South African archaic human species
Homo naledi
is an
extinct
species
of
archaic human
discovered in 2013 in the
Rising Star Cave
system,
Gauteng
province, South Africa (See
Cradle of Humankind
), dating to the
Middle Pleistocene
335,000?236,000 years ago. The initial discovery comprises 1,550 specimens of bone, representing 737 different skeletal elements, and at least 15 different individuals. Despite this exceptionally high number of specimens, their classification with other
Homo
species remains unclear.
Along with similarities to contemporary
Homo
, they share several characteristics with the ancestral
Australopithecus
as well as early
Homo
(
mosaic evolution
), most notably a small cranial capacity of 465?610 cm
3
(28.4?37.2 cu in), compared with 1,270?1,330 cm
3
(78?81 cu in) in modern humans. They are estimated to have averaged 143.6 cm (4 ft 9 in) in height and 39.7 kg (88 lb) in weight, yielding a small
encephalization quotient
of 4.5.
H. naledi
brain anatomy seems to have been similar to contemporary
Homo
, which could indicate comparable cognitive complexity. The persistence of small-brained humans for so long in the midst of bigger-brained contemporaries revises the previous conception that a larger brain would necessarily lead to an evolutionary advantage, and their mosaic anatomy greatly expands the known range of variation for the genus.
H. naledi
anatomy indicates that, though they were capable of long-distance travel with a humanlike stride and gait, they were more
arboreal
than other
Homo
, better adapted to climbing and
suspensory behaviour
in trees than
endurance running
. Tooth anatomy suggests consumption of gritty foods covered in particulates such as dust or dirt. Though they have not been associated with stone tools or any indication of material culture, they appear to have been dextrous enough to produce and handle tools, and therefore may have manufactured
Early
or
Middle
Stone Age
industries
since no other human species in the vicinity at that time has been discovered. It has also been controversially postulated that these individuals were given funerary rites, and were carried into and placed in the chamber. They may also have carved crosshatched rock signs in a passage to what could be a funeral chamber (2023).
[2]
Discovery
[
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]
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On 13 September 2013 while exploring the
Rising Star Cave
system in the
Cradle of Humankind
, South Africa, cavers Rick Hunter and Steven Tucker found
hominin
fossils at the bottom of the
Dinaledi Chamber
.
[3]
On the 24th, they returned to the chamber and took photos, which they showed to South African
palaeoanthropologists
Pedro Boshoff and
Lee Rogers Berger
on October 1.
[3]
Berger assembled an excavation team which included Hunter and Tucker, the so-called "
Underground Astronauts
".
[4]
The chamber had been entered at least once before, by cavers in the early 1990s. They rearranged some bones and may have caused further damage, although much of the floor in the chamber had not been walked on prior to 2013.
[5]
It lies about 80 m (260 ft) from the main entrance, at the bottom of a 12 m (39 ft) vertical drop, and the 10 m (33 ft) long main passage is only 25?50 cm (10 in ? 1 ft 8 in) at its narrowest.
[5]
In total, more than 1,550 pieces of bone belonging to at least fifteen individuals (9 immature and 6 adults)
[6]
have been recovered from the
clay
-rich sediments. Berger and colleagues published the findings in 2015.
[7]
The fossils represent 737 anatomical elements ? including portions of the skull, jaw, ribs, teeth, limbs, and inner ear bones ? from old, adult, young, and infantile individuals. There are also some
articulated
or near-articulated elements, including the skull with the jaw bone, and nearly complete hands and feet.
[7]
[5]
With the number of individuals of both sexes across several age demographics, it is the richest assemblage of associated fossil hominins discovered in Africa. Aside from the
Sima de los Huesos
collection and later
Neanderthal
and modern human samples, the excavation site has the most comprehensive representation of skeletal elements across the lifespan, and from multiple individuals, in the hominin fossil record.
