Neanderthals Flashcards
- How old is the earliest fossil with Neanderthal autapomorphies?
And when did they finally go extinct?
Earliest >150kya but maybe 400kya
Controversial, fossils from 28kya but have disappeared almost everywhere else and meta analysis (Douka, 2014) suggests by 40kya extinction had occurred
Where are Neanderthals generally thought to have existed
Tend to think of Neanderthals as being predominantly in Europe but realistically it could be further east as Europe is simply the most excavated area in the world
Was there any overlap of Neanderthals and humans
yes: 26-54ky overlap
ample time for flow of genes and culture
Give the key diagnostic features of a Neanderthal skull (6)
large anterior teeth, often with labial wear
midfacial prognathism
retromolar space
receding frontal bone
NO chin, but multiple mental foramina
Taurodontism (this can occur in sapiens)
Give key features of Neanderthal post-crania
wide body with short limbs
barrel chest
wide pelvis with long, thin superior pubic ramus
pronounced muscle insertions eg olecranon and tibial tuberosity
What does Neanderthal morphology mean? (4)
Stronger, more robust skeleton + More pronounced muscle insertions = biomechanically more powerful and efficient
Hands are particularly strong (large hook of hamate, large phalangeal tubercles)
Different pelvic geometry - different pattern of (rotation at ) birth
Hyperpolar limb proportions and prognathic face/nose = cold adaptation
What is Bergmann’s Rule
body mass increases in colder climates (reduce surface/volume ratio)
What is the evidence Neanderthals were cold adapted
What is the impact of these adaptations
Bergmann’s rule=
Neanderthals: high BMI, short stature, wide pelvis – suggests high ratio of surface:volume
Allen’s Rule:
Neanderthals: short distal extremities – similar to cold adapted humans
Thompson’s Rule:
large nasal cavity, warms air before inhaling
Ruff’s Rule:
short and stocky - heat retention
This all means they may have been able to survive in temp 4 degrees colder than modern humans
What is Thompson’s Rule?
Allen’s Rule?
Ruff’s Rule?
Thompson: nasal aperture will be larger in colder climates
Allen: heat loss reduced by changing limb length
Ruff: cylinder body plan, independent of stature, associated with heat dissipation
Would Neanderthals have worn clothes
considering higher cold temperatures, Neanderthals would have had to cover up to 80% of body in winter as well as hands and feet
Compare the brains of Neanderthals and sapiens
similar absolute size
larger visual system in Neanderthals
Neanderthals had smaller adjusted cranial capacity
Pearce (2013) argues sapiens invested in ecological problem solving while Neanderthals invested in enhanced vision
How does Neanderthal ontogeny differ from humans
Grow more slowly than early hominins but faster than modern humans
Resorption and deposition during development is different - SH hominin sub adults show bone deposition on the maxilla where humans show resorption (evident from ~5yo) - this bone modelling is more similar to early Homo meaning modern humans have derived faces
What did Zollikofer et al argue about Neanderthal ontogeny
Using reconstructions of Neanderthal neonates and infants, they estimated Neanderthal brain size at birth was similar to that in recent Homo sapiens and most likely subject to similar obstetric constraints.
Neanderthal brain growth rates during early infancy were higher, however. This pattern of growth resulted in larger adult brain sizes but not in earlier completion of brain growth. Because large brains growing at high rates require large, late-maturing, mothers
it is likely that Neanderthal life history was similarly slow, or even slower-paced, than in recent H. sapiens.
Zollikofer et al 2008
Give specifics of Neanderthal ontogeny
use computerized fossil reconstruction and geometric morphometrics to show that characteristic differences in cranial and mandibular shape between Neanderthals and modern humans arose very early during development, possibly prenatally, and were maintained throughout postnatal ontogeny.
Postnatal differences in cranial ontogeny between the two taxa are characterized primarily by heterochronic modifications of a common spatial pattern of development.
Evidence for early ontogenetic divergence together with evolutionary stasis of taxon-specific patterns of ontogeny is consistent with separation of Neanderthals and modern humans at the species level.
Zollikofer 2001
Why is the hyoid important
What does it look like in Neanderthals
its size defines the range of tongue and laryngeal movement, affecting sound production
similar size and shape to sapiens
What is the importance of the hypoglossal canal?
