (running adaptations) Range of Motion - range and rate of motion increase with proximity to pivot point Application example - COR = temporo-mandibular joint (TMJ); Fi = masseter; Fo = teeth; Li = distance from COR to point of action of masseter; Lo = dist. From COR to bite force o On a face out lever arm lines up with snout angle/direction - Gorilla has higher out-force = more chewing potential; Shrew has higher velocity rate = faster chewing Lab 2: Comparative Anatomy Cranium/Mandible: Size of face relative to braincase – size order: humans, monkeys, apes Browridges – face grows anteriorly, face tucked under braincase humans so no browridges Foramen magnum position – monkeys and apes have a posteriorly placed foramen magnum – humans anteriorly b/c of bipedalism and balancing over center of gravity Prognathism – longer jaw associated with greater range of motion, orthognathic animals have greater MA in many chewing muscles Sagittal Crest – fxn of temporalis muscle size relative to braincase; sexually dimorphic Temporal fossa – size related to size of temporalis muscle – chewing Mandibular Corpus Depth/breadth – deeper corpus bending, breadth of corpus twisting Dentition: 2, 1, 3, 3, pattern Diastema – space to accomadate lower canine Incisor size – related to degree of initial food preparation and manipulation – surface for biting and holding food; apes and monekys large, humans small Canine fossa – male/female dimorphism male-male competition Pre-molars – blades and cusps in apes, flat and molariform in humans Molars – 4-5cusps; small and cuspy in monkeys and apes, large and flat in hominids – relate to food quality Enamel thickness – diet quality (amt of fiber)– monkeys and apes have thin enamels; humans – thicker Lab 3: Australopithecine Cranial Morph. And Diets Ways to infer diet fxnl morphology, dental microwear, isotopes Dental microwear – pits and scratches (more on Pan. than Anthro.) Isotopic study – ratio of C12 and C13 Masticatory Apparatus Temporalis – postorbital constriction and widely flaring zygomatic arches = larger temporal fossa (thin muscle) – closes the mouth Masseter – originates on z-arch and inserts @ right angle joining of ramus and corpus Abuctor muscles = temporalis, masseter Adductor = pterygoids (lateral and medial) aligns incisors for incision of food Digastric – originates on mandible goes through to voicebox and then up to styloid process (by the ears) allows you to open your jaw Increasing bite force – 1) increase size or cross sectional area and generate greater Fi 2) shortening Lo lengths 3) lengthening Li lengths 4) combination Degree of postorbital constriction means increase in temporalis muscle size Widely flaring z arches changes angle of masseter and makes it able to move vertically or horizontally (grinding) Anterior placement of z arch increases Li and MA, reduced prognathism also increases MA Pterygoids – elevating and side-to-side movements of mandible Mandible – chewing is a unilateral fxn; asymmetric motion generating bending and twisting; thickness resists twisting and bending; one-sided chewing = more force and less stress Fossil diet – adapted to fallback foods Sexual Dimorphism – sexual selection based on male dimorphism Lab 4: Australopith Postcranial Morphology & Locomotion Anatomical feature Ape Human Australopithecine 1] Action of the small gluteals Hip abduction [2] Structure and orientation of ilia [4] Hamstrings mechanical advantage [5] Quadriceps action and energetic efficiency [6] Bicondylar angle Present [7] Lateral lip on femoral condyle [8] Femoral head dimorphism Minimal [9] Calcaneus and talus structure [10] Glenoid fossa orientation Dorsal and superior [11] Torso shape and scapula morphology Funnel-shaped torso, dorsally placed scapula [12] Relative vertebral body size [13] Number of lumbar vertebrae [14] Transverse processes of lumbar vertebrae, MA in lateral flexion and extension [15] Leg length, locomotory energetics and balance Ape hip extension Human Hip abduction Australopith Hip abduction Flat Curved In between Small Greater Similar to humans Costly Flex or extend hip None Present None Present Greatest, shorter femur, wider hip Less pronounced Minimal High dimorphism Smaller Robust Similar to humans Dorsal and superior Inferior later Superior dorsal Barrel Funnel Funnel Little wedging Wedging Wedging 4 5 6 Dorsal, greatest MA Dorsal Dorsal, wider, greater MA Short Longer Short Marked 16-18, no knuckle walking in Australopiths but distal radius primitive Lab 5: Early Homo Introduction - there was a much more complete fossil record before and after the time of “early Homo” - period characterized by isolated fossil finds of teeth and crania, few postcranials - originally thought to be one species – Homo habilis – now thought to be more than one - possibly more related to australopithecines than to homos - 3 major questions about early Homo material o Variability – how many species? o Adaptive grade – ape like or human like o Phylogeny – are these species ancestral or side branches to humans Taxonomy – How many species? - phenotypic data can be poor indicators of specific differences - Brain Size variability – some early Homos have brain volume close to H. ergaster, others closer to male afarensis - Tooth size – pretty consistent across early Homo specimens Grade: Similar to Australopithecines or Homo? - Australopiths – apelike body proportions, smallish brains, large teeth - Homo – modern human body proportions, changes in locomotion, enlarged brains, small postcanine teeth - Limb proportions – early Homos had proportions more like Austs than like ergaster - Index of Dimorphism (ID) closer to Austs than like erectus - Encephalization Quotient (EQ) – half of specimens like Austs, half like ergaster What grade? Probably closer to Austs than Homo, but more than 1 species present – Homo habilis doesn’t adequately capture all the differences Lab 6: H. Ergaster/Erectus