Homo Flashcards
What new evidence has come to light regarding the age of the oldest Homo Sapiens fossils in Africa
found in Omo-Kibish and Herto, Ethiopia, dating of deposites from an eruption of the Shala volcano, which overlies the fossils, shows a new minimum age to be 233+/-22kya
(Vidal et al. 2022)
Describe the juvenile pelvis of H naledi
showed incipient development of features expressed inadultH. nalediilia. The proportional height of the Lesedi ilium was within the rangeof human juveniles between 4–11 years of age
australopith-like iliac blade morphology
expanded auricular surface more similar to humans
Where have Homo erectus fossils been found
Europe (AKA Homo antecessor) between 1.6 and 0.5Mya
West Asia (georgicus) 2-1.6mya
Africa (ergaster) 2-0.7mya
E Asia - 2-0.4mya
SE Asia - 1.5-0.2mya
Who first defined H erectus
What was important about this fossil
Dubois based on fossil from Java
Large cc (950cc) from 0.5mya
How long could erectus have lived in Java
Oldest found in Sangiran ad is 1.6-1.3Ma
Youngest could be as young as 0.1Ma
Survived for 1.5My
What is the problem with fossils found in Sangiran
Sangiran hard to date cos often found and moved by farmers
How does erectus molars differ from habilis molars
in erectus M3 is reduced compared to the very large M3 of habilis
Describe the evolutionary trends in Javan Homo erectus morphology.
Early – many primitive features, large teeth, narrow dental arcade
Later – larger in cc, frontal less flat, AP extended cranial base, short temporal muscle attachment
How old is the Ngandong erectus material?
key features?
found along river so can be hard to date - Riza 2020 suggests ~110kya
CC:1132cc
Flat horiztontal orbital torus
Dubois found 2 femora and a calotte in Java which were all claimed to be erectus. Why is this doubted?
Trinil I femur is much closer to a late Pleistocene H. sapiens morphology (therefore not used in erectus analyses)
Trinil II is likely contemporaneous with the calotte and both are erectus
What is Meganthropus
Dubois’ paratype of Pithecanthropus erectus from 1891 NOT HOMININ BUT APE(S), probably related to Lufengpithecus, and possibly including Gigantopithecus
Give some typical erectus features of the fossils in Sangiran
Large SO torus with supratoral sulcus
Long receding forehead
bregmatic eminence
Large face, with reduced alveolar prognathism
Salient nasal bones
Where were key erectus fossils found in 1920s and 30s
How old?
China
The fossils from Zhoukoudian, China were lost during WWII
Estimated age: 780 – 400 Ka
How does the Zhoukoudian erectus material differ from that of Java? (3)
- Size: 915-1225cc (upper is almost sapiens)
- Forehead: not receding – arched and curved – allometry?
- Sagittal keel: much less pronounced
Why is the material from Hexian interesting?
V young – 400ky
Affinities to early java material – more similar to older erectus – survival of more primitive erectus in some parts of china
Some dental (EDJ) affinities to early African Homo
What is the temporal pattern of Homo erectus occupation of Asia? What questions does it raise?
Early Pleistocene has fossils and stone tools but then large temporal gap – repopulation?
Fossils in Africa from ~2mya classed as erectus but that species is based off Dubois’ skull cap from ~0.5mya in SE Asia – surely there had been selective pressures etc – problems here
What is the type specimen of ergaster
KNM-ER 992 from Koobi Fora (1.7ma)
mandible with no retromolar space and a slender corpus
What is the retromolar space
The retromolar space or retromolar gap is a space at the rear of the mandible, between the back of the last molar and the anterior edge of the ascending ramus where it crosses the alveolar margin.
This gap is generally small or absent in modern humans, but it was more often present in Neanderthals
Not seen in ergaster but is in erectus and earlier hominins
What is the importance of the Turkana boy
first humanlike narrow skeleton (90%) complete
How old was the Turkana boy
8‐9 years old [developmental stage equivalent to a human 12 year old child]
What were the key features of the Turkana boy (4)
Resembles a very robust modern human from
the neck down
Brachial and humero‐femoral index in the
modern human range
Barrel‐shaped rib‐cage
Narrow bi-iliac breadth
What is interesting about KNM-ER 42700?
