A cremation from the Early Holocene of Malawi.

All modern Human societies feel the need to dispose of the dead in specific, often highly ritualised, ways, and this appears to have been the case for much of the history of our species, and potentially other members of the genus Homo. Cremation is an effective way to achieve this, transforming the recognisable remains of a member of the community into a small amount of ash and calcined bone fragments in a relatively short period of time. However, this process also requires the effort and co-ordination to build a funeral pyre of sufficient size, and does not appear to have become common or widespread until the Middle Holocene.

The oldest known assemblage of burned Human remains comes from Lake Mungo in New South Wales and have been dated to about 40 000 years before the present, although these remains appear to have gone through a multi-stage ritual process which included burning, rather than being a single cremation event as the main way to deal with the remains. The oldest evidence of a body being burned on a pyre comes from a site called Xaasaa Na′ (Upward Sun River) in the Alaskan Arctic, where a child of about three years, who was burned about 11 500 years ago. The oldest known burned remains in Africa are about 7500 years old and come from Egypt, but it is unclear whether these were deliberately cremated. The oldest known evidence for deliberate cremation in Africa comes from Njoro River Cave in Kenya, and is only about 3300 years old.

In a paper published in the journal Science Advances on 1 January 2026, a team of scientists led by Jessica Cerezo-Román of the Department of Anthropology at the University of Oklahoma present evidence for a deliberate cremation, from the Hora 1 archaeological site at the foot of Mount Hora in the Mzimba District of Northern Malawi.

Mount Hora is a 110 m high granite inselberg (isolated hill) rising above the Kasitu River Valley, which forms a distinctive regional landmark. The Hora 1 archaeological site is a rock shelter at the base of this hill, opening to the east and incorporating a flat dry area of about 80 m². This site is easily accessed from all directions, and contains no enclosed areas which could act as a natural furnace. No other natural rock shelters could be found in the region. Hora 1 has produced a record of Human occupation stretching over more than 21 000 years, and appears to have been used for mortuary practices between about 16 000 years ago and about 8000 years ago.

HOR-1 site (11°39′S, 33°39′E; 1470 m above mean sea level, white stipples) relative to published excavated Later Stone Age (LSA) sites. Light green in inset outlines the Zambezian Biome. Cluster of sites near Mpunzi (or Mphunzi) Mountain includes six additional sites, three with human remains (Mtuzi, Changoni Bible School, and Chencherere II). Cerezo-Román et al. (2026).

This site was first excavated by Desmond Clark and William Rangeley in the 1950s, who excavated an area of 7-8 m² (Clark and Rangeley did not record measurements of the area they excavated, but did take photographs, which have been used to reconstruct the extent of their work). These excavations produced the almost complete remains of two adult Humans, HOR-1 (or UTC-242), a male directly dated to between 9081 and 8725 years before the present, and Hora 2 (or UTC-243), a female directly dated to between 8172 and 7875 years before the present. Both of these skeletons have yielded ancient DNA, suggesting a relationship to modern hunter-gatherer populations in Southern Africa.

Between 2016 and 2019, the Malawi Ancient Lifeways and Peoples Project carried out further excavations at the site, removing and sieving 7.8 m³ of sediment from two blocks which produced about 46 500 objects, including the almost incomplete skeletons of two male infants (Kahora 1 and Kahora 2), which have been indirectly dated (i.e. dated from material found with them, not directly from the skeletal material) to about 14 000 and about 16 000 years before the present, and which have also yielded ancient DNA, as well as fragmentary remains attributed to at least four additional adults (Hora 4 to Hora 7) and five additional non-adults (Kahora 2 to Kahora 3), as well as numerous animal remains. Examination of these fragmentary remains has suggested that they may have been subjected to complex mortuary practices, including secondary burials (i.e. burials either after being buried once and then excavated, or after having been either left unburied for an extensive period, or having been subjected to ritual behaviours which substantially altered the remains), which may have included ritual token-taking.

Views of HOR-1. (A) Plan view of the HOR-1 site georectified onto drone imagery, showing the open overhang, accessibility, and Malawi Ancient Lifeways and Peoples Project (MALPP) excavation Area I and II. (B) View from the north of the open site access. (C) Plan view of the Malawi Ancient Lifeways and Peoples Project Area I excavations, Kahora 1 and Kahora 2 infant burials, and Hora 3 remains, with the outline of the original 1950 excavations and position of the Hora 2 skeleton by Clark and Rangely, reconstructed using an archival site photograph. Cerezo-Román et al. (2026).

Cerezo-Román et al. report the discovery of a large ash feature at Hora 1, along with an associated set of remains attributed to a single cremated adult, who they designate as Hora 3. Hora 3 is calculated to have been chronologically intermediate between Hora 2 and Kahora 1. As such it appears to form part of a genetically continuous population which lived in the area across the Pleistocene-Holocene boundary, although (unsurprisingly) it was not possible to extract DNA from the cremated remains.

The 'ash feature' forms a large cemented block, dated to about 9500 years before the present, which overlies a Pleistocene sediment sequence which begins 17 000 years ago and ends about 12 000 years ago, and is overlain by younger Early Holocene deposits. At the core of this feature they found a set of Human remains identified as 'Cluster 1), which lie on top of an ash layer 10-15 cm thick, and roughly 2.5 m by 1.5 m in extent. Below this are a series of deeper lenses of consolidated ashes, charcoal, and rubified sediment, together referred to as the 'Lower Ash', which are interpreted as evidence for a series of consecutive burn events at the same location.

The Cluster 1 remains sit on top of a layer of incompletely combusted charcoal, and are in turn overlain by several more stratified lenses of ash and rubified sediment. A second set of remains was identified to the northeast of Cluster 1 and 5-10 cm deeper. These were identified as Cluster 2, but appear to be a part of the same individual which became detached during the cremation event.

Plan view of the exposed ash complex, showing the large spatial extent. Jessica Thompson in Cerezo-Román et al. (2026).

Twenty radiocarbon dates were obtained from the core pyre and larger ash feature, providing a chronology for the site. Dates obtained below the core feature begin around 12 699 years before the present, with the latest coming from 9918 years before the present. Layers of ash above the core feature produced dates ranging from 9540 to 9454 years before the present, while a layer of ash on the eastern excavation wall was dated to between 9538 and 9455 years before the present. A Land Snail bead from a layer immediately above the core combustion event provided a date of between 9537 and 9441 years before the present, and is taken as evidence of this layer being covered over quickly by non-combustion sediments. Another large combustion event was indicated by a stratigraphically higher layer of ash, which produced a date of between 9452 and 9142 years before the present, i.e. less than 400 years after the core event, although this layer produced no Human remains. Above this were further layers of non-combustion deposits dating from between less than 9403 and 9031 years before the present. 

Spatial relationships of dated materials, samples, and human remains. HOR-1 excavations in profile (A) and plan (B) views showing positions of dated materials and adult cremated remains with different degrees of certainty (LH indicates 'likely human' based on size, texture, and preservation, but not morphologically diagnostic). (C) shows phytolith (green) and micromorphology (white boxes) sample positions. Thick blue line shows how the profiles correspond in each view. C1 indicates Cluster 1 and C2 indicates Cluster 2. Jessica Thompson in Cerezo-Román et al. (2026).