H. Erectus was first discovered by Eugene Dubois in Java Now, H. erectus is believed to be in Old World in China, India, Europe, the Middle East, Georgia, and Africa. Earliest specimens ar found in Africa and Georgia at roughly 1.8-1.9 Ma and are sometimes referred to as H. ergaster. The latest H. erectus fossils are from Indonesia and may date to as recent as 50,000 years ago Their increased geographic range into more seasonal and temperate environments is related to changes in body size, shape compared to austral. Mainly, these changes reflect an adaptation to be long distance runners or trekking and an increase in diet quality. Also, technology may have come into play with new tools and controlled use of fire Another adaptation that may have occurred is the diet in that there was now more increased meat-eating/hunting and social or mating stems For this reason, they are small-brained and hidden size dimorphic and are believed to be the first "human-like species" Postcranial: Adaptations/Expanations Retention of arboreal adaptations in Austalopth.-H. Ergaster is clearly proportioned like a modern human with long legs and shorter arms. Found in more temperate and seasonal environments and the long legs suggest ranging behavior had changed, as an adaptation for longer distance movements. Low dimorphism of H. Ergaster/eretus are affected mostly by female body size. Females are much more constrained by energetics than males because of the costs of gestation, lactation, and child-raising. Increased size of females suggests strongly that diet quality changed dramatically to account Thorax shape: Barrel shaped chest-Associated with smaller gut size, reorientation of the ilia, and changes in lung volume and capacity. Rib cage is narrow at the bottom to match that changed shape of the pelvis. Guts and brains are both expensive tissues, so any reduction iin gut size would free up developmental energy for selection to act on. Iliac Pillar: Thicker bone in the form of a "pillar just behind hip joint. This feature may serve as a buttress to the hip joint and is related to physical exertion and habitual posture/locomotion. Cranial/Dentition Adapations/Functions Nasal region: Bones around nose project outwards. Presence of external nose to humidify and warm outside air before entering lungs. Cortical thickness: Robusticity of the overall skeleton Facial Dimorphism: Browridges in male and female specimens or supraorbital torus are quite pronounced and more dimorphic. Male-male competition, placement of face and scaling effects. Tooth Size: Smaller, because of increase in female body mass and smaller gut size shows major shift in diet quality. Increased meat eating or cooking Also, have saggital keel, occipital torus, reduced prognathism. Lab 7: Archaic Homo, Neanderthals, and Modern Humans Regional Continuity (Multiregional) – modern jhumans evolved from Homo erectus populations spread throughout Africa, Europe, and Asia. Rapid Replacement – modern humans differentiated in a more restricted area (Africa) around 200-100 Ka, then eventually dispersed throughout the Old World, Australia, and the Americas-replacing existing populations with little or no hybridization. Evidence (particularly from genetics) supports replacement model. Archaic Homo: Homo Heidelbergensis “Archaic Homo” is the term given to hominids ranging in time depth from 600 Ka in Africa (e.g. Kabwe), to 350-400 Ka in Europe (e.g., Mauer, Petralona), and 100-200 Ka in Asia (e.g., Dali). These specimens are distinct from H. erectus and later H. sapiens in their overall cranial, dental, and postcranial morphology. What sets it apart from erectus: Result of a larger brain ((large as H sapiens relative to body mass) : Parietals are expanded Rounded occipital Frontal is broader Large sep. ridges above the orbits Large lower limb joint- walking long distances Homo Neanderthalensis: Existed in Europe and Southwest and Central Asia from at least 125 Ka until just under 30 Ka. Thick, double arched brow ridges Projecting face: anteriorly in the midline Large nose Occipital bun Large incisor teeth “retromolar” gap behind the third molar Weak chin *Brains were large, if not larger, than the brains of living H. Sapiens Broad rib cage, long clavicle, wide pelvis Stockier than humans Robust hands and feet *many features have little functional meaning Size and wear of incisors suggest that the Neanders using anterior teeth as “tools” Possessed considerable physical strength – perhaps elevate levels of activity similar gestation and birthing process compared to modern humans slightly different forms of locomotion capable of both power and precision grip Homo sapiens Arguably the oldest known specimens are from Bouri, Ethiopia (150-160 Ka.), Skhul and Qafzeh in Isreal (100-90Ka.), and from Klasies River and Border Cave in South Africa (120-80Ka.) Appearance of H. sapiens coincided with all the advances in behavior associated with the “Upper Paleolithic Revolution”. – elaborate speech w/ conceptual and manipulative skills necessary to design and manufacture fine stone and bone tools i.e. needles, fishing hooks, symbolic objects and art. Suggestion that cultural evidence dates back to 200Ka. globular cranium orthognathic and reduced face canine fossa diminutive brow ridges rounded occiput flexed cranial base presence of a chin slightly smaller teeth in relation to body size compared to earlier hominids (including other members of Homo) elongated distal limb bones long limbs relative to the trunk narrow trunk and pelvis low body mass relative to stature relatively large pelvic inlet expansion of the braincase reduction of the face flexion of the basicranium expansion of the braincase may be related to elaborated and fully modern language capabilities. Presence of a chin may be related to the reduction of the face, where the buttress of modern humans suggest a tropical origin for the species