East African H erectus
Young adult – not fully fused synchondrosis
Mosaic – projecting glabella, distinct keels, no occipital torus, rounded back, max length at lambda, modern arrangement: tympanic bones orientated coronally(?) while petrous bones are sagittally
Thin bones in skull while erectus has thick bones – no prominent SO torus or trigones – outside range of erectus? But not habilis either
What was the range of cc in E African erectus
from 691 cc (KNM‐ER 42700) to 1067 cc (OH 9)
How does E African occipital compare to that of non-African erectus
Occipital: less angular than in non‐African Homo erectus (except OH9)
What is important about the Buia and Bouri skulls?
Very different skulls but contemporaneous (1mya)
Bouri has the most features shared with later Middle Pleistocene hominins
Regardless of whether we classify them as populations of the same species or separate species, what can we learn from the mosaics of traits seen in different groups of “erectus”?
Tall, slender body, with narrow shoulders, barrel‐shaped rib‐cage, narrow pelvis, legs longer than arms
Larger cranial capacity than Homo habilis
Why is StW 53 interesting
has been classified as Homo, Au. africanus, and P. robustus
existed 1.78‐1.49 Ma
• Small cc
• Large teeth
• M3 > M2
What is the temporal distribution of African H erectus
Homo ergaster
2-0.8mya
What evidence is there of regional erectus fossils being different species
morphometric studies show:
The frontal bone of African “erectus” fossils is different
The parietal bones of African “erectus” fossils are different
The occipital bone of African “erectus” is similar to Chinese “erectus”, but not Javanese erectus fossils
Different skull bones are different in ‘erectus’ from different regions – different trajectories?
Compare erectus and ergaster (8)
ergaster has thinner cranial bones No sagittal keel No bregmatic eminence More pronounced supratoral sulcus Less developed supraorbital torus Less developed occipital torus Slender mandibles Relatively gracile TMJ
What does Homo erectus sensu lato include
- All post‐habiline hominins prior to major brain enlargement in the Middle Pleistocene
- Widely distributed in Asia and Africa
- 2.0 – 0.1 Ma
What is Homo erectus sensu stricto
Javanese fossils only, perhaps
including East Asian remains
• >1.6 Ma – 0.1 Ma
What is H ergaster
African lineage only • Ancestral to H. erectus ? • 2.0 – 1.0 Ma • Less specialized features • Very variable
What does a) the fossil evidence and b) lithic evidence suggest about when hominins first left Africa
a) Crania from Dmanisi, Georgia ~1.77Ma
Longgudong - >2.14ma?
b) Stone tools from Southern China 2.12Ma – only 17 artefacts
N India – 2.6Ma
Indication but not substantial but does suggest OoA 2-3ky earlier than fossils suggest
Who was the first hominin in Georgia
How long were they there
Did they have tools
What was the cranial capacity of these hominins
Homo georgicus (Georgian Homo erectus)
1.77 Ma (fossils)
• ~ 100,000 years of occupation of the site
(from ~1.85 Ma)
Oldowan artefacts (Mode 1)
Small cranial capacity: 546 cc to 730 cc - lots of variety
• D4500: 546 cc (EQ ~ Australopithecus)
What can we infer about the behaviour of Homo georgicus (Georgian Homo erectus)
toothless individual 9D3444/D3900) would not have had easy access to food so must have been looked after
o Paramasticatory use of anterior dentition of D2600 – teeth worn labially – used to hold objects
o Lateral extension of body of mandible near molars – typical of early hominins absent in ergaster and later Homo
What was the face shape of H georgicus
Zygomatics?
Prognathic – could be part of developmental process as D4500 has most pronounced cf D2282 and D2700 which are both not fully developed (adolescent and juvenile respectively) - same can be said for the supraorbital torus
Flared zygomatics cf not flaring in erectus
Less angular head – more similar to ergaster
What did Lordkipanidze et al. (2013) suggest about taxic diversity of early Homo, based on Dmanisi?
Early homo out of Africa were very diverse and should be considered as a single highly variable species
This implies the existence of a single evolving lineage of early Homo, with phylogeographic continuity across continents.
- What might the link between A.L. 666, SK847, and Dmanisi?