Phytoliths are produced by plants as a way of handling silica absorbed with water from the soil. Most plants produce phytoliths to some extent, but Monocotyledons, such as Grasses and Palms, which utilise phytoliths both as structural support and a defence against herbivores (silica phytoliths quickly wear down the teeth of animals which lack specific adaptations to deal with them), produce phytoliths which can often be used to identify the maker to genus or even species level. Phytoliths recovered from the Hora 1 pyre deposits imply that the majority of the ash was generated by the burning of wood, but that non-woody plant fibres were also present. 

Sediments and ash layers of the pyre deposit. (A) view southwest of the feature 50 cm south of Cluster 1 of the remains. In both (A) and (B) black arrows indicate the top of micromorphology sample MALAPP 834 and white arrows show the distal radius from Cluster 1 in situ. Dashed box is the profile in (C), which shows layered ash with flecks of charcoal overlying rubified sediment. Distinct root and termite disturbances are visible crossing the boundaries of the intact ash layers. The brownish (grey when dry) sediment above and below is also ash-rich but more mixed (homogenised) and darker due to a higher content of finely comminuted organic material and microcharcoal. The sediments were dampened with a mist of water prior to taking the photos and image levels were adjusted to emphasise colour differences. Jessica Thompson in Cerezo-Román et al. (2026).

These pyre deposits comprise a series of laminated ash layers extending horizontally for more than a metre, and containing ash, charcoal, rubified sediments, and Human remains. These deposits have been penetrated by insect burrows and/or plant root tunnels in places, presumably before the ash layers became cemented. Examination of the rubified sediments suggests that these contain fragments of baked clay which probably originated from structures built on trees by termites, suggesting that deadwood was collected to build the pyre(s). Also present were carbonised fragments of the Bracket Fungus Ganoderma.

Thin section of sample 834. Sample 834 was collected from the pyre, about 1m away from the Hora 3 remains (A). One thin section was selected from the small block sample (B) with reddish disturbed sediments in the lower part, and bright ashes in the top (C). Termite galleries and shelter tubes are very common in woodland areas, especially on deadwood, which presents ready-for-use fuel and was likely preferentially selected to build the pyre (D); (E) displays the clear boundary between compact laminated ashes (lower half of photomicrograph) and mixed ashy sediment on top, which are mixed with sandy clay soil aggregates. A very coarse sand fragment (feldspar) in the lower right likely represents roof spall; (F) A closer view of wavy laminated ash topped by sandy ash; Wavy layers of intact ashes (grey) and elongate, sickle-shaped clay aggregates (orange, red), possibly resulting from termite shelter tubes on tree branches (G); The clear lower contact of the compacted ash is characterised by a sudden increase of sand in the underlying ashy sediment (H); Phosphatic ashes indicate some mineral alterations and weathering of the ashes (I); Termite channels, fortified by well-sorted sandy clay, run through the lower part of the sample and contain burned organic matter (J); Articulated ash fragment in compacted ash (K); The sediment consists of a mix of fire residues, clay aggregates, and sand (L). Flora Schilt in Cerezo-Román et al. (2026).

The ash deposits generally contain fewer artefacts (such as flakes from toolmaking) than non-ash deposits, probably relating to the fact that these were laid down much more rapidly. However, the Human-remains-containing core layer contains many more such objects, suggesting that they were deliberately included within the pyre, possibly for symbolic reasons.

Unifacial points and cores from square E11-a core pyre ash contexts. (B) and (F) Unmodified points; (D), (E), (G), (H), (I), and (J) broken point fragments (dashed red line shows breaks); (A), (C), (E), and (D) point showing unifacial retouch; (H) point fragment showing basal tang. Red stars indicate point areas with possible residue preservation. Justin Pargeter in Cerezo-Román et al. (2026).

A total of 170 Human bone fragments were recovered from the pyre area. Of these, 112 could be clearly identified as specific parts of the Human skeleton, with the remaining 18 identified as 'likely human' on the basis of their size, texture, and preservation. The majority of these bone fragments (160) were recovered within the ash feature, with 152 found within the core feature. Of the elements which could definitely be identified as Human, 97 could be assigned to a specific element, with 90 of these being long bone (i.e. arm or leg) fragments. Other fragments from the core pyre include partial left and right calcanei, an unsided patella, the lower right articular facet of a lumbar vertebra, and four phalanges. All are consistent with a single individual.

Skeletal inventory and thermal alteration colour changes. Preserved skeletal elements and changes in bone coloration in relation to temperature and fire exposure. Additional fragments not shown in the figure: an unsided patella, the lower right articular facet of a lumbar vertebra, and unidentified long bone fragments. Jessica Cerezo-Román in Cerezo-Román et al. (2026).

Analysis of the Hora 3 skeleton suggests that this individual was a female aged between 18 and 60. The individual is presumed to be an adult on the basis that it appears to be skeletally mature, with the distal femur and radius having fused. The articular margin of the distal femoral epiphysis showed marginal lipping, which may indicate the presence of a mild degenerative disease, giving the upper age estimate. No other potential age-markers were preserved. Since it was impossible to recover DNA from the burned and fragmented remains, sex was estimated using overall small size and gracility and the vertical diameter of both preserved femoral heads. On the basis of the diameter of the femora, the individual is estimated to have been between 145 and 150 cm tall in life (about 5 feet). Based upon the maximum and minimum midshaft diameters of the right humerus and femur, Hora 3 it thought to have been less mobile than most modern Southern African hunter gatherers, and more accustomed to manual labour.

(Top) Right distal humerus with cemented/encrusted termite tubes. (Bottom) Cemented elements that include the left proximal femur, a radius shaft, a tibia shaft, and an unidentifiable long-bone shaft. Cerezo-Román et al. (2026).

About 585 g of material thought to be derived from the skeleton of Hora 3 was recovered. This is notably less than is typically produced by a Human cremation, which is usually more than 1500 g. Notably, no head elements were found within the core pyre area, although a fragment of a sphenoid (the base of the skull) was found within the larger ash feature. Fragments of cranium, and in particular teeth, are usually among the most easily preserved and identified elements in cremations, and it is surprising not to have found any such fragments, given that sediment and ash removed from the site were wet sieved to 1 mm.

All of the skeletal remains show transverse, curved-transverse, and longitudinal cracks, which are typical for the burning of a fleshed corpse, but none show signs of the checking pattern fractures which are often seen when dry remains are burned. Most fragments were grey to black, with some brown, bluish-white, and white hues. Fragments from closest to the torso were typically the most blackened and charred, while more distal elements were grey or white - which indicates more exposure to flame in areas which would have been covered by less flesh. Many elements are white on their external surface, while internal surfaces are grey, which implies these elements were intact when the burning occurred. These colours suggest that the temperature to which the distal elements were exposed exceeded 500°C, while elements closer to the torso typically reached maximum temperatures lower than 500°C. Fourier-transform infrared spectroscopy is sensitive to low temperature heating of bone, which often does not involve recrystallisation, and therefore can provide an accurate palaeothermometer for low temperature cremations. Four white/grey fragments from Hora 3 were analysed in this way, with all producing spectral peaks indicative of heating to over 500°C.