What does it look like in Neanderthals?
What does this suggest?
carries CNXII which allows fine muscle control in tongue
similar size/ larger cf. modern humans
Neanderthals could have had a similar level of tongue control
How does the Neanderthal FOXP2 gene compare to that of humans
Neandthals had the 2 human specific aminoacid substiutions but may have been regulated differently
Describe the hearing capacities of the SH hominins
the skeletal anatomy in these hominids is compatible with a human-like pattern of sound power transmission through the outer and middle ear at frequencies up to 5 kHz, suggesting that they already had auditory capacities similar to those of living humans in this frequency range
(NB argued to be phylogenetically close to the Neandertals and are attributed to the species Homo heidelbergensis)
Did Neanderthals make sophisticated tools?
yes: Mousterian stone tool industry (Mode 3)
Levallois points shafted onto spears for thrusting
made tools out of bone (inc human)
Is there evidence of Neanderthals hunting large game
high levels of trauma
fractures of femur, fibula, and skull, thought to stem from hunting large game at close range
Shanidar 1: healed humerus fracture
Shanidar 3: fractured ribs, from horn or tusk?
Give evidence of Neanderthals having abstract practices
Perforated bone in Mousterian site from 55kya
- flute or tooth marks?
personal ornaments frmo 30kya (eg feathers, shells)
innovation or imitation
anterior teeth often have labial wear - para-masticatory use as seen in the Selk’nam
use of ochre and colour
NB: All of above dated between 50-40kya, very last populations of Neanderthals and could have been interacting with humans
Hyena bone with parallel lines from 60kya - counting?
How did Neanderthals use pigment
mostly MnO2 (black pigment) but red was occasionally used
> 500 blocks of pigment have been found in France >43kya
Zilhao (2010) argued ochre was used as an antiseptic
Why do we think Neanderthals used feathers decoratively
from ~44kya, many bird bones (eg golden eagle), which are not dietary, have been found with cut marks exclusively on wings
Perensani (2011)
Describe Neanderthal use of shells
perforated and pigment stained marine shells in Spain ~50kya
preference for coloured shells in middle Paleolithic layers/ Mousterial layers in Italy
Peresani, 2013
What social practices do we think Neanderthals took part in
Care of the elderly and infirm
interpersonal violence
cannibalism
burial
Why do we think Neanderthals cared for the infirm
Shanidar 1 - at a young age suffered a blow which damaged the left eye (possibly blinding him) and the brain area controlling the right side of the body, leading to a withered right arm and possible paralysis that also crippled his right leg. survived to ~35 and thought to have required daily care for over a decade
Neanderthal 1 - healed frontal bone, chronic inflammation, metastatic disease. died ~40 so must have been cared for
These are not isolated cases of care - such injuries are very common and it seems rare to reach adulthood without injury (Spikins, 2019)
this may have evolved in a similar way to cooperation in sapiens
small related groups likely led to inbreeding and high rates of genetic disease, requiring care
Give an example of Neanderthals using medicinal plants
poplar, which contains salicylic acid and may have been used as a painkiller (Weyrich et al., 2017).
Is birth attendance uniquely human?
NO - has been found in bonobos
As in humans, birth in bonobos was a social event, where female attendants provided protection and support to the parturient until the infant was born. Moreover, bystander females helped the parturient during the expulsive phase by performing manual gestures aimed at holding the infant, despite no physical restraint hindering bonobo birth
might be related to the high level of female gregariousness in these species.
the capacity of unrelated females to form strong social bonds and cooperate could have represented the evolutionary pre-requisite for the emergence of human midwifery.
When does the earliest widely accepted evidence for care for illness and injury dates to ?
Give examples (3)
Give an earlier possible example
when pro-social changes emerged (1.8mya/ emergence of Homo)
hominin from Dmanisi with tooth-loss and periodontal disease dated to 1.8mya
Homo ergaster with hypervitaminosis A or treponemal disease (1.6mya)
erectus with juvenile disc herniation (1.6mya)
2mya - Au. sediba juvenile from Malapa (MH1) with a bony tumour of the spine which would have limited movement and caused chronic pain and muscle spasm
How north may Neanderthals have stretched
How did this affect diet
far north as 55 degrees N
although highly debated Mousterian assemblages found near the Arctic Circle
Dependence on animal food for survival increases with increasing distance from the equator (Johnson, 2014), and moreover average prey size increases with cold and increasing latitude (Rodríguezet al., 2008) bringing elevated risks of injury and mortality through injury from hunting as well as through competition withother predators
selective pressures to adapt to the strains imposed by a high protein diet has also been argued to be the explanation for the large lower thorax of Neanderthals (Ben-Dor et al., 2016).