[7]
The
holotype specimen
, DH1, comprises a male partial
calvaria
(top of the skull), partial
maxilla
, and nearly complete
jawbone
. The
paratypes
, DH2 through DH5, all comprise partial calvaria. Berger and colleagues named the species
Homo naledi
in 2015, the specific name meaning "star" in the
Sotho language
, because the remains came from Rising Star Cave.
[7]
The remains of at least three additional individuals (two adults and a child) were reported in the Lesedi Chamber of the cave by
John Hawks
and colleagues in 2017.
[8]
Classification
[
edit
]
In 2017, the Dinaledi remains were dated to 335,000?236,000 years ago in the
Middle Pleistocene
, using
electron spin resonance
(ESR) and
uranium?thorium
(U-Th) dating on three teeth, and U-Th and
paleomagnetic
dating of the sediments they were deposited in.
[1]
The fossils were previously thought to have dated to 1?2 million years ago
[7]
[9]
[10]
[4]
because no similarly small-brained hominins had previously been known from such a recent date in Africa.
[11]
The smaller-brained
Homo floresiensis
of Indonesia lived on an isolated island, and apparently became extinct shortly after the arrival of modern humans.
[12]
The ability of such a small-brained hominin to have survived for so long in the midst of bigger-brained
Homo
greatly revises previous conceptions of
human evolution
and the notion that a larger brain would necessarily lead to an evolutionary advantage.
[11]
Their mosaic anatomy also greatly expands the range of variation for the genus.
[13]
H. naledi
is hypothesised to have branched off very early from contemporaneous
Homo
. It is unclear whether they branched off at around the time of
H. habilis
,
H. rudolfensis
, and
A. sediba
, are a
sister taxon
to
H. erectus
and the contemporaneous large-brained
Homo
, or are a sister taxon to the descendants of
H. heidelbergensis
(modern humans and Neanderthals). This would mean that they branched off from contemporary
Homo
at latest before 900,000 years ago, and possibly as early as the
Pliocene
. It is also possible their ancestors speciated after an interbreeding event between
Homo
and late australopithecines.
[11]
Looking at the skull,
H. naledi
has the closest
affinities
to
H. erectus
.
[13]
It is unclear if these
H. naledi
were an isolated population in the Cradle of Humankind, or if they ranged across Africa. If the latter, then several
gracile
hominin fossils across Africa which have traditionally been classified as late
H. erectus
could potentially represent
H. naledi
specimens.
[14]
Anatomy
[
edit
]
Skull
[
edit
]
Two male
H. naledi
skulls from the Dinaledi chamber had cranial volumes of about 560 cm
3
(34 cu in), and two female skulls 465 cm
3
(28.4 cu in). A male
H. naledi
skull from the Lesedi chamber had a cranial volume of 610 cm
3
(37 cu in). The Dinaledi specimens are more similar to the cranial capacity of australopithecines. For comparison,
H. erectus
averaged about 900 cm
3
(55 cu in),
[8]
and modern humans 1,270 and 1,130 cm
3
(78 and 69 cu in) for males and females respectively.
[15]
The Lesedi specimen is more within the range of
H. habilis
and
H. e. georgicus
. The
encephalization quotient
of
H. naledi
was estimated at 3.75, which is the same as the pygmy
H. floresiensis
, but notably smaller than all other
Homo
. Contemporary
Homo
were all above 6,
H. e. georgicus
at 3.55, and
A. africanus
at 3.81.
[16]
It is unclear if
H. naledi
inherited small brain size from the last common
Homo
ancestor, or if it was evolved secondarily more recently.
[17]
The skull shape is more similar to
Homo
, with a slenderer shape, the presence of
temporal
and
occipital lobes
of the brain, and reduced
post-orbital constriction
, with the skull not becoming narrower behind the eye-sockets.
[7]
[17]
The
frontal lobe
morphology is more or less the same in all
Homo
brains despite size, and differs from
Australopithecus
, which has been implicated in the production of tools, the development of language, and sociality.