Similar shape of subnasal clavus and maxillary pillars and angle of zygomatics which are slightly flared – possible association
This would put Dmanisi material as descent from fossils in Africa that predate erectus or ergaster
Which species has the most similar body proportions to the Dmanisi postcrania
H sapiens
What are the derived traits of Homo georgicus (Georgian Homo erectus)
• Biomechanically efficient locomotion, both for long‐range walking and energy
storage/return during running
Leg length and morphology similar to modern humans
Adducted hallux and plantar arch
Homo georgicus (Georgian Homo erectus) is a mosaic. What are the primitive features
Retention of primitive characters not seen in later hominins More medial orientation of the foot Absence of humeral torsion Small body size Low encephalisation quotient (EQ)
Why may the Dmanisi hominins not be a single species
Significant differences between individuals • Significantly larger size variation than in humans • Significantly larger shape variation that any extant ape species
Established early in ontogeny
Independent of size or sexual dimorphism
Not synchronic deposition of fossils
2 different palaeodemes?
a) What is the oldest hominin fossil out of Africa
b) Oldest sites with stone tools or cut‐marks on fauna
a) Longgudong, China (>2.14 Ma? controversial)
Otherwise, Dmanisi, Georgia, W Asia 1.77 Ma
b) Quranwala, Siwaliks, N India 2.6 Ma (controversial)
When was there faunal exchange 1.5-2.5mya
2.6 – 2.5 Ma: Afro‐Eurasian faunal exchange
Africa->Eurasia: Giraffa, Struthio
Eurasia->N. Africa: Ursus
Eurasia->Sub-Saharan Africa: equus
1.8 – 1.6 Ma: important Afro‐Eurasian faunal exchange (mostly out of Africa) - carnivores, primates, hippos
From Eurasia to Africa: wolves
What is the significance of the faunal exchange 1.5-2.5mya
• 1.8-1.6 Ma: out of Africa mostly – main point of dispersal
Why does it matter? – hippos are v water dependent like humans so not only was there a corridor but also fresh water was available
What might the relationship between hominins and carnivores have been? Why does this matter for dispersals?
Used same range bc humans acted as scavengers utilizing the carcasses left behind by giant carnivores
Why might the earlier dispersal (2.5mya) only have been a limited dispersal?
Were exchanges of fauna but limited – no archaeology
Did not colonise Maghreb(?) (Western N Africa) and Nile was not strong corridor of water to support the dispersal
Why do researchers suggest that ATD6-69 might be the LCA of Neanderthals and modern humans?
Sima del Elephante - maxilla of child 1.2Ma – LCA of neanderthals – v controversial
Intersection of Zygoma and maxillary has sharp depression – typical of early humans
Why in spain? – recurrent connection with Africa through middle east? – maybe just part of erectus diversity
What do proteomes tell us about Homo antecessor?
Antecessor’s ancient proteomic data (small sample size) suggests it is the sister group of sapiens
Describe Model 1 of the dispersals of early Homo out of Africa
MODEL 1 – one single dispersal early (2.5‐2.2 Ma)
Early dispersal of early Homo; tethered to highlands? (stone; water)
Giving rise to Homo georgicus and a population ancestral to Homo erectus in eastern Asia
Describe Model 2a of the dispersals of early Homo out of Africa
Main dispersal at 19‐1.8 Ma
[either no early dispersal, or extinction of those very early Asian populations]
Dispersal of early Homo, giving rise to Homo georgicus (and later European pops., including H antecessor) and ancestor of
Homo erectus in East Asia
1.6 ‐ 0.8 Ma EMPT
Regional isolation in Asia
Expansion in Europe
When did H erectus live?
What happened during this time to this species?
2-0.1mya
it regionalised, acquiring local differences due to different pressure in each local environment, as well as due to founder effects and drift
Many researchers argue that erectus differences should be recognized taxonomically, and several local populations have been given species names:
ergaster to the African fossils
georgicus to the Dmanisi fossils
antecessor to the fossils from Spain
erectus to the Javan and Chinese populations
What are the key features of erectus’ skull
what ergaster does not have
thick cranial bones (not ergaster), angled occipital (variable), with strong
superstructures, including a sagittal keel (not ergaster) and a pronounced
supraorbital torus
With which early members of Homo did the erectus group overlap temporally? And what about members of other hominin genera?
Rudolfensis and habilis
Boisei
How does the cranial capacity of erectus vary
546-1250cc
What are the body proportions of erectus
1.3-1.7m
Modern proportions
KNM-WT – share proportions of most recent Homo and v different from habilis and Lucy
Erectus achieved greater stature earlier on than modern humans
What can we learn from the hominin footprints from Ileret, East Turkana?