How frequent was Neanderthal nutritional stress?
enamel hypoplasia suggests it was common and within the upper range of that observed within modern hunter-gatherer
Why does Spikins argue it is unsurprising that Neanderthals were caring
In modern contexts high risk environments increase propensities to altruism through normal behavioural plasticity ac-cording to context (Li et al., 2013) as does dependence on collaborative hunting of very large and dangerous game (eg whale hunting, (Heinrich et al., 2004)
What is a selective trade-off in the TRPM8 gene
whilst protecting from effects of cold is also associated with migraine susceptibility
How does Spikins link autism to Neanderthal care
Positive selection for autism genes may also imply a level of social support for those who bring additional skills but may need more support
Give evidence of cannibalism in Neanderthals
cut marks on bones in 12 individuals from 49kya in Spain
Give evidence of interpersonal violence in Neanderthals
healed cranial vault fracture from a sharp implement
36kya
Zollikofer (2002) hypothesized that the high intra-group damage potential inherent to weapons might have represented a major factor during the evolution of hominid social behavior.
What is the evidence of Neanderthal burial
60kya: Shanidar 4 – pollen found in high density around the flexed body – bed of flowers?
70kya: 9yo child, Ibex burial + fire marks
Ritualistic or pragmatic (ie disposal)
Why did Neanderthals go extinct? Give the four key hypotheses.
- Climate – swift extreme climatic shifts
- Disease – demographic attrition through truama and hardship + introduction of new disease by modern humans cf native americans
- Direct competition – Steve Jones, UCL – sapiens’ more advanced technology meant that the conservative Neanderthals didn’t stand a chance – cf colonization of south america
- Indirect competition - modern humans outcompeted them for food, causing their spatial contraction and population collapse
What have we learnt about Neanderthals from aDNA?
Smaller historical effective population size – through time Neanderthals had smaller populations than humans
Long runs of homozygosity – highly inbred and the individual who was sequenced had parents who were v closely related (double first cousins or grandfather grand daughter, half-siblings, or uncle and niece)
Neanderthals went through boom or bust – expand massively then cannot sustain and local populations go extinct – genome from Chagyrskaya cave (Altai mountains, Russia) is more closely related to younger European genomes than to older indivduals from Denisova – independent dispersals to the east – lots of expansions and extinctions
Culturally different expansions
Describe assimilation events between sapiens and Neanderthals
Human ancestor impacted neanderthals’ genomes – early on Neanderthals assimilated humans
Humans later assimilated neanderthals – children grew up human not N
NE also interacted with DEN – first gen offspring of one of each N mother and D father
Neanderthals were assimilated into very first movement out of Africa
Never found a fragement of DEN inside N genome
When did Neanderthals exist?
Where did they live?
Brain size?
Daily calorie intake?
~150kya-40kya (could be 430-28kya)
Europe, West and Central Asia, Siberia
1470cc
~4000calories/day
Give a summary of Neanderthals
shared LCA with DEN who expanded OoA ~450kya (NE/DEN LCA had separated by 450kya)
NE evolved in W Eurasia and share features with middle Pleistocene European hominins
Become extinct ~40kya, possibly longer in Gibraltar
fluctuating demography with expansions followed by extinctions
complex behavior likely influenced by humans in Eurasia
What does the Neanderthal auditory tube look like
Neanderthal adults exhibiting primitively tall and narrow nasopharynges with infant-like horizontal CET and choanal orientation.
As horizontal CET orientation is associated with increased Otitis Media incidence in infants and children until around age six, its appearance in Neanderthal adults strongly indicates persistence of high Otitis Media susceptibility at this time. This could have compromised fitness and disease load relative to sympatric modern humans, affecting Neanderthals’ ability to compete within their ecological niche, and potentially contributing to their rapid extinction
(Pagano 2019)
When did the LCA of sapiens and Neanderthals and Denisovans live
What does this suggest
in Africa 550‐700 Ka
- A recent African origin of all humans 200,000 – 300,000 years ago
- An African phase prior to global expansion
- Reduced non‐African diversity
What is Ruff’s suggestion for how body shape has changed throughout human evolution in response to temperature changes
Heat dissipation is principally achieved through sweating.
