The Blood Customs of Dahomey.

The West African coast on the Gulf of Guinea comprises a network of lagoons and waterways stretching from the Volta River to Lagos, with hundreds of miles of navigable waterways, good agricultural land, and both access to the Atlantic Ocean shelter from its storms. People have lived in this area for at least two-and-a-half thousand years, potentially much longer, although we do not really know how long. The archaeological record of this area is poor. The wet tropical climate does not favour preservation, much of the landscape is made up of shifting islands and waterways, which appear and vanish over time, and many old occupation sites are seen as sacred by the local population.

During the twelfth and thirteenth centuries AD, Aja-speaking peoples migrated from the Tado area on the River Mono in what is now eastern Togo migrated southeast onto what is now southern Benin, conquering or intermingling with the peoples already living their, and forming a new ethnic identity, the Fon. By around 1600 AD, this are had been split into three kingdoms, Allada (or Greater Arda), on the southeastern coastal plain of modern Benin, Ajatche (or Lesser Arda) on the southwestern coastal plain, and Dahomey on the Abomey Plateau, inland and to the north of both.

Initially Allada appears to have been dominant to the other two kingdoms, which paid it tribute, but sometime between 1620 and 1645, however, this relationship was overturned when a King of Dahomey, probably Dakodonu, overthrew the king of Allada. The name Dahomey itself is probably a misnomer, meaning, roughly 'The King had his stomach ripped out' in the Fon language, in reference to this overthrowing of Allada; the people of the kingdom probably always referred to it as 'Abomey', the name of both the capitol city and the plateau on which it stands, but European travellers favoured Dahomey, which was the name eventually given to the French colony which was formed in 1894; this later gained independence as the Republic of Dahomey in 1960, changing its name to the People's Republic of Benin in 1975 (confusingly, this refers to the Bight of Benin, part of the Gulf of Guinea; the city of Benin is far to the east in Nigeria), and the Republic of Benin in 1990.

Dahomey appears to have embraced the opportunity to trade with Europeans early. It initially traded extensively with Portugal, importing firearms, cloth, beads and Cowrie shells, cloth, and alcohol, and exporting local goods and slaves. This trade appears to have been profitable to both sides, relations were good between the nations, and Dahomey established a permanent diplomatic mission in Brazil, so important was the trade relationship. Portuguese traders recorded Dahomey as a civilised country with a ruling class living in walled cities and an agricultural economy, based largely on serf labour, not notably different to its neighbours. Most of the slaves purchased there were local serfs, with some prisoners of war captured from neighbouring states.

Over time, the Portuguese were replaced as the main trading partners of Dahomey by the British, whose merchants told a very different story. They recounted Dahomey as a fierce warrior kingdom, with a standing army of over ten thousand, including regiments of female amazon warriors. This Dahomey waged constant war against its neighbours, and exported thousands of slaves every year, all apparently taken in battle. The agricultural system had been transformed, with a plantation system imported from the Americas and manned by more captured slaves, the Dahomeans themselves appear to have been almost entirely employed in the business of war (even the women). Furthermore, each year a great ceremony, known to Europeans as the 'Customs of Dahomey' took place in which hundreds of slaves were sacrificed to the local deities of the Voodún (or Voodoo) religion (despite their willingness to trade with Europeans, the Kings of Dahomey appear to have had little time for Christian missionaries). 

An illustration depicting victims awaiting sacrifice at a ceremony in Abomey in 1793, from Archibald Dalzel's The history of Dahomy, an inland Kingdom of Africa. New York Public Library/Wikimedia Commons.

There are a number of potential explanations for this difference. Possibly, the Portuguese were unaware, or untroubled, by the ferocious nature of the Dahomeans, although given the apparent closeness of the two nations, and the general enthusiasm of the Portuguese Empire for spreading the Catholic faith (which was willing to put up with quite a bit of brutality in the name of Christ, but nor any pagan deity), this seems unlikely. Alternatively, British merchants may have greatly exaggerated the behaviour of the Dahomeans. This would have had some advantages, if Dahomey was such a wild and savage place, then the merchants who ventured there would be seen as correspondingly more brave and adventurous by their peers. Also, slavery was much easier to justify if you were taking slaves from brutal masters in Africa, who might sacrifice at any moment on the whims of the priests of some terrible pagan god, and selling them on into the comparatively benign stewardship of a Christian planter in the New World. A third explanation is that the size of the slave industry had a brutalising affect on the kingdoms of West and Central Africa, as over time each nation had to raise a larger-and-larger army to protect its own citizens from raids by its neighbours, and was forced to fund its armies by carrying out more-and-more raids on its neighbours. Under this scenario, which was favoured by abolitionists in Britain and elsewhere, smaller African states were wiped from the map, with their citizens being shipped en mass to the Americas, while the remaining nations needed not just to be well armed, but to appear sufficiently alarming to be left alone.

The Voodún religion is not a doctrinal monolithic faith with a single religious text like Christianity or Islam, but rather a polytheistic religion with a large (and variable) number of deities, with different stories told about them in different places. The religion has historically been followed in the southeast of Ghana, southern parts of Togo and Benin, and the southwest of Nigeria (although the variable nature of the religion means that there can be some doubt about where it starts and finishes), and has spread to areas of the New World, including parts of Brazil, the Caribbean, and the Southwestern United States. 

Voodún is split into a large number of cults, each dedicated to a specific deity, with different forms of worship, although most commonly through 'fetishes', idols in which divine spirits are believed to reside, and to which sacrifices (typically of food) are made. Some are open in their worship, organising large public festivals and parades; others are secretive in nature. A persistent rumour among Voodún worshippers, and neighbouring peoples, is that some of these more secretive cults practised Human sacrifice (a popular trope in both Hollywood and Nigerian movