Neanderthals Flashcards
Earliest European remains
Barranco Leon, 1.4mya - Hominin deciduous molar
Sima del Elefante, Atapuerca, 1.2mya - mandible and phalanx, Mode 1 stone tools, classified as Homo sp.
Gran Dolina, Atapuerca, 800kya - Neanderthal-modern human divergence
Sima de los Huesos, Atapuerca, 430kya - early Neanderthals
Routes to Europe
Most accepted view: through the Near East, Anatolia, Southeast Europe, up to Western Europe
Earliest hominins in the Iberian peninsula - alternative route through the Gibraltar strait? Not likely, but possible
ATD6-96 - extremely gravcile mandible from Atapuerca vs Tighennif 3 - extremely robust mandible from North Africa
Homo antecessor
Antecessor means explorer
Described as last common ancestral species to Neanderthals and modern humans
Gran Dolina: >170 remains, 11 individuals, mostly subadults
~1000cc
More primitive than Neanderthals and modern humans, but more derived than H. erectus
Canine fossa
Primitive dentition but some Neanderthal traits
(Dental traits shared by Neanderthals and H. antecessor are considered Neanderthal autapomorphic traits)
Homo antecessor in Africa?
UA 31 (Buia cranium) and BOU-VP 2/66 (Daka calvaria)
From Ethiopia, dated to ~1.0mya (similar age to H. antecessor)
Related to the last common ancestor of Neanderthals and H. sapiens?
H. antecessor behaviour
Bones have cutmarks.
Defleshing and breaking of bones for marrow.
Treatment of human remains indistinguishable from fauna.
Nutritional cannabilism?
Mainly infants and subadults.
Result of intergroup aggression?
H. heidelbergensis - temporal range, holotype, sites
~640-250kya
Holotype - Mauer mandible
Sites - Europe:
Arago, France - 450kya - similar to African material
Mauer, Germany - 610kya - holotype
Ceprano, Italy - 400kya - affinities with H. antecessor, surprisingly late for primitive features
Petralona, Greece - 400kya - similar to African specimens
Sites - Africa:
Kabwe, Zambia - 300kya - similarities to H. sapiens but also archaic features
Saldanha, SA - 400-800kya - similar to Kabwe and Mauer
Ndutu, Tanzania - 350kya - smaller face, gracile features
Bodo, Ethiopia - 640kya - similar to Petralona and Kabwe, more primitive
Sites - Asia:
Jinniushan, China - >200kya - large but gracile
Dali, China - 260kya - similar to H. sapiens and Neanderthals
Narmada, India - >236kya
Disputed taxonomy, phylogenetic relationships, and validity
H. heidelbergensis anatomy
Similarities to H. erectus:
Broad faces
Big browridges (but more swollen)
Relatively long, low cranium
Derived cranial features
Similar to Neanderthals:
Divided browridges
Nasal shape
Similar to H. sapiens: Reduced midfacial prognathism (variable) Canine fossa (variable)
Similar to both Neanderthals and H. sapiens:
Brain size (1100-1400cc - first hominin within modern range)
Increased convexity of frontal and parietal
Reduced post-orbital constriction (wider frontal)
Neanderthal-modern H. sapiens LCA
Early H. sapiens - Djebel Irhoud redated to 315kya
Early H. neanderthalensis - Sima de los Huesos dated to 430kya
LCA must be substantially older; H. heidelbergensis is an unlikely ancestral species
Denisovans
First species identified through ancient DNA - very little fossil evidence. Know a lot about genetics but almost nothing about anatomy.
Sister branch to Neanderthals - early divergence within Neanderthal lineage
4-6% Denisovan DNA in present-day Melanesians
Neanderthal-Denisovan interbreeding e.g. 90kya female
Some Asian fossils usually classified as H. erectus or H. heidelbergensis may be Denisovans
Need to find Denisovan aDNA in other fossils OR find more fossil evidence from Denisova
Early Neanderthals
Sima de los Huesos, Atapuerca, 400kya
> 6000 fossils, 28 individuals. Whole bodies deposited, no evidence of habitation (not easy to reach).
Most 10-18 years, absence of very young and very old
Only carnivores and hominins - predation?
Not young or old - catastrophic event?