Although the body of Hora 3 shows signs of having been manipulated after the cremation, many of the recovered fragments were recovered together, covered in ash, and in some cases cemented together. This includes a right distal humerus and right proximal radius cemented together along with long bone fragments and a possible metacarpal fragment. This appears to show that intact joints were present at the time of final disposal of the remains. Many of the bone surfaces are covered by calcium carbonate, with none of the recovered Human bones showing more than 50% exposure. Nevertheless, eight of the bones showed stone tool marks consistent with defleshing, while none showed any signs of having been scavenged by carnivores. 

Bone modifications made with stone tools. Bone surface modifications made with stone tools. (A), (B), (C), (D), and (G) are cut marks. (E) is a percussion mark. (F) is an indeterminate mark made with a sharp object. (B) to (G) show the variable bone colours and superficial calcination with carbonisation at the interior. (C) shows the minor 'skin' of calcium carbonate overlying the marks. (G) shows remaining adhering ash. (B) to (G) show two aspects of the fragment, with close-ups of the modifications in the boxed area. Jessica Thompson in Cerezo-Román et al. (2026).

Cerezo-Román et al. reconstruct a sequence of events in which people began using the HOR-1 as a habitation site about 21 000 years ago. The earliest recorded burials at the site occurred 16 000 and 14 000 years ago, both being infants. Small ash features consistent with campfires appear around the end of the Pleistocene, with larger pyres appearing by 10 240 years before the present.

Geology of HOR-1 rock shelter.(A) Pinkish and gray walls of the rock shelter at the site location. (B) Exfoliating bedrock higher above the excavation trench. (C) Example of spheroidal exfoliation of the bedrock near the rock shelter. Flora Schilt in Cerezo-Román et al. (2026).

Some time between 9540 and 9454, a pyre comprising at least 30 kg of deadwood and grass, with some hairy leaves from herbaceous plants, was assembled, something which would have represented a significant investment of time and effort by the local community. The body of a small, probably female, adult individual was then burned on this pyre, probably within a few days of their death. The distribution of the bones, particularly those of the arms and legs, suggests that this individual was placed on this pyre in a flexed or pugilistic position (i.e. with the knees and elbows brought in close to the torso). Cut marks on some of the bones show signs of defleshing, although the preservation of the bones and some joints suggests that the flesh of the body (or some other covering) was present at the time of burning. It is likely that the skull was removed before the burning, since neither scavengers nor Early Holocene hunter gatherers seem likely to have been able to remove all of the fragments which would have been produced by burning a skull, while leaving other elements of the skeleton in situ. The ritual removal of skulls, and other body parts, has been documented in a range of modern African populations, but never at a site of such antiquity. 

Spatial locations of femora and right tibia fragments. Cerezo-Román et al. (2026).

The fire achieved was able to reach temperatures above 500°C, although this doesn't appear to have been consistent, and the lamination of the deposits suggests that the fire-makers continued to add fuel to the pyre for some time. The detachment of Cluster 2 from Cluster 1, and the greater degree of burning seen on Cluster 1, may indicate that the body was manipulated during the cremation process to detach parts of it. Knapped material appears to have been added to the pyre, either at the outset or during the process, possibly in association with other funerary objects. Multiple further fires were lit at the same site over the next few hundred years, although no further cremations appear to have occurred.

Reconstruction of the cremation ritual. Sequence of events leading to the formation of the cremation feature at HOR-1. (A) Site location at an inselberg, a natural monument. (B) A large quantity of wood was collected to construct the pyre, suggesting communal labor. (C) Cutmarks on bone show parts of the body were defleshed. (D) Human remains display black coloration and curved transverse fractures, indicating some moisture in the remains. (E) The pyre and body at Cluster 1 were actively disturbed during burning, creating Cluster 2. (F) High temperatures were maintained by attendees adding additional fuel. (G) Convergent points are uniquely associated with the cremation. (H) Bipolar reduction dominates a lithic assemblage that occurs in higher concentrations with the remains than in the rest of the ash feature. (I) The presence of carbonised Ganoderma and the remnants of termite tunnels indicates the use of deadwood as the primary fuel. (J) The absence of cranial and dental remains suggests these may have been collected and removed. (K) Multiple fires were relit atop the original pyre location within communal memory. Patrick Fahey in Cerezo-Román et al. (2026).

Evidence of cremation among African hunter-gatherer populations is extremely rare in the archaeological record, and has not previously been found south of the Sahara. The oldest previously documented example come from the Nabta Playa site in southern Egypt, where a single individual appears to have been burned between 7800 and 7300 years ago (i.e. at least 1300 years before the earliest date for the Nabta Playa stone circle), and this has been interpreted as a 'burned inhumation' rather than a true cremation, possibly caused by the accidental ignition of material placed within the grave. Burned Human remains, possibly as much as 7000 years old, have also been recovered from an ancient midden near Lake Besaka in Ethiopia, although again this does not represent an in situ pyre like that seen at Hora, and again may not represent an intentional burning. The previous oldest known intentional cremations from Africa are associated with Neolithic Elmenteitan pastoralists in Kenya, about 3300 years ago.

A open-air, pyre cremation such as that seen at Hora requires a significant investment in time and labour by the local population, which may be why the practice is so rare among hunter-gatherer populations. In an enclosed furnace, a Human corpse can be burned in about two hours, but open air cremations require considerably longer, during which time the temperature must be maintained, typically by adding more fuel. Nevertheless, such a cremation would be a notable community event, particularly if, as seems to have been the case at Hora, parts of the body were removed during the process for ritual disposal elsewhere.

While the Hora site only records a single event, which seems to have been highly unusual in nature, it adds to a growing picture of a culturally diverse tropical African hunter-gatherer population in the Early Holocene, displacing an earlier impression that these peoples were likely to have been culturally homogeneous, even over great distances and long periods of time. The people at Hora did not erect megaliths or other architectural features. Nevertheless, they appear to have utilised chosen a site of natural prominence to carry out a significant mortuary ritual, a site which appears to have been utilised repeatedly over an extended period of time (at least 16 000 years), albeit with changing rituals during that time.

Evidence for evolving social cooperation and complexity has been recorded from many ancient hunter-gatherer populations around the world, although until now this has largely been absent from Africa. The Hora Rockshelter in Northern Malawi records a population showing both a diversity of behaviour and the ability to stage large events which would have required the investment of time and effort by many individuals. This appears to have been carried over multiple generations, with repeated fires being lit, including at least one subsequent major pyre event, suggesting a communal memory in which the significance of the location was maintained over many generations.

Cerezo-Román et al. conclude that around 9500 years ago the remains of an adult female were burned on a substantial pyre at the Hora Rocksheter in Northern Malawi. This is the oldest known pyre cremation in Africa, and the oldest known adult pyre cremation in the world. The fire appears to have been one of a sequence of fires at the same location, which persisted for centuries after the cremation event, signifying the importance of the site to the population. This site demonstrates the emergence of complex funerary rights, communal projects involving large numbers of people, and the recognition of certain sites as culturally significant long before the emergence of agriculture and organised food production, challenging previous conceptions about the extent to which community co-operation occurred among ancient hunter-gatherer populations in tropical Africa.