[17]
Like modern humans, but unlike fossil hominins, including South African australopithecines,
H. erectus
, and Neanderthals, the permanent 2nd molar erupted comparatively late in life, emerging alongside the premolars instead of before, which indicates a slower maturation unusually comparable to modern humans.
[18]
The tooth formation rate of the front teeth is also most similar to modern humans.
[19]
The overall size and shape of the molars most closely resemble those of three unidentified
Homo
specimens from the local
Swartkrans
and East African
Koobi Fora
Caves, and are similar in size (but not shape) to Pleistocene
H. sapiens
. The necks of the molars are proportionally similar to those of
A. afarensis
and
Paranthropus
.
[20]
Unlike modern humans and contemporary
Homo
,
H. naledi
lacks several accessory dental features, and has a high frequency of individuals who present main
cusps
, namely the
metacone
(midline on the tongue-side) and
hypocone
(to the right on the lip-side) on the 2nd and 3rd molars, and a Y-shaped hypoconulid (a ridge on the lip-side towards the cheek) on all three molars. Nonetheless,
H. naledi
also has many dental similarities with contemporary
Homo
.
[21]
The
anvil
(a
middle ear bone
) more resembles those of chimps, gorillas, and
Paranthropus
than
Homo
.
[22]
Like
H. habilis
and
H. erectus
,
H. naledi
has a well-developed brow-ridge with a fissure stretching across just above the ridge, and like
H. erectus
a pronounced
occipital bun
.
H. naledi
has some facial similarities with
H. rudolfensis
.
[21]
Build
[
edit
]
The
H. naledi
specimens are estimated to have, on average, stood around 143.6 cm (4 ft 9 in) and weighed 39.7 kg (88 lb). This body mass is intermediate between what is typically seen in
Australopithecus
and
Homo
species. Like other
Homo
, male and female
H. naledi
were likely about the same size, males on average about 20% larger than females.
[16]
A juvenile specimen, DH7, is skeletally consistent with a growth rate similar to the faster ape-like trajectories of MH1 (
A. sediba
) and
Turkana boy
(
H. ergaster
). Because dental development is so similar to that of modern humans, a slower maturation rate is not completely out of the question. Using the faster growth rate, DH7 would have died at 8?11 years old, but using the slower growth, DH7 would have died at 11?15 years old.
[23]
Concerning the
spine
, only the 10th and 11th
thoracic vertebrae
(in the chest region) are preserved from presumably a single individual, which are proportionally similar to those of contemporary
Homo
, though are the smallest recorded of any hominin. The two transverse processes of the vertebra, which jut out diagonally, are most similar to those of Neanderthals. The
neural canals
within are proportionally large, similar to modern humans, Neanderthals, and
H. e. georgicus
. The 11th rib is straight like that of
A. afarensis
, and the 12th rib is robust in cross-section like that of Neanderthals. Like Neanderthals, the 12th rib appears to have supported strong
intercostal muscles
above, and a strong
quadratus lumborum muscle
below. Unlike Neanderthals, there was weak attachment to the
diaphragm
. Overall, this
H. naledi
specimen appears to have been small-bodied compared with other
Homo
species, though it is unclear if this single specimen is representative of the species.
[24]
The shoulders are more similar to those of australopithecines, with the
shoulder blade
situated higher on the back and farther from the midline, short
clavicles
, and little or no humeral torsion.
[7]
Elevated shoulder and clavicle bones indicate a narrow chest.
[24]
The
pelvis
and legs have features reminiscent of
Australopithecus
, including anterposteriorly compressed (from front to back)
femoral necks
, mediolaterally compressed (from left to right)
tibiae
, and a somewhat circular
fibular neck
;
[25]
[26]
which indicate a wide abdomen. This combination would preclude efficient
endurance running
in
H. naledi
, unlike
H. erectus
and descendants. Instead,
H. naledi
appears to have been more
arboreal
.