1.5mya
Modern human functional anatomy – adapted hallux, medial longitudinal arch and medial weight transfer before push-off – size and distance between are consistent with stature of erectus
Fully committed to terrestiality
Describe the arboreality of H ergaster
Homo ergaster fully committed to life on the ground
Adapted to long‐distance running
Useful in long‐distance scavenging and hunting in open country
Is there sex dimorphism of erectus
If all not habilis = erectus then there is dimorphism – size
Describe the life history pacing of Homo erectus. How do we know?
Looking at microstructure of enamel can give age – KNM-WT15000 was 8yo (consistent with dental eruption and fusion of epiphyses) – same as chimp trajectory – stark contrast to human child (closer to a 10-15yo human child’s anatomy) – erectus still growing more like an ape than human but there is a shift to slowing of LHS
How does the human brain grow after birth
Humans have sustained brain growth after birth – unique to humans (6 years after birth at foetal growth rate – fast and demanding)
How long ago do we have evidence of humans eating meat
Efficiency through hunting, not just scavenging [cut‐marks on elephant bones at
Olorgesailie (~1 Ma) with no underlying carnivore marks]
When is there evidence of controlled use of fire
Swartkrans, South Africa: burnt bones
in 20 levels (1.5‐1.0 Ma)
• Koobi Fora, East Turkana, Kenya (1.5 Ma)
How long ago have we got evidence of fireside entertainment
15kya
plaquettes from France with ancient paintings as well as colouring suggesting they were placed by a fire
when AI was used to see what they would have looked like by the fire originally they moved
Needham (2022)
What were key factors contributing to erectus’ LHS (2)
What did this mean for its LHS
lived in a complex environment – broader diet, larger ranges, competition with carnivores
was encephalised – larger brain to body size
developed more slowly than australopithecines but still faster than modern humans
had a reduced childhood in comparison to humans ‐ short period of learning
Bigger at an earlier chronological age than humans
Homo erectus developed more slowly than australopithecines but still faster than modern humans
had a reduced childhood in comparison to humans ‐ short period of learning
Bigger at an earlier chronological age than humans
What allowed this to occur
high energy diet using cooking and hunting
What was the importance of erectus being bigger at an earlier chronological age than humans
protect against predation
Erectus juveniles were energetically independent and may have contributed to the group - high energy environment
How much more energy could cooked starch contribute to hominin diet?
What else did fire allow hominins to access?
How did the introduction of cooking affect time
> 30%
honey -> >100c more
> 4hr/day saved on chewing
What is the temporal trend in encephalization in Homo erectus?
Australopithecines constant trend of encephalization, with increase in early homo, and progressive trend towards larger size from erectus
Complete pelvis of female suggests they could give birth to big headed babies – but not as big as humans
Different proportions of brain areas
Comment on the cognition of Homo erectus.
Different cognition from other hominins – increased carnivory and archaeological sites
Care of old and young
Cannibalism?
Trinil, java – geometric scratches on shell from 0.54mya - notation/ symbolism
Problem solving in complex environment – wider ranges, new predators, new competitors, climate change
What is the evidence for erectus’ care for elderly
toothless skull of individual >40y/o
Dmanisi 1.77mya
What is the evidence for hominin cannibalism
cut marks on skulls
1.2mya in Spain
H antecessor
What were the conditions and constraints on H erectus becoming a fully terrestrial carnivore ape, with opportunistic range expansions and dispersals
CONDITIONS: carnivory and stone‐ tool making; slender, narrow body with longer legs than arms
CONSTRAINTS: Permanent freshwater and raw materials for stone‐ tool manufacture; relatively fast growth
What were the causes and consequences on H erectus becoming a fully terrestrial carnivore ape, with opportunistic range expansions and dispersals
CAUSES: Increased competition due to niche reduction as a result of climate change and inter‐specific hominin diversity; competition with carnivores
CONSEQUENCES:
Adaptive radiation of Homo
with new ancestral body plan
and large body size achieved
early in development, leading to geographical diversification, increased genetic variance, and new subsistence strategy (social predators) and a dependence on technology and high energy diet. These
consequences set the adaptive platform through which all descendant populations benefitted from slower growth, encephalisation, and greater social cognition, leading to convergent evolutionary trends
Is cooking still important
Even now across the world it is v hard to find anyone who has not cooked in the last 3-4 waking hours
Evening meal is universally cooked
hard to thrive on raw food
Cooking helps more than eating meat
What happens if modern humans don’t cook food
As raw food increases to 100% - amenorrhea even with domesticated food and there is no shortage of food – won’t be able to have baby – we are adapted to eat cooked food
Starch – when cooked v similar to just eating glucose – more productive – does this fit with digestive anatomy – different processes in large and small intestine
How much honey to great apes eat
None
hunter gatherers eat lots – humans use cooking – honey guides Apis honey, reducing time looking for honey from 8 to 3 hours
Evolved behavior not learnt from parents
- ancient adaptation to high glycaemic load
5 things we have learnt from ancient genomes about the evolution of hominins in the last 1 million years?