See also...

Genetic analysis of individuals from the Oakhurst Rockshelter suggest 9000 years of genetic isolation in South Africa.

Modern Southern African populations contain genetic diversity which records the deepest branching events known in the genetic history of extant Humans. The region also has a long archaeological record, with archaic Homo sapiens first appearing here around 260 000 years ago and Anatomically Modern Humans around 120 000 years ago. Because of this, the ancient Human populations of Southern Africa have been the subject of numerous archaeological and palaeogenomic studies, by scientists hoping to gain insights into population structures during the later stages of Human evolution. However, this interest in the most ancient Human genomes recoverable has left somewhat of a gap in the study of more recent, Holocene populations.

The Holocene has seen significant changes in technology and culture within Southern Africa. During the last 2000 year new populations have migrated into the area, bringing with them pastoralism and crop-farming. This began with the arrival of herders from East Africa, and was followed by farming populations from West Africa, who also brought the Bantu language group to the region. As well as  setting up  new communities with new ways of living, both of these groups contributed to the genetic structure of the original populations, so that all  extant San and Khoe populations draw at least 9% of their genetic material from outside Southern Africa. 

In a paper published in the journal Nature Ecology & Evolution on 19 September 2024, Joscha Gretzinger of the Department of Archaeogenetics at the Max Planck Institute for Evolutionary Anthropology, Victoria Gibbon of the Division of Clinical Anatomy and Biological Anthropology at the University of Cape Town, Sandra Penske also of the Department of Archaeogenetics at the Max Planck Institute for Evolutionary Anthropology, Judith Sealy of the Department of Archaeology at the University of Cape Town, Adam Rohrlach, also of the Department of Archaeogenetics at the Max Planck Institute for Evolutionary Anthropology, and School of Computer and Mathematical Sciences at the University of Adelaide, Domingo Salazar-García of the Department of Geological Sciences at the University of Cape Town, and the Departament de Prehistòria, Arqueologia i Història Antiga at the Universitat de València, and Johannes Krause and Stephan Schiffels, again of the Department of Archaeogenetics at the Max Planck Institute for Evolutionary Anthropology, present the results of a study in which they obtained genetic samples from a series of individuals from the Oakhurst rockshelter in South Africa, and compared these to other genetic samples from historic and living populations in South Africa.

The Oakhurst Rock Shelter is located 7 km from the southern coast of South Africa, close to the town of George in Western Cape Province. It was first excavated in the 1930s, and has yielded a remarkable sequence of archaeological remains, now known to represent about 12 000 years of accumulation. The Early Holocene layers here have yielded an assemblage of macrolithic tools which has been named the 'Oakhurst Complex' in reference to the site, which has been discovered at many sites across South Africa. Around 8000 years ago, this Oakhurst Assemblage was replaced by a set of microlithic tools, which have been named the Wilton Assemblage, which persisted throughout the remainder of the Middle and Late Holocene, with minor variations. Around 2000 years ago, ceramics also begin to appear at the site. 

As well as the numerous cultural artefacts, the Oakhurst Rock Shelter has also yielded 46 sets of Human remains, adult and juvenile, deposited throughout the archaeological sequence, including the oldest dated set of Human remains to have yielded DNA in South Africa, which are 10 000 years old. Gretzinger et al. obtained genetic material from 13 individuals from the Oakhurst Rock Shelter, all of which have been radiocarbon dated from their bone collagen, yielding ages of between 10 000 and 1300 years; nine of these dates are from previous studies, while four are new dates obtained by Gretzinger et al.. The generic sex was determined for all thirteen individuals, with the mitochondrial haplogroup obtained for nine individuals and the Y chromosome haplogroup for five.

Because mitochondrial DNA is found in the mitochondria, organelles outside the cell nucleus, it is passed directly from mother to child without being sexually recombined each generation, enabling precise estimations of when individuals shared common ancestors, at least through the female line. It is also possible to trace direct ancestry through the male line, using DNA from the Y chromosome, which is passed directly from father to son without sexual recombination.

Gretzinger et al. next created a haplotype population tree including ancient DNA from the nine Oakhurst individuals with mitochondrial DNA haplotypes, as well as samples from other archaeological sites in Africa, and modern populations. Most of the samples used were from previous studies, and are publicly available, however, some of the sequences were obtained from San skeletal material held by the University of Cape Town, and used only with permission of the San communities from which they were obtained. Access to this data is only available to other researchers with the permission of the University of Cape Town Skeletal Repository Committee and the relevant San communities. 

This recovered the Oakhurst individuals as being on the deepest branching limb of the living Human tree, which also includes living San populations, but closest to other ancient individuals from South Africa than to any living population. They also note that they recovered an ancient divide between San populations living north and south of the Kalahari, and that all the ancient South African populations, including the Oakhurst individuals, are on the same branch as the San populations from south of the Kalahari.

Maximum likelihood tree showing genetic affinities between ancient and present-day southern Africans, generated using TreeMix of genome sequences from present-day and ancient populations, excluding populations with evidence of asymmetrical allele sharing with non-Africans indicative of recent gene flow. Branches of ancient individuals/groups are truncated for better readability. Gretzinger et al. (2024).

Looking at the wider genomes, Gretzinger et al. found that San and Khoekhoe populations split into three principle groups, with the Kx`a-speaking Ju|’Hoan and !Xuun forming a northern cluster, Khoe-Kwadi-speaking Nama, and Tuu-speaking ‡Khomani and Karretjiemense forming a southern cluster (Karretjiemense is an Afrikaans word meaning 'people of the cart', but is how these people self-identify), while the Tuu-speaking Taa, Kx`a-speaking ǂHoan, and Khoe-Kwadi-speaking Gǀui and Gǁana form a central group. Eight of the Oakhurst individuals lie within the southern cluster, as do four other Later Stone Age skeletons with published genomes from South Africa, although the oldest individual in the dataset show a slightly greater affinity for the northern cluster. Notably, within the southern cluster, the Oakhurst individuals showed the greatest affinity to populations still living close to the area today, with the youngest individual, OAK007, dated to 1344 years before the present, showing the greatest affinity for living populations, sharing more and longer identical by descent segments with the Karretjiemense and ‡Khomani than with any other tested population.

Comparison of the genomes of the Oakhurst individuals to other, previously published, ancient African genomes, Gretzinger et al. found that all South African Later Stone Age genomes were closer to one-another that to those of any other ancient African. The youngest individual, OAK007, was most closely related to two other Later Stone Age individuals, from St. Helena and Faraoskop, both of which have been dated to about 2000 years before the present. Together, these three individuals form a sister group to two further individuals from Ballito Bay on the eastern coast of KwaZulu-Natal, thought to be of similar age. Older genomes from Oakhurst become steadily less closely related to these individuals as they get older, as well as less closely related to the genomes of historical San samples from Sutherland, Western Cape Province. However, the genome of a 1200-year-old pastoralist from South Africa clustered with Later Stone Age genomes from Malawi, while those of four Iron Age farmers from South Africa clustered most closely with Later Stone Age genomes from Cameroon. 

