Neanderthals

Question 1

1. How old is the earliest fossil with Neanderthal autapomorphies?

And when did they finally go extinct?

Answer 1

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

Question 2

Where are Neanderthals generally thought to have existed

Answer 2

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

Question 3

Was there any overlap of Neanderthals and humans

Answer 3

yes: 26-54ky overlap

ample time for flow of genes and culture

Question 4

Give the key diagnostic features of a Neanderthal skull (6)

Answer 4

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)

Question 5

Give key features of Neanderthal post-crania

Answer 5

wide body with short limbs

barrel chest

wide pelvis with long, thin superior pubic ramus

pronounced muscle insertions eg olecranon and tibial tuberosity

Question 6

What does Neanderthal morphology mean? (4)

Answer 6

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

Question 7

What is Bergmann’s Rule

Answer 7

body mass increases in colder climates (reduce surface/volume ratio)

Question 8

What is the evidence Neanderthals were cold adapted

What is the impact of these adaptations

Answer 8

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

Question 9

What is Thompson’s Rule?

Allen’s Rule?

Ruff’s Rule?

Answer 9

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

Question 10

Would Neanderthals have worn clothes

Answer 10

considering higher cold temperatures, Neanderthals would have had to cover up to 80% of body in winter as well as hands and feet

Question 11

Compare the brains of Neanderthals and sapiens

Answer 11

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

Question 12

How does Neanderthal ontogeny differ from humans

Answer 12

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

Question 13

What did Zollikofer et al argue about Neanderthal ontogeny

Answer 13

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

Question 14

Give specifics of Neanderthal ontogeny

Answer 14

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

Question 15

Why is the hyoid important

What does it look like in Neanderthals

Answer 15

its size defines the range of tongue and laryngeal movement, affecting sound production

similar size and shape to sapiens

Question 16

What is the importance of the hypoglossal canal?

What does it look like in Neanderthals?

What does this suggest?

Answer 16

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

Question 17

How does the Neanderthal FOXP2 gene compare to that of humans

Answer 17

Neandthals had the 2 human specific aminoacid substiutions but may have been regulated differently

Question 18

Describe the hearing capacities of the SH hominins

Answer 18

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)

Question 19

Did Neanderthals make sophisticated tools?

Answer 19

yes: Mousterian stone tool industry (Mode 3)

Levallois points shafted onto spears for thrusting

made tools out of bone (inc human)

Question 20

Is there evidence of Neanderthals hunting large game

Answer 20

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?

Question 21

Give evidence of Neanderthals having abstract practices

Answer 21

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?

Question 22

How did Neanderthals use pigment

Answer 22

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

Question 23

Why do we think Neanderthals used feathers decoratively

Answer 23

from ~44kya, many bird bones (eg golden eagle), which are not dietary, have been found with cut marks exclusively on wings

Perensani (2011)

Question 24

Describe Neanderthal use of shells

Answer 24

perforated and pigment stained marine shells in Spain ~50kya

preference for coloured shells in middle Paleolithic layers/ Mousterial layers in Italy
Peresani, 2013

Question 25

What social practices do we think Neanderthals took part in

Answer 25

Care of the elderly and infirm

interpersonal violence

cannibalism

burial

Question 26

Why do we think Neanderthals cared for the infirm

Answer 26

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

Question 27

Give an example of Neanderthals using medicinal plants

Answer 27

poplar, which contains salicylic acid and may have been used as a painkiller (Weyrich et al., 2017).

Question 28

Is birth attendance uniquely human?

Answer 28

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.

Question 29

When does the earliest widely accepted evidence for care for illness and injury dates to ?

Give examples (3)

Give an earlier possible example

Answer 29

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

Question 30

How north may Neanderthals have stretched

How did this affect diet

Answer 30

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).

Question 31

How frequent was Neanderthal nutritional stress?

Answer 31

enamel hypoplasia suggests it was common and within the upper range of that observed within modern hunter-gatherer

Question 32

Why does Spikins argue it is unsurprising that Neanderthals were caring

Answer 32

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)

Question 33

What is a selective trade-off in the TRPM8 gene

Answer 33

whilst protecting from effects of cold is also associated with migraine susceptibility

Question 34

How does Spikins link autism to Neanderthal care

Answer 34

Positive selection for autism genes may also imply a level of social support for those who bring additional skills but may need more support

Question 35

Give evidence of cannibalism in Neanderthals

Answer 35

cut marks on bones in 12 individuals from 49kya in Spain

Question 36

Give evidence of interpersonal violence in Neanderthals

Answer 36

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.

Question 37

What is the evidence of Neanderthal burial

Answer 37

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)

Question 38

Why did Neanderthals go extinct? Give the four key hypotheses.

Answer 38

1. Climate – swift extreme climatic shifts
2. Disease – demographic attrition through truama and hardship + introduction of new disease by modern humans cf native americans
3. Direct competition – Steve Jones, UCL – sapiens’ more advanced technology meant that the conservative Neanderthals didn’t stand a chance – cf colonization of south america
4. Indirect competition - modern humans outcompeted them for food, causing their spatial contraction and population collapse

Question 39

What have we learnt about Neanderthals from aDNA?

Answer 39

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

Question 40

Describe assimilation events between sapiens and Neanderthals

Answer 40

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

Question 41

When did Neanderthals exist?

Where did they live?

Brain size?

Daily calorie intake?

Answer 41

~150kya-40kya (could be 430-28kya)

Europe, West and Central Asia, Siberia

1470cc

~4000calories/day

Question 42

Give a summary of Neanderthals

Answer 42

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

Question 43

What does the Neanderthal auditory tube look like

Answer 43

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)

Question 44

When did the LCA of sapiens and Neanderthals and Denisovans live

What does this suggest

Answer 44

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

Question 45

What is Ruff’s suggestion for how body shape has changed throughout human evolution in response to temperature changes

Answer 45

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).

Question 46

How does pelvic width change with latitude

Answer 46

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.

Question 47

Give an interpretation of the labial wear on Neanderthal teeth

Answer 47

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.

Question 48

Why might climate have contributed to Neanderthal extinction

Answer 48

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

Question 49

Who called Neanderthals the last true conservatives

Answer 49

No cumulative culture – the ‘last true conservatives’ (Steve Jones, 2006)

Question 50

Key citations for each of the following reasons for Neanderthal extinction
direct competition
indirect competition
climate
pathogen

Answer 50

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)

Question 51

Give example of inter-sapiens conflict in early Holocene

Neanderthal conflict?

Answer 51

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)

Question 52

What genetic evidence is there of humans being more aggressive than Neanderthals

Answer 52

 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

Question 53

What are arguments for Neanderthals being wiped out by indirect competition

Answer 53

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)

Question 54

How important was climate change to Neanderthal extinction

Answer 54

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)

Question 55

What is a problem for the division of labour hypothesis

Answer 55

Rosas (2015) found cultural striations in Neanderthal dental wear were greater in females - use of mouth as 3rd hand?

Question 56

Evidence of very early symbolism

Answer 56

Excalibur in SH hominins - ritual