- LCA of modern humans – 280kya
- 300kya is when sapiens separated
- Neanderthals and denisovans separated 450kya
- Sapiens separated from N and D 750-550kya
- 950-750kya when N/D/Sapiens separated from other hominins
What happened 800kya
Why did this happen
MAJOR shift in climate (early to middle Pleistocene)
Long‐term cooling of sea‐surface temperatures
• Increase in amplitude of glacial cycles
• Shift from 41,000 to 100,000 year mode
Due to changes in Earth’s orbit -
What is eccentricity and how does this affect climate
Eccentricity refers to the shape of the Earth’s orbit around the sun, and its
natural variation from more or less elliptical.
When the orbit is more elliptical, Earth receives greater variation in energy from the sun throughout the year than when the orbit is circular
What is obliquity
Obliquity refers to the axial tilt of the Earth, or angle between the axis and plane of rotation.
Earth’s obliquity oscillates between 22.1o and 24.5o on a 41,000 year cycle.
Obliquity is what causes the seasons.
What is the precession of equinoxes
Why is it important
The precession of the equinoxes (the shift between a hemisphere being tilted away from or toward the sun) occurs on a 19,000‐23,000 year cycle.
Precession cycles define the timing and intensity of solar insolation.
What are the 2 configurations of Milankovitch cycles
Glacial/Ice‐Growth configuration
Interglacial/Ice‐Melt configuration
Describe the Interglacial/Ice‐Melt configuration of the Milankovitch cycles
High eccentricity (elliptical orbit)
High obliquity (more seasonal contrast)
Small Earth‐Sun distance in summer (more
ice melting, increased heat absorption)
EFFECT: warmer summers, cooler winters
Describe the glacial configuration of the Milankovitch cycles
Low eccentricity (circular orbit)
Low obliquity (tilt)
Large Earth‐Sun distance in summer (no ice
melting, increased heat reflection)
EFFECT: less seasonal contrast
What happened to Climate change in the Early‐to‐Middle Pleistocene Transition (EMPT)
• Long‐term cooling of sea‐surface temperatures
• Increase in amplitude of glacial cycles
• Shift from 41,000 (dominated by obliquity) to 100,000 year (dominated by eccentricity) mode
• Shift to asymmetric glacial cycles: short warm and long cold phases with abrupt endings
(terminations)
What are glacials/interglacials defined by?
Glacials/Interglacials are defined by insolation at 65 degrees N latitude
What are “terminations”?
When a glacial cycle ends in an abrupt increase in temperatures
What happened to the length of temperature cycles after the EMPT
Shift from 41,000 to 100,000 yr long cycles
How do processional cycles affect the Sahara
• Hydrological response to eccentricity and its interaction with precessional cycles (ie, how the
amount of precipitation in the tropics [monsoons] relates to the interaction between the shape of
the Earth’s orbit around the sun and the timing and intensity of solar insolation at high latitudes)
- EFFECT: amplification or dampening of tropical monsoons
- eccentricity minima: weak monsoons at precessional timescales, increased aridity in East Africa, “Yellow Sahara” phase
- eccentricity maxima: increase in tropical climate variability at 100,000 and 400,000 yr cycles [independent of continental ice volume] with alternating strongest and weakest monsoons [alternating “Green Sahara” and “Desert Sahara” phases]
When was the most recent Green Sahara event
12 to 6kya – when monsoons are at max for short window of time – rained so much animals could cross and live there
Do inferred population splits tend to happen in dry or wet Sahara episodes?
s-D-N split from other hominins was during dry spell – moment of fragmentation of African populations – sufficiently long to keep populations separate to maintain different evolutionary trajectories
s from N and D – intense wet episode with green Sahara – during contact between sub-Saharan and northern Africa
N from D during dry – fragmentation event again
S and other hominin again in aridity then followed by green Sahara event
Why is the Middle Pleistocene a “muddle”?