Next Gretzinger et al. looked for potential ingression of non-San genetic sequences into the Oakhurst individuals, finding no trace of affinity to populations in either East of West Africa, and a consistent grouping with southern rather than northern San groups, the genetic gulf between which groups appears to have been widening steadily since their split around 20 000 years ago, before the drying of the Lake Makgadikgadi palaeo-wetland, which once covered most of central Botswana.

All of the Oakhurst individuals dating to between 10 000 and 1344 years before the present, form part of a single clade (group with shared common ancestry), which also includes individuals from St. Helena, Faraoskop, and Ballito Bay dating to between 2200 and 1300 years before the present. However, the genomes of individuals from South Africa from between 1300 and 1200 years before the present show a significant discontinuity with earlier individuals, with a second discontinuity observed between 1200 and 400 years before the present. Gretzinger et al. attribute these discontinuities to the influxes of first pastoralists from East Africa and then farmers from West Africa into the region. However, they find no trace of West African ancestry in three individuals from Sutherland dating to the late nineteenth century, while about 11% of their genome appears to be of East African ancestry, a proportion similar to that seen in living ‡Khomani individuals from the Northern Cape Province, who typically have genomes comprising about 9% East African genetic material.

Based upon this, Gretzinger et al. observe that no evidence of any genetic influx from outside of modern South Africa recorded at Oakhurst Rock Shelter between 10 000 and 13 000 years before the present, a remarkable period of genetic continuity lasting almost 9000 years. Despite this, the Oakhurst individuals show no signs of being genetically isolated, The level of conditional nucleotide diversity (the  extent to which each member of a pair of chromosomes differs from its partner, used as a measure of inbreeding within a population( maintained within the Oakhurst samples is lower than that found in  Later Stone Age population from Malawi, Kenya and Cameroon, but comparable to other Later Stone Age populations from Western Cape and KwaZulu-Natal, and greater than is seen in ancient hunter gatherer populations from Serbia, Japan, and Brazil, as well as modern San and Khoe populations. This is non consistent with a model of long-term isolation, instead indicating to the presence of a much larger population of Later Stone Age hunters in South Africa before about 1300 years before the present, when other groups are generally accepted to have begun to arrive in the region, and a subsequent dramatic reduction in the size of that population.

Reconstructing the demographic history of South Africa over the past 2000 years is complicated, with at least two significant prehistoric population influxes, and substantial genetic exchange with both other parts of Africa and other continents following the establishment of the first European settlements in about 1650. To try to address this, Gretzinger et al. created a model using genomes from Later Stone Age hunter-gatherers in South Africa, the Luxmanda archaeological site in Tanzania, which has been dated to about 3000 years before the present, and modern Mende populations from West Africa.

They then developed a best-fit model which enabled them to group populations into primarily West Africa or Primarily East African (excluding populations with a substantial amount of genetic material from both sources), in order to estimate dates for the admixtures of the West and East African components. They found that San and Khoe populations began to absorb genes from East Africa substantially before those from West Africa, with an estimated date of 1068 years before the present. This is consistent with the East African ancestry recovered the 1200-year-old pastoralist remains from Kasteelberg, on the southwest coast of South Africa near St. Helena Bay, and the estimated date of admixture of 1228 years before the present recovered from the nineteenth century Sutherland material. 

The arrival of West African genes in South Africa appears to have been considerably more recent, with living Bantu-speaking groups such as the Herero, Tswana, and Kgalagadi, producing an estimated admixture date around 808 years before the present, while 400-year-old remains attributed to Iron Age farmers from KwaZulu-Natal yielded an estimated admixture date around 832 years before the present. Living San and Khoe groups yielded a more recent estimated admixture date, of about 578 years before present. Gretzinger et al. suggest that this may reflect either several waves of West African arrivals, or a continuous flow, with an initial admixture of San and Khoe genetic material into the ancestors of modern Bantu-speaking groups and a subsequent flow of West African genes into the ancestors of modern San and Khoe populations.

All groups show considerably more Later Stone Age ancestry on their X chromosomes than on their autosomal (non-sex determining) chromosomes, with this signal being stronger in San and Khoe populations than the Bantu-speaking groups. This implies that in most cases, the contribution from Later Stone Age hunter-gatherers was from the female side. The extent to which this is true appeared to vary between living populations, with the living Damara (a Khoekhoe-speaking people from northwestern Namibia) having had about 1.4 female Later Stone Age hunter-gatherers in their ancestry for each male, the ǂHoan (a Kxʼa language-speaking group from Botswana) having about 2.28 Later Stone Age hunter gatherer females per male in their ancestry, the Shua (a Khoe-speaking group from central Botswana) having about 4 Later Stone Age hunter gatherer females per male in their ancestry, the Haiǁom (a Khoekhoe speaking group from Namibia) having about 5.2 Later Stone Age hunter gatherer females per male in their ancestry. This also applies to South Africa Bantu-speaking groups (for whom the overall contribution of Later Stone Age hunter gatherer genetic material is lower), with about 2.1 females per male having contributed genetic material to the extant population. 

This female bias can also be seen in the historical Sutherland genomes and the 1200-year-old pastoralist remains from Kasteelberg, although, surprisingly, not to the four Iron Age KwaZulu-Natal individuals, who have a higher proportion of Later Stone Age hunter gatherer genetic material on their autosomal chromosomes than on their X chromosomes, indicating a higher proportion of male Later Stone Age hunter gatherer ancestors than female ones. This is different to the situation seen in all other groups in South Africa and Botswana for which a trend could be determined, and may reflect a change in the way different groups were integrating in the past 400 years compared to the nature of such interactions during the arrival of the first farmers into the region.

Gretzinger et al. finally note a recent admixture of male northwest European DNA into San/Khoe and mixed groups from Colesberg and Wellington. The estimated date for these ingressions is 199 years before the present, despite the known arrival of Dutch and British migrants into the region from the mid-1600s onwards, something which led to a collapse in San and Khoe genetic, linguistic and cultural diversity. In addition to severely disrupting existing population structures, the European arrivals introduced a range of new populations into the region, all of which have contributed to modern population structures to some extent. As an example, Gretzinger et al. note that mixed-ancestry South Africans from Colesberg drew an average of 24.4% of their ancestry from South Asia, 2.8% from East Asia, 8.2% from Northern Europe, and about 35.5% from Later Stone Age hunter gatherers. Some San and Khoe groups also have a significant proportion of European ancestry, with the Karretjiemense drawing an average of 5.61% of their ancestry from Europe, the ‡Khomani on average 9.45%, and the Nama on average 6.83%. This suggests that southern San populations were particularly affected by intermixture with Europeans, with these groups having a higher proportion of European ancestry that other San or Khoe populations, comparable to that of the mixed-ancestry South Africans sampled. Thus, the modern populations most closely related to the Oakhurst individuals appear to be particularly affected by genetic ingression from other populations.