[24]
Limbs
[
edit
]
Fossil right hand of
H. naledi
(bottom view left, top view right) ? Scale 5 cm (2 in)
Fossil right foot of
H. naledi
, A) top view, B) left-side view, C) arch ? Scale 10 cm (3.9 in)
The
metacarpal bone
of the thumb, which is used in holding and manipulating large objects, was well-developed and had strong crests to support its
opponens pollicis muscle
used in precision-pinch gripping, and its
thenar muscles
. This is more similar to other
Homo
than
Australopithecus
.
H. naledi
appears to have had strong
flexor pollicis longus muscles
like modern humans, with humanlike palm and finger pads, which are important for forceful gripping between the thumb and fingers. Unlike
Homo
, the
H. naledi
thumb metacarpal joint is comparably small relative to the thumb's length, and the thumb
phalangeal
joint is flattened. The distal thumb phalanx bone is robust, and proportionally more similar to those of
H. habilis
and
P. robustus
.
[27]
The metacarpals of the other fingers share adaptations with modern humans and Neanderthals to be able to cup and manipulate objects, and the
wrist joint
is broadly similar to that of modern humans and Neanderthals. Conversely, the proximal phalanges are curved and are almost identical to those of
A. afarensis
and
H. habilis
, which is interpreted as an adaptation for climbing and
suspensory behaviour
. Such curvature is more pronounced in adults than juveniles, suggesting that adults climbed just as much or more so than juveniles, and this behaviour was commonly done. The fingers are proportionally longer than those of any other fossil hominin, other than the arboreal
Ardipithecus ramidus
and a modern human specimen from
Qafzeh cave
, Israel, which is consistent with climbing behaviour.
[27]
H. naledi
was a
biped
and stood upright.
[7]
Like other
Homo
, they had strong
insertion
for the
gluteus
muscles, well-defined
linea aspera
(a ridge running down the back of the femur), thick
patellae
, long tibiae, and gracile fibulae. These indicate that they were capable of long-distance travel.
[26]
The
H. naledi
foot was similar to that of modern humans and other
Homo
, with adaptations for
bipedalism
and a humanlike gait. The
heel bone
has a low orientation, comparable to those of non-human great apes, and the
ankle bone
has a low declination, which possibly indicate the foot would have been subtly stiffer during the stance phase of walking before the foot pushed off the ground.
[28]
Pathology
[
edit
]
The adult right
mandible
U.W. 101-1142 has a bony lesion, suggestive of a
benign tumour
. The individual would have experienced some swelling and localised discomfort, but the tumour's position near the
medial pterygoid muscle
(likely causing discomfort on the jaw hinge) may have impeded function of the muscle, and changed elevation of the right side of the jaw.
[29]
Dental defects in
H. naledi
specimens during 1.6?2.8 and 4.3?7.6 months of development were most likely caused by seasonal stressors. This may have been due to extreme summer and winter temperatures causing food scarcity. Minimum winter temperatures of the area average about 3 °C (37 °F), and can drop below freezing. Staying warm for an infant of the small-bodied
H. naledi
would have been difficult, and winters likely increased susceptibility to respiratory diseases. Environmental stressors are consistent with present-day
flu seasons
in South Africa peaking during winter, and paediatric
diarrhoea
hospitalisation being most frequent at the height of the rainy season in summer.
[30]
Local hominins were likely preyed upon by large carnivores, such as lions, leopards, and hyaenas. There seems to be a distinct paucity of large carnivore remains from the northern end of the Cradle of Humankind, where Rising Star Cave is located, possibly because carnivores preferred the Blaaubank River to the south which may have offered better hunting grounds with a greater abundance of large prey items. Alternatively, because many more sites are known in the south than the north, carnivore spatial patterns may not be well-represented by the fossil record (
preservation bias
).
[31]
Culture
[
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]
Food
[
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]
Dental chipping and wearing indicates the habitual consumption of small hard objects, such as dirt and dust, and cup-shaped wearing on the back teeth may have stemmed from gritty particles. These could have originated from unwashed roots and tubers. Alternatively, aridity could have stirred up particulates onto food items, coating food in dust. It is possible that they commonly ate larger hard items, such as seeds and nuts, but these were processed into smaller pieces before consumption.