- Processes: different dispersals – lots of different interactions – complex
- Analytical techniques: not enough organic stuff to date properly
Describe African fossil record from EMPT
KNM-OL 45500 – one of smallest hominin fossils ever found - + the other two v different crania (Buia and Bouri)
When/ where are Buia and Bouri from
erectus/ergaster crania
Buia
• Eritrea
• ~ 1.0 Myr
• 750‐800 cc
Bouri, Middle Awash, Ethiopia
~ 1.0 Myr
995 cc
What are 2 of the oldest fossils of hiedelbergensis
fragmentary cranium from Algeria ~800‐700 Ka ?
Casablanca
• Morocco
• 600‐400 ?
• Cranial fragment
Give the primitive and derived traits of the hiedelbergensis fossils found in North Africa from 400kya
Primitive traits: • ~930 cc • Low position of max breadth • Pronounced supramastoid crest • Bregmatic eminence
Derived traits: • Frontal convexity • Rounded temporal squama • Vertical wall of parietals • Pattern of meningeal vessels
What is the oldest evidence of heidelbergensis in South Africa?
What is the cc
> 600 Ka
1225 cc
What the flora and fauna like in South Africa ~600kya
Over 50 different spp of mammals, as well as birds, reptiles and amphibians.
Vegetation: included bush and trees, at least around the spring‐fed water
bodies, and more open bush and grass than at present, but appears to have been fynbos.
Describe the features of heidelbergensis in East Africa ~600kya
Bodo
Middle Awash, Ethiopia
630 Kya
- ~1100 cc
- broad, massive face
- extremely broad nose (the largest of any Pleistocene hominin)
- thick supraorbital torus
CUT MARKS - prehistoric defleshing/ cannibalism?
When did Lahr place the LCA for humans and Neanderthals
virtually reconstructed 3 different scenarios and found one placing vLCA 700kya is most consistent with fossil record
suggest an Afro-European ancestral population in the Middle Pleistocene (Homo heidelbergensis sensu lato)
What is the importance of Gesher Benot Ya’aqov (GBY)
Israel
800‐700 Ka
• ~ 100 Ka use of area, lacustrine
• Rich Acheulean industry
rich evidence of fire – points to north Africa dispersal
Did heidelbergensis make it to the UK
Boxgrove - found a large tibia suggested to be heidelbergensis from 500kya
Give a key heidelbergensis fossil from Greece
Petralona ? [700‐200 Ka] • 1220 cc ~90o angle nuchal/occipital planes • very large, orthognatic face • prominent transverse torus
Describe the cranium from Ceprano Italy from ~400kya
Homo heidelbergensis ?
430‐385 Ka
• 1185 cc - large
- Very pronounced, tall supraorbital torus
- Angled occipital
- No sagittal keeling
- Very thick bones
- No derived Neanderthal character
Nothing here preempts Neanderthals
Could have had different populations with different amounts of derived features around this time
Give the cave in Spain that holds lots of individuals
Who are these individuals
Sima de los Huesos
> 29 individuals at bottom of cave shaft
More women than men – bar 1 child most are teens/young adults
Died simultaneously
May individuals from single population
together with bones of cave bears and 1 handaxe
What is the cc of the crania in the Sima de los Huesos
Any other important bones?
from 1125cc to 1390cc (larger is average for model humans)
Complete pelvis – much more flared than modern – very tall -165-170cm
Radiating fractures in crania – interpersonal conflict
handaxe
child with serious pathology - care?
Have heidelbergensis fossils been found in France?
yes in Arago from 400kya
smaller cc: 1166cc
What is the importance of Steinheim
Partial cranium – not very derived and has small cc (~900cc) – survivor ship of an older population which was present before N arrived
What is heidelbergensis broadly
Heidelbergensis has become associated with any middle Pleistocene specimen in Europe that fits the description of being very robust with teeth similar to the type specimen from Mauer
Were Homo heidelbergensis hunters ?