Demographic changes in the San and Khoe populations of southern Africa: Summary of the inferred population history of the San and Khoe in southern Africa. Sex symbols indicate male- and female-biased reproduction. Note that pastoralism and farming both appeared in present-day South Africa at about the same time, 2,000 years ago. Gretzinger et al. (2024).

The question of population continuity within Later Stone Age communities in Southern Africa has engaged archaeologists for over a century. In the past two decades, the application of genetic methodology to archaeological problems has helped to unravel the demographic histories of Stone Age populations in Europe, Asia, and North Africa, revealing episodes of large-scale migration in these regions, during which indigenous populations were either replaced by or absorbed into the new population. These biological replacements of populations also appear to have been vectors for the spread of new technologies. In South Africa, in contrast, there appears to have been a surprisingly long period of genetic continuity, with on detected influx of genetic material from elsewhere for at least 9000 years, from the beninning of the Holocene till around 1200 years ago, during which time the Southern San remained isolated from Northern and Central San populations as much as from other populations elsewhere in Africa.

This implies that the cultural changes seen at the Oakhurst Rockshelter, such as the transition from the Oakhurst to the Wilton technocomplex, were a result of entirely local inovation. It has previously been observed that there have been slight fluctuations in craniofacial size in Later Stone Age populations in coastal South Africa, something which has been interpreted as a sign of genetic discontinuity, something which Gretzinger et al.'s results contradict. Since the population was not a small isolated one which might be subject to strong effects from genetic drift, it seems likely that these variations were driven by changes in the local environment.

The 9000 years of genetic and cultural isolation experienced by Later Stone Age hunter gatherers in South Africa seems to have ended quite abruptly, with the spread of herding communities from East Africa shortly followed by the arrival of farming communities from West Africa. Most parts of South Africa do not record any genetic trace of these arrivals before about 1300 years ago. However, there is evidence for changes in settlement patterns and other cultural behaviours in this coastal South Africa from about 2000 years ago, which have been interpreted as a response to the arrival of herding in the area. In Europe, a similar cultural shift is seen at the Neolithic-Mesolithic boundary, with a genetic admixture between the incoming farming population and the extant hunter gatherer population not being recorded for about 2000 years after the cultural shift. This implies that in Europe at least, farming and hunter gatherer populations were able to live alongside one-another for a long period of time before beginning to intermix, something which may also have been true in South Africa. Alternatively, pastoralism may have been culturally transmitted from East Africa to Southern Africa long before the spread of East African populations into the region.

However, from about 1200 years ago onwards, there has been substantial migration into Southern Africa from other regions, and substantial ingression of new genetic material into all populations, with the effect that all living San and Khoe populations are admixed with one or both of East African Pastoralist and West African Farmer ancestry. This process was accelerated by the arrival of European settlers in the mid-seventeenth century, which led to widespread population collapse among hunter gatherer populations in Southern Africa. Combined with a loss of oral traditions, these events have greatly obscured the prehistoric population structure of southern Africa. 

Genetic methods such as those used by Gretzinger et al. provide a way to study these ancient population structures, showing that the San and Khoe inhabitants of South Africa are the direct decendants of the Early Holocene inhabitants of the region, despite considerable disruption to their lifestyle and population structure by later migrants to the region.

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Evidence for Late Pleistocene net-hunting in southern Somalia.

Humans have been farming for about 12 000 years, a period which was preceded by hundreds of thousands of years of living a hunter/gatherer existence. It is logical to assume, therefore, that the social networking skills which made modern civilization possible developed largely before the event of farming. The advent of specialist hunting techniques such as net-hunting, which require advanced planning and co-operation between groups of hunters, and which can produce large amounts of meat in single events, are thought to have been a significant step in the development of Human social networks, connected to the emergence of social hierarchies and concepts of land ownership.

The lifestyle of Homo sapiens in East Africa from our emergence as a species onwards has been heavily investigated for decades, although curiously, despite the importance placed on the role of hunting as a driver of Human evolution by many early archaeologists, the techniques used to hunt have been relatively neglected, with an assumption that big game hunting was the dominant behaviour. Recent studies, however, have suggested that while big game hunting was important on the grassy plains of East Africa, in the coastal forests of the region, remote capture devices, such as snares, traps, or nets, used to target small Animals have been important for at least 125 000 years.

Given that such methods are thought to be closely associated with the emergence of social networks, the nature these hunting practices, and the ways in which they evolved over time, must be seen as an important part of the Human story. Examination of Later Stone Age sites in the semi-arid environment of southern Somalia, have shown that nets were being used to intensively hunt Dwarf Antelopes between about 26 000 and about 6000 years ago.

In a paper published in the journal Archaeological and Anthropological Sciences on 2 December 2023, Mica Jones of the School of Archaeology at the University of Oxford, presents the results of a study of Animal bones from the Late Pleistocene Guli Waabayo rock shelter in the Buur Heybe inselberg cluster, and the implications of this for the use of net-hunting by the Later Stone Age hunter/gatherers of the area.

Inselbergs are hills made from hard volcanic rock, which persists when other deposits are eroded away, typically forming elongate structures. In Somalia such structures are known as 'buurs', and in the inter-riverine region of southern Somalia, where rainfall is typically between 400 mm and 600 mm per year, clusters of these buurs can help to trap nutrient-rich soils and collect rainwater in pools and ponds, providing favourable habitats for a range of Plants and small Animals.

A previous study of the Rife Range Site at Buur Hakaba found that Later Stone Age hunters there specialised in hunting Dwarf Antelope, probably with nets, leading to the development of a more sedentary lifestyle and the holding of territories in the Early-to-Middle Holocene, a period of increased rainfall. However, the small number of remains recovered from this site, combined with an absence of dates, limits the usefulness of this study. The Guli Waabayo rock shelter at Buur Heybe, about 25 km to the northeast of Buur Hakaba, has yielded a much more extensive collection of Animal remains, combined with a series of robust dates, covering a period of 20 000 years, potentially enabling a good test site for hunting practices in the region before the start of the Holocene.

Map of inter-riverine southern Somalia with locations of the Buur Heybe and Buur Hakaba inselberg clusters. Jones (2023).

More than a hundred rock shelters and other archaeological sites have been identified in the inter-riverine region of southern Somalia since the 1930s. However, only three of these sites have been subject to any organised excavation work. The Gogoshiis Qabe site was excavated in the 1940s by the Italian archaeologist Paolo Graziosi. In the 1950s British archaeologist John Desmond Clark investigated the Guli Waabayo and Rife Range sites, establishing that there was a degree of cultural continuity between the two. Following Clark's activities, no further organised archaeological work was carried out in the region until the 1980s, when Steven Brandt and the Buur Ecological and Archaeological Project returned and carried out further excavations at all three sites. These excavations were brought to an abrupt halt with the onset of the Somali Civil War in 1989, although, with the consent of the Somali Academy of the Arts and Sciences, much of the excavated material was exported to the US, where it has been held in the collections of several institutions. 