Single handaxe - ‘Excalibur’ - ritual behaviour? Would be the earliest case of ritual behaviour on in the fossil record
Single population? Intragroup variation, sexual dimorphism, allometry
Relatively tall, robust, similar body proportions to Neanderthals
Most traits show clear Neanderthal affinities, but to different degrees
Teeth are extremely derived - even more Neanderthal than the teeth of classic Neanderthals
Brain 1240cc
Primitive brain organisation
Mosaic evolution
Traits related to mastication are derived in Neanderthal direction, non masticatory traits are much more primitive
Accretion model for the origin of Neanderthals
Dean, 1998
Neanderthal features appear gradually throughout European Middle and Late Pleistocene
4 stages:
- Early pre-Neanderthals e.g. Arago, Mauer, Petralona
- Pre-Neanderthals e.g. Sima de los Huesos, Swanscombe, Steinheim
- Early Neanderthals e.g. Ehringsdorf, Krapina, Saccopastore
- Classic Neanderthals e.g. Neanderthal, Spy, Gibraltar
SH 14
Sima de los Huesos. ‘Benjamina’
Craniosynostosis - early closure of cranial sutures
Perhaps behavioural anomalies
Conspecific care likely involved
Earlier example from Dmanisi
Conspecific care - early acquisition in hominin behavioural repertoire
SH 17
Sima de los Huesos.
Cranium with two fractures, probably produced with the same object
Can be the result of intentional and repeated blows
Lethal act of interpersonal violence?
Perhaps related to the origin of accumulation?
Co-operative hunting - Sima de los Huesos
Huge accumulation of bison remains in a layer associated with SH hominins
More than 25,000 remains with cutmarks
Preferential use of axial/meaty elements
Evidence of mass communal hunting - hunting strategies plus preservation of meat?
Symbolic behaviour and lineage - Sima de los Huesos
SH hominins had auditory capacities similar to H. sapiens (compatible with language) BUT this does not mean that they were capable of producing language
Classic Neanderthals - temporal range, holotype
150-30kya - older limit difficult to establish because origins can be traced to the Middle or even late Lower Pleistocene
Holotype - Feldhofer 1 (Neanderthal 1)
Neanderthal vs modern human anatomy
Neurocranial shape - lower and flatter in N, globular in S
Forehead - low and receding in N, high and bulging in S
Supraorbital anatomy - double-arched in N, minimal and divided in S
Anterior mandible - receding jaw in N, chin in S
Posterior mandible - retromolar gap in N, not in S
Occipital shape - occipital bun in N, flatter occipital in S
Facial prognathism - S has flatter face
Canine fossa - in S, not N
Parietal bossing in S - house-shaped from behind
N brain size - 1500cc, EQ 6.9
S brain size - 1300cc, EQ 7.6
Paramasticatory use of anterior teeth
Were the anterior teeth adapted for gripping and tearing flesh?
Extremely worn anterior teeth
Midfacial prognathism, no canine fossa, swept back zygomatics
Due to high stress on anterior dentition?
Postcranial anatomy
Larger body mass than H. sapiens - 65-76kg Shorter and stockier - 150-170cm Large, wide, conical rib cage Large joints Shorter distal limb segments
Cold adaptions in Neanderthals
Glacial cycles during Neanderthal occupation
70-40kya - mean temperature 6C colder than today
200-40kya - generally colder than present, although variable
Bergmann’s Rule
Warm-blooded animals have larger body-mass in colder climates
Large size -> less surface area -> less heat loss
Neanderthals have:
short, wide pelvis
high body mass -> low surface:volume ratio
Allen’s Rule
Warm-blooded animals in cold climates have shorter appendages - limbs, tails, ears etc.
Reduced surface area -> less heat loss
Neanderthals have:
short distal limb segments
Asian Neanderthals
West Asia: Tabun, 120kya Shanidar, 70kya Teshik Tash, 70kya Amud, 50kya Kebara, 50kya
Less extreme anatomy than classic Neanderthals
Some similarities with H. sapiens, near early H. sapiens sites
Result of hybridisation in the Near East region?
Intentional burials?
Shanidar -flower burial?