[32]
[33]
H. naledi
occupied a seemingly unique
niche
from previous South African hominins, including
Australopithecus
and
Paranthropus
. The teeth of all three species indicate that they needed to exert high shearing force to chew through perhaps plant or muscle fibres. The teeth of other
Homo
cannot produce such high forces perhaps due to the use of some food processing techniques, such as cooking.
[32]
Technology
[
edit
]
H. naledi
could have produced
Early Stone Age
(
Acheulean
and possibly the earlier
Oldowan
) or
Middle Stone Age
industries
because they have the same adaptations to the hand as other human species which are implicated in tool production.
[11]
[16]
H. naledi
is the only identified human species to have existed during the early Middle Stone Age of the
Highveld
region, South Africa, possibly indicating that this species manufactured and maintained this tradition at least during this time period. In this scenario, such industries and stone cutting techniques would have evolved independently several times among different
Homo
species and populations, or were transported over long distances by the inventors or apprentices and taught.
[11]
Funerals
[
edit
]
In 2015, archaeologist Paul Dirks, Berger, and colleagues concluded that the bodies had to have been deliberately carried and placed into the chamber by people because they appear to have been intact when they were first deposited in the chamber. There is no evidence of trauma from being dropped into the chamber nor of predation, and there is exceptional preservation. The chamber is inaccessible to large predators, appears to be an isolated system, and has never been flooded. That is, natural forces were not at play.
[5]
There is no hidden shaft through which people could have accidentally fallen in, and there is no evidence of some catastrophe which killed all the individuals inside the chamber. They said it is possible that the bodies were dropped down a chute and fell slowly due to irregularity and narrowness of the path down, or a soft mud cushion to land on. In whatever scenario, the morticians would have required artificial light to navigate the cave. The site was used repeatedly for burials as the bodies were not all deposited at the same time.
[5]
In 2016, palaeoanthropologist Aurore Val countered that such preservation may have been due to
mummification
rather than careful burial, and the absence of
long bone
heads is reminiscent of predation. She believes that discounting natural forces such as flooding for depositing the bodies is unjustified. She identified evidence of damage done by beetles, beetle larvae, and snails, which facilitate decomposition. The chamber does not present ideal conditions for snails, nor does it contain snail shells, which would indicate decomposition initiated before deposition in the chamber.
[34]
In 2017, Dirks, Berger, and colleagues reaffirmed that there is no evidence of water flow into the cave, and that it is more likely that the bodies were deliberately deposited into the chamber. They said it is possible that they were deposited by contemporary
Homo
, such as the ancestors of modern humans, rather than other
H. naledi
, but that the cultural behaviour of funerary practises is not impossible for
H. naledi
. They proposed that placement in the chamber may have been done to remove decaying bodies from a settlement, prevent scavengers, or as a consequence of social bonding and grief.
[11]
In 2018, anthropologist Charles Egeland and colleagues echoed Val's sentiments, and stated that there is insufficient evidence to conclude that a hominid species had developed a concept of the
afterlife
so early in time. They said that the preservation of the Dinaledi individuals is similar to those of baboon carcasses which accumulate in caves, either by natural death of cave-dwelling baboons, or by a leopard dragging in carcasses.
[35]
In 2021, following the analysis of the bone fragments of an immature individual, Juliet Brophy and Berger again stated that the
H. naledi
remains were purposefully interred by some human species.
[36]
This would make
Homo naledi
the oldest evidence of burial by hominids.
[37]
The findings are disputed.
[38]
Gallery
[
edit
]
-
-
-
-
-
-
-
-
-
-
-
-
-
(A,B,C,D) Views of one
tibia
-
-
(1) adult right foot, (2) juvenile left, (3,4) adult left, (5) juvenile right
See also
[
edit
]
References
[
edit
]
- ^
a
b
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Further reading
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