• Large numbers of stone tools associated with large animal carcasses:
elephants, rhinos, buffalo, etc.
•‘Butchery’ sites, such as Torralba & Ambrona, Spain
•Hunting weapons – Schöningen spears, Germany
•Hunting & Scavenging ?v
Is there any suggestion heidelbergensis could control fire
Schöningen, Germany ~ 400 Kyr 2‐3 m long wooden spears Tip hardened by fire Associated with butchered horse carcass
Temp would be so cold they would have needed control of fire and needed high energy diet
When was the Neanderthal/ Denisovan split
450kya
What are the possible Homo erectus remains found to be younger than Neanderthal/ Denisovan split in E Asia
Jinniushan - female (from pelvis) with 1400cc
What is important about fossils found in Xujiayao
from 160‐220 Kya
Mosaic:
• Different from archaic and recent humans
• Some features shared with Neanderthals, but not exclusive
• Some primitive traits shared with older eastern Asian fossils
Give some evidence for populational interactions across Eurasia during the late Middle and early Late Pleistocene.
fossils from Linjing; 105-125kya
overall cranial shape, and especially the combination of the wide cranial base and low neurocranial vault, indicates a pattern of continuity with the earlier, Middle Pleistocene eastern Eurasian humans. Yet the presence of two distinctive Neandertal features—one (iniac and nuchal morphology) unknown among earlier eastern crania, and the other (labyrinthine proportions) evident in only one similarly aged eastern Eurasian fossil
What is the importance of fossils in Xiahe, Tibet
> 160kya mandible
proteomics indicate it is Denisovan
indicate that archaic hominins occupied the Tibetan Plateau in the Middle Pleistocene epoch and successfully adapted to high-altitude hypoxic environments long before the regional arrival of modern Homo sapiens.
Why does a fossil found in Taiwan highlight Pleistocene Asian diversity
unexpectedly late survival (younger than 450 but most likely 190–10kya) of robust, apparently primitive dentognathic morphology in the periphery of the continent
cannot be simply explained by clinal geographic variation of Homo erectus between northern China and Java, and suggests survival of multiple evolutionary lineages among archaic hominins before the arrival of modern humans in the region.
When was LCA of modern humans in Africa
280kya
Which fossils may be attributed to Homo rhodesiensis
Ndutu
400kya, 1100cc
Eliye Springs, Kenya
350kya, 1300-1450cc, Chronic anemia in childhood?
both have no sagittal keeling and relatively rounded occipital
Kabwe, Zambia - Type specimen of Homo rhodesiensis
299kya
Bregmatic eminence like erectus so primitive but present 300kya
What could Homo rhodesiensis be part of
Homo heidelbergensis
What is Homo helmei
found in Florisbad, South Africa
• 259 ± 35 Ka
• Large orthognatic face • Large orbits and nose • Large supraorbital ridges, but not shaped like a torus • Very broad
(Homo aff. Sapiens)
What is important about fossils found in Jebel Irhoud
315kya
Very small face with many modern features – has beginnings of a chin
Who is a possible contender for LCA of mid-Pleistocene hominins
Bodo, dating to 630 ka, shares features with later old world mid Pleistocene hominins pointing to that connectivity in time
In pre-aDNA days, where did researchers place Homo heidelbergensis on a hominin phylogeny?
What did Krause et al. (2010), using aDNA, add to the phylogeny—and where on the phylogeny?
Believed to emerge out of Africa from erectus bottleneck
Bottleneck in heidelbergensis led to sapiens and a distinct bottleneck led to neanderthals who then died off
• Heidelbergensis as LCA between Neanderthals and sapiens
OR
• Rhodesiensis and heidelbergensis as sister taxa stemming from antecessor with the former leading to sapiens and the latter leading to Neanderthals
OR
• Rhodesiensis being LCA of neanderthals and sapiens with no hiedelbergensis
Separate branch of ghost hominin – Denisovan
What other hominin genes does the Denisovan genome contain
0.0% modern human
• 0.5% Neanderthal (*** HLA)
• 0.5‐8% ANCIENT (Homo erectus?)