Jones examined the material brought back from Guli Waabayo to establish the identity of the small Antelope bones present; these had previously all been assigned to the genus Madoqua (Dik Diks), but this is not the only genus of small Antelopes in the Horn of Africa, and, since different Antelope have different ecologies, the identity of these bones has implications for the techniques which would have been effective when hunting them. Next Jones looked at the overall assemblage, and calculated the proportion of remains coming from each type of small Mammal, and then the proportion of each age-set within the overall sample.

Studying hunting methods used to on large prey can be achieved by looking at the weapons involved, or the use and modification of the landscape by the hunters, such as cliff jumps or kite structures. However, the study of how small game is hunted requires less direct approach, as such Animals are often captured with snares, nets, or traps made from perishable materials such as plant fibres, wood, or leather. In order to address this Zooarchaeologists have developed methods of analysing the ways in which people in the past hunted small game, based upon analysis of the variety of remains.

The composition of collections of small Animal remains will vary depending on the hunting method used. Studies conducted in places where net-drives (driving Animals into net traps) are still used, such as the Congo Basin, Australia, and the southwestern United States, has shown that this method will tend to target only one or two species, but will trap almost all members of those species within the target area. This method is most useful for targeting small Animals in forested, bushy or rocky habitats. Pursuit hunting and setting of individual traps such as snares both tend to produce a more diverse range of prey, which will be reflected in the bone assemblage left behind. 

Furthermore, while net hunting captures individuals of all ages, both snare hunting and persuit hunting tend to produce a high proportion of remains from a single age group, since the young of small Animals generally lack the weight to trigger snares designed for adult Animals, and Humans hunting with bows or clubs will generally take a higher proportion of slower, juvenile Animals.

The excavations carried out at Guli Waabayo comprised 11 one metre by one metre pits, each excavated to a depth of 2.5 m. These uncovered a sequence of three Middle Stone Age/Later Stone Age technologies, although, the lower part of the exposed sequence could not be dated due to an absence of Animal remains. Dates were obtained from Ostrich shell fragments for the upper 1.5 m of the sequence, demonstrating at least sporadic occupation of the shelter from 26 000 to 6000 years ago. Two distinct Later Stone Age technologies could be seen within the sequence, the Eibian, which was used during the arid Marine Isotope Stage 2 from about 29 000 to about 14 500 years ago, and the Bardaale, used in the wetter African Humid Period from about 14 500 to about 6000 years ago.

Analysis of the faunal remains found at Guli Waabayo showed that Later Stone Age hunters across the Pleistocene/Holocene boundary targeted a diverse range of game, including large and small Mammals, Birds, Reptiles, and Fish, although they had a clear preference for Mammals massing less than 20 kg. This preference for small Mammals seems to have increased slightly in the African Humid Period, although this change in climate appears to have made relatively little difference to the site's occupants, probably because the African Humid Period was a lot less pronounced in southern Somalia than in other areas.

Of the bones present at Guli Waabayo, 3104 could be identified as coming from small Mammals, with 2111 identifiable to a lower taxonomic level. Of these 1263 have previously been identified as coming from Dwarf Antelope, with the assumption that they originated from Dik Dik. However, this assumption was never actually put to the test, and there is another form of Dwarf Antelope present in the Horn of Africa, the Suni, Neotragus moschatus.

Jones selected all of the first phalanges assigned to Dwarf Antelopes in the Guli Waabayo collection, and compared them to first phalanges from Dik Dik and Suni in the collection of the Field Museum in Chicago. The size ranges of the first phalanges did not overlap, and all of the specimens from the Guli Waabayo collection conformed with the Dik Dik form, although it was not possible to tell which species of Dik Dik they belonged to.

The Dik Dik genus, Madoqua, first appears in the fossil record in the Miocene. There are four living species within the genus, Madoqua guentheri, Madoqua saltiana, and Madoqua piacentinii are endemic to the Horn of Africa, while the fourth, Madoqua kirkii is found from Somalia south to Tanzania and eastern Uganda, as well as on the west coast of Southern Africa in Namibia and Angola. All species favour rocky environments with low thicket vegetation, with the inselbergs of southern Somalia providing an excellent environment, that today supports a large population of Dik Dik, which live in monogamous pairs, with each pair occupying and defending a territory with an area of about 1 km².

A Salt's Dik Dik, Madoqua saltiana, in the wild. David Castor/Wikimedia Commons.

Dik Dik are not widely hunted in Africa today, although in places they are targeted for their meat and skins, which are used to make gloves and cloaks. The Hadza people of northern Tanzania occasionally hunt Dik Dik with bows and arrows, while the Mukogodo people, who lived in the area around Mount Kenya in the eighteenth century are believed to have hunted Dik Dik both with snares and by stalking. 

Modern Somali pastoralists have been recorded to chase down Dik Dik, which seems surprising, given that these Dwarf Antelopes can reach speeds of about 42 km per hour, but is possible because they are so reluctant to leave their territories that they will often run around the perimeter when chased, allowing much slower Human pursuers to keep them in sight until they become exhausted and are easily captured. In the 1940s and 1950s several agropastoralist groups in Somalia were reported to capture Dik Dik in net hunts.

During these hunts, the men used nets about 5 m in length, strung between 2 m long poles. These nets would be used to create semi-circular traps, into which Dik Dik could be driven by men and Dogs, before being dispatched with bows and arrows, a methodology which could conceivably have been used by Later Stone Age hunters in the same region.

Having established the Small Antelope remains at Guli Waabayo came from Dik Dik, Jones next sorted the remains into age groups, based upon examination of complete and partial tooth rows, as well as loose lower 3rd molars and 4th premolars. The dental aging of modern Dik Dik has not been extensively studied, but it was still possible to separate the remains into juvenile, older juvenile, adult, and older adults, based upon the relative eruption and wear seen in the teeth. 

Madoqua mandibles of different age sets from Guli Waabayo. Jones (2023).

Dik Dik remains dominated the faunal assemblage at Guli Waabayo in both Marine Isotope Stage 2 and the African Humid Period. Roughly twice as many remains assignable to Madoqua were found in Marine Isotope Stage 2 selection as in the African Humid Period selection, but this was also true of other Animals present, ab probably reflects the fact that Marine Isotope Stage 2 lasted about 14 000 years, while the African Humid Period lasted only about 6000 years. Nevertheless, the proportion of the assemblage made up by Dik Dik bones did increase, from 55.2% in Marine Isotope Stage 2, to 71.9% in the African Humid Period.

Small Mammal frequencies from Guli Waabayo, presented as % (number of identifiable specimens). Abbreviations: MIS 2, marine isotope stage 2; AHP, African humid period; ka. thousand years ago. Jones (2023).

In both intervals, the age-profile of the Dik Dik remained fairly constant, with all age groups present, but adults making up almost half of the assemblage in both cases, while the other three age groups each made up between 13.3% and 21.3% of the assemblage, although Jones notes that the sample size is very small for the African Humid Period, which may mask greater variation than is recorded.

Dik-dik mortality profiles from Guli Waabayo, presented as % (minimum number of individuals). Abbreviations: MIS 2, marine isotope stage 2; AHP, African humid period; ka, thousand years ago. Jones (2023).