In the closed, humid environment of a forest, sweating is not a very efficient process. Thus a forest-adapted hominid would need to generate less heat ( ie to be small.)
Hence Australopithecus has a small stature well suited for forest living.
However, taller Homo erectus would be restricted to open country
Neanderthals have broader pelvises bc individuals should have proportionally more body mass compared with their SA (broader trunk).
How does pelvic width change with latitude
As the average temperature decreases, the need to regulate body temperature
requires the ratio of surface area to mass to decrease; in other words, individuals should have proportionally more body mass compared with their surface area. This is achieved by having a fatter cylinder – a broader trunk.
The pelvic width therefore increases in populations at higher latitudes – in Europeans and north Asians, for instance.
Give an interpretation of the labial wear on Neanderthal teeth
they held animal hides with their teeth while scraping off the flesh with stone tools. They could then perforate these skins and wear them as clothing.
This interpretation is based on the labial wear observed in the Selk’nam people of Patagonia, Argentina.
Why might climate have contributed to Neanderthal extinction
o Neanderthals already adapted for cold – did not need to make clothes
Sapiens did – used needles – helped survive MIS3 (Gilligan, 2008)
Neanderthals lacked the cultural capabilities to adapt
• No cumulative culture – the ‘last true conservatives’ (Steve Jones, 2006)
o Eruption of Italian volcano
Disruption to vegetation (indicated by low pollen in layers of Mezmaiskaya Cave ~40kya) - Shackley 2010
• Neanderthals couldn’t adapt
o Important regionally
In Central Europe, Dansgaard-Oeschge events lead to an earlier Neanderthal extinction by ~ 500-1,500 years, but globally absence of DO events leads to only ~2kya difference
Who called Neanderthals the last true conservatives
No cumulative culture – the ‘last true conservatives’ (Steve Jones, 2006)
Key citations for each of the following reasons for Neanderthal extinction direct competition indirect competition climate pathogen
direct competition:
Lahr (2016) - Holocene HGs
Churchill (2009) - Shanidar 3
Parra (2014) - ADSL & aggression
indirect competition:
Kuhn (2006) - sex DoL
Fa (2013) - dogs helped hunt rabbits for sapiens
climate:
Gilligan (2008) - clothes
Shackley (2010) - used pollen to show volcano disrupts vegetation
Timmermann (2020)
pathogen:
Underdown (2016) -AMH had protection from bacterial sepsis
Pagano (2019)
Give example of inter-sapiens conflict in early Holocene
Neanderthal conflict?
in Kenya
• Articulated skeletons of individuals killed in conflict
o At least 5 have sharp force trauma and one cranium still has an obsidian bladelet (Lahr, 2016)
Shanidar 3
• Only 1 rib properly damaged, most consistent with injury by a low-mass, low-kinetic energy projectile weapon (36kya) (Churchill, 2009)
What genetic evidence is there of humans being more aggressive than Neanderthals
Neanderthals lack genes eg ADSL which is associated with aggression (Parra et al., 2014)
• Increased predation in Africa meant sapiens were more aggressive (Ko, 2016)
Unclear whether absence of ADSL would increase or decrease aggression
What are arguments for Neanderthals being wiped out by indirect competition
inferior cognition sexual DoL Dogs • Altai dog – 33kya • Need to shift to rabbits – sapiens could but Neanderthals could not – still reliant on less common large mammals (Fa, 2013)
How important was climate change to Neanderthal extinction
o Important regionally
In Central Europe, Dansgaard-Oeschge events lead to an earlier Neanderthal extinction by ~ 500-1,500 years, but globally absence of DO events leads to only ~2kya difference (Timmermann, 2020)
What is a problem for the division of labour hypothesis
Rosas (2015) found cultural striations in Neanderthal dental wear were greater in females - use of mouth as 3rd hand?
Evidence of very early symbolism
Excalibur in SH hominins - ritual