Denisovan genes in what other hominins:
0.0% Denisovan in Neanderthals
• 0.2% Denisovan in modern Han Chinese, Dai of S China, Karitiana of Brasil
• ~4‐6 % Denisovan genes in PNG, Australia, Philippine Negritos
What is strange about the genetic material from Sima de los Huesos (Homo heidelbergensis)?
mtDNA Clustered with Denisovans
Nuclear genome: Neanderthal ancestor
What is the likely temporal range of the LCA of humans, N and D
900-700kya
extreme aridity
What was the Competitive context (other hominins?) for the LCA of humans, Neanderthals, and Denisovans
north vs sub Saharan? Homo antecessor? At least co-existed with another hominin
No fitting fossil has been found
When is genetic split of
a) humans x NEA/DEN
b) genetic split NEA x DEN
Where did each occur
a) 750-550kya (within Africa)
b) 450kya (in Eurasia)
When did the African ancestors of NEA‐DEN exist?
What climatic event does this fit with?
How does this meet archaeological findings (2)
What does this suggest
Probably between 650‐550 Ka
Minor “Green Sahara” events between 600‐500 Ka
Consistent with introduction of Acheulean into Europe
Consistent with the similarities observed between African and European Middle
Pleistocene fossils
Implies the rich, African looking (full of cleavers, absent in Europe) Acheulean at GBY
was an earlier (limited?) dispersal
- Is a gene coalescence date the same as a population separation?
not necessarily
Model 1 suggests Neanderthal and Denisovan lineages separate during the during out of Africa event. What does this imply?
What is the problem with this
the split would date to date of dispersal out of Africa , and implies that the LCA was not Eurasian, but African, and lived between 700-450 Ka.
Too late to fit fossil record
Achulean would have to be earlier dispersal
What is model 1 of the LCA of Neanderthals and Denisovans
: Neanderthal and Denisovan lineages separate during the process of out‐of‐Africa
dispersal
time of split = time of dispersal out of Africa
Implies that ancestor of NEA‐DEN lived in Africa between 700 and 450 Ka
Too late to fit the fossil record? Earliest ages (Ceprano, Sima) ~430 Ka; Petralona?
Implies the rich, African looking (full of cleavers, absent in Europe) Acheulean at GBY
was an earlier (limited?) dispersal
Give model 1 of the LCA of Neanderthals and Denisovans in the context of the related climatic events
900‐700 Ka
AFRICAN MEGADROUGHT
Populations fragment and
differentiate
700‐450 Ka
Populations expand
“Green Sahara” events
450 Ka
Dispersal out of Africa
and split between NEA
and DEN
What is model 2 of the LCA of Neanderthals and Denisovans
: Neanderthal and Denisovan lineages separate in Eurasia some time after the out
of Africa dispersal
time of dispersal out of Africa much before split NEA x DEN
Implies that ancestor of NEA‐DEN lived somewhere in Eurasia between 700 and 450 Ka
What Eurasian fossils match the LCA of NEA & DEN?
Earliest “non‐erectus” fossils in China ~350 Ka – where were the Denisovans before?
What is the problem with the idea of an Eurasian LCA of Neanderthals and Denisovans?
No Eurasia fossis match this
Earliest non-erectus fossils are ~350kya – where were Denisovans before this?
Why could one argue that the non-erectus (e.g. Jinniushan, Dali) fossils in China are Denisovans?
temporal overlap between Homo erectus and a different hominin in China who shares
similarities with contemporary western Eurasian populations and Neanderthals
0.5‐8% ANCIENT genes in Denisovans - Perhaps this ancient gene is Asian erectus and occurred as these two populations met when the west entered china
Why was the evolutionary geography complex in the middle Pleistocene
Afro‐Eurasian dispersal(s)
Euro‐East Asian dispersal(s) – Siberian corridor
Regionalisation of evolutionary trajectories
Who were the hominins in Africa and Eurasia in the Middle Pleistocene
Homo heidelbergensis, Homo rhodesiensis (?) & Denisovans
Acheulean in Europe and Africa
Complex behaviours
Who were the hominins in China in the Middle Pleistocene
late survivorship of H. erectus (~400 Kyr)
China: co‐existence of H. erectus with a heidelbergensis‐related
hominin, presumably related to Denisovans
Who were the hominins in SE Asia in the Middle Pleistocene
: late survivorship of H. erectus (~100 Kyr)
Denisovans in Southeast Asia?
Co‐existence with Homo floresiensis – the hobbit