There was a relatively high proportion of Carnivore remains in both Marine Isotope Stage 2 (4.4%) and the African Humid Period (5.4%). This could potentially indicate that the site had at times been occupied by Carnivores (for example Hyenas), and that they would have been responsible for some of the other remains present. Signs of such denning behaviour are considered to include juvenile Carnivore bones, gnaw marks on the bones of other Animals. Of 88 identified small Carnivore bones present in the total collection from Guli Waabayo, only three had unfused epiphyses (two from indeterminate Carnivores and one from a small Felid). In addition a mandible was identified with an erupting third molal (a sign that it came from a juvenile Animal), which could be assigned to a Dwarf Mongoose, Helogale sp.. No bones could be attributed to juvenile medium or large Carnivores, such as Hyenas or Lions, and no neonatal Carnivore bones of any type were found. A single Dik Dik bone from Marine Isotope Stage 2 showed signs of gnawing, but this was clearly by a Rodent Three Dik Dik bones from the African Humid Period also showed signs of gnawing, with two of these again clearly having been gnawed by Rodents. Conversely, none of the Dik Dik bones showed signs of butchery with tools, although this would not typically be expected with the bones of an Animal this small. Signs of burning, another clear indicator of a Human predator, were found on 12.3% of the Marine Isotope Stage 2 Dik Dik remains, and 7.8% of the African Humid Period Dik Dik bones.

Jones's investigation confirmed that Later Stone Age hunters had extensively targeted Dik Dik, small territorial Bovids, during both Marine Isotope Stage 2 and the African Humid Period, something in line with previous research at the site. However, unlike previous researchers, Jones did not concentrate on the toral diversity of remains at the site, but specifically upon the most abundant item, the Dik Dik, with a view to understanding the importance of these Antelopes to Later Stone Age hunting communities in the area. The evidence strongly suggests that the people here became specialist hunters of Dik Dik during the arid Late Pleistocene. 

This is different to the pattern observed at the Riffle Range Site, where a specialisation in net-hunting of Dik Dik appears to have developed during the African Humid Period, alongside a more general shift towards a more settled lifestyle and the permanent holding or territory by Later Stone Age groups. At Guli Waabayo the proportion of Dik Dik among the Animal remains increased in the African Humid Period, suggesting that they became a more important resource during this time, but they were clearly already a major prey species in Marine Isotioe Stage 2. This suggests that the change in climate did not provoke a major change in hunting methods or other behaviours at Guli Waabayo. This raises the possibility that the apparent change in behaviour at the Riffle Range Site 12 000 years ago may be actually indicate that the site was only sporadically inhabited prior to this, or simply that earlier remains have not been preserved. An increase in usage of the Riffle Range Site during the Holocene appears to be the more likely scenario, but why that should be is unclear.

The hunters of Guli Waabayo appear to have taken far more Dik Dik than any other Animal, even similar sized small Mammals, such as Hares or Hyraxes. Thus suggests that they had developed a specialised hunting technique which enabled them to reliably capture large numbers of Dik Dik, bur which did not work as readily on other prey. Furthermore, the Dik Dik remains have a distinct age profile, with all age groups present but adults being the most abundant. Based upon studies of modern Dwarf Antelope hunters, the most likely explanation for this is that communal net drives were used to target Dik Dik on a regular basis, but that other hunting methods were also used to catch other prey.

The selection of faunal remains present at Guli Waabayo makes it likely that both snare and pursuit hunting were also practiced, but the large proportion of Dik Dik in the remains cannot be explained by this. Snares will tend to target a range of Animals of the same size, including small Carnivores, which are present in the Guli Waabayo assemblage, but do not tend to capture juvenile Animals, which are typically too small to trigger them. Had snares been the primary hunting method at Guli Waabayo, then the overall assemblage of Animals would have been more varied, without any single species dominating, and juvenile Dik Dik would have been largely absent. Pursuit hunting, in contrast, tends to produce a far higher proportion of juvenile Animals, which are easier to run down, but again does not tend to produce an assemblage dominated by a single species. 

The bone assemblage from Guli Waabayo was accumulated over thousands of years, making it unlikely that all of the Dik Dik remains there are the result of a single hunting method. Nevertheless, it does appear likely that net hunting played a significant role in the life of Later Stone Age hunters in the region.

The most egalitarian hunter/gatherer societies tend to invest little effort in the manufacture of material goods. However, as societies become more complex, an increasing emphasis is placed upon resource management, and delayed-return technologies, which ensure the future supply of food. Such technologies include both devices used to store food, and to capture Animals for consumption, and are in turn associated with a decrease in mobility, with groups tending to remain in one place for longer. In environments with strong seasonal variations in the availability of food, this can lead to the development of flexible political systems or large resource-sharing networks. 

The Holocene Kansyore people of the Lake Victoria Basin are known to have used pottery and Fish weirs d to exploit seasonal Fish stocks, a specialization which resulted in long-term occupation of sites, and presumably a sense of ownership of those sites. The connection between subsistence, land use, and technology during earlier periods in East Africa is less clear, although the specialisation in hunting Dik Dik in the pre-Holocene Later Stone Age at Guli Waabayo suggests that these people had developed a lifestyle in Marine Isotope Stage 2 which was tied to long term resource management within a single environment, and the use of specialist equipment, in the form of nets, which would have required considerable investment in time to construct and maintain.

The traditional view of Late Pleistocene hunters in East Africa has been one of highly mobile groups which placed an emphasis on the hunting of large game. However, the evidence from s coastal and forested regions suggests that small game may have been a far more important resource than was previously thought, to at least some groups of hunter/gatherers. The use of delayed-return hunting technologies, such as snares, traps, or nets, would have allowed more settled populations in coastal and forested environments, enabling a diversity of survival strategies to survive alongside one another in East Africa during the Late Pleistocene. The evidence for specialist Dik Dik hunting at Guli Waabayo adds to this understanding, giving an example of a community surviving in a semi-arid environment by adopting a more settled lifestyle and a specialist method of hunting.

Most hunting methods target one Animal at a time, but net drives can trap numerous Animals at the same time, potentially leading to surpluses of meat following the hunt. Among modern communities which practice net hunting in the Congo Basin, hunts are typically communal activities, involving men, women, and children, and the re-enforcement of social bonds. In Australia the organisation of net-hunts has been associated with the emergence of political hierarchies, while in the southwestern US the organisation of communal net hunts for Rabbits has been associated with both forming co-operative bonds between different groups, and the establishment of temporary leaders.

No direct evidence of netting has been found at Guli Waabayo, however, the pattern of remains suggests that the hunters here were using communual net-drives to capture large numbers of Dik Dik during the Later Stone Age. Such hunts would likely have resulted in considerable amounts of meat becoming available at certain times, enabling the surplus to be shared among large extended communities.

If this were the case, then Dik Dik hunting would have enabled large communal gatherings, promoting social cohesion between groups and promoting further co-operation. However, the faunal data is not enough in itself to make many assumptions about these ancient societies, and further analysis of Human burials from Guli Waabayo and Gogoshiis Qabe, combined with studies of lithic tools from these sites and the Riffle Range Site, should help us gain a better understanding of these ancient East African communities.

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