Neanderthal DNA
What we know:
- Homo sapiens is not the closest sister group of Neanderthals - it is the Denisovans
- Modern human and Neanderthals, and Denisovans, hybridised
- Neanderthals and modern humans diverged at least 500kya - exact time debated
What we can infer:
- Homo sapiens may also have other more closely related sister group(s)
- Hybridisation between different hominin lineages was probably common during human evolution
Neanderthal/modern human hybridisation
Pestera cu Oase, Romania, 40kya
Suggested to be a hybrid based on unusual combination of Neanderthal and modern human traits
2016 DNA analysis demonstrates that Oase individual is 4th-6th generation Neanderthal-modern human hybrid
Pre-40kya Neanderthal, Denisovan, & Sapiens specimens - 13% showed evidence of hybridisation = very common
Neanderthal DNA in present-day humans
Non-uniform distribution
High concentration in skin and hair genes -> adaption to cold climates
Neanderthal alleles associated with disease risk
Deficiency of Neanderthal alleles implies removal through selection
Genes expressed in X chromosome - Neanderthal DNA reduced fertility in H. sapiens
Neanderthal extinction
- Climate-driven extinction
- Lived in a climatically very unstable period
- Ice Ages in Europe interspersed with warmer periods
- Difficult climatic conditions + isolation -> fragmentations and local extinctions
- Model supported by geographic distribution of last Neanderthals - southern Europe - Competition with H. sapiens
- Disappear soon after entrance of H. sapiens to Europe
- H. sapiens is a highly invasive species, related to many faunal extinctions
- Competitive exclusion?
- H. sapiens behaviourally superior to H. neanderthalensis? - Genetic assimilation/negative consequences of interbreeding
- No Neanderthal mtDNA in modern humans - most hybrids had Neanderthal father and H. sapiens mother?
- Little Neanderthal DNA in X-chromosome of modern humans
- Interbreeding affected Neanderthal fertility and they were genetically assimilated by modern humans?
- Late Neanderthals associated with Chatelperronian tools, similar to H. sapiens
- Interactions between Neanderthals and Sapiens may have been co-operative rather than antagonist
- Possibly didn’t recognise themselves as different - are they different species?
Neanderthal extinction likely a result of several factors
Neanderthal tools
Mousterian - Mode 3
Prepared core technology more complex than Acheulean
Some stone tools were hafted (attached to a handle), probably to make spears
Wooden spears - Schöningen, 400kya
Neanderthals and Middle Pleistocene hominins hunted large animals
Specialised toolkits
Neanderthal burials
Ritual nature or carried out to dispose of decomposing bodies?
Indicators: burial position, grave goods, lack of carnivore activity, relative completeness and articulation
Neanderthal ornamentation?
Found in association with Neanderthal remains
- Independent invention of Neanderthals?
- Evidence of acculturation associated with incoming modern humans?
- Result of modern human activity?
e.g.
Gibraltar - feathers from corvids and raptors
Cueva Antón ~50kya - ochre stained shell pendant
Maastricht Belvédère, 200-250kya - imported ochre. Medicine/insect repellent/curing hides/adhesive?
Neanderthal rock art and music
Cave art normally associated with H. sapiens
Art from Iberia has been shown to be ~40kya e.g. abstract symbol in Gorham’s Cave, Gibralta, >39kya; ‘seals’ in Nerja caves, Spain, ~42kya; hand prints in El Castillo cave, Spain, >37kya
Recent research has dated some sites to over 65kya (20kya before H. sapiens arrive in Europe) - La Pasiega Gallery, Spain
Symbolism and abstract thinking more developed in H. sapiens?
Divje Babe Cave, Slovenia, 50kya - Neanderthal ‘flute’ made from juvenile bear femur
Neanderthal behaviour
‘Modern’ traits found in Neanderthals:
- Complex tool technology
- Ambush hunting at close range using spears
- Evidence for burial but no/few grave goods
- Personal ornamentation, but in association with the appearance of H. sapiens in Europe
‘Modern’ traits not (yet) found in Neanderthals:
- Microblades and projectiles
- Ritual burial?
- Representative art, figurines
- Long distance trade networks
Anthropocentric to think these traits are important?
Need to compare Neanderthals with H. sapiens of the same period, not with contemporary modern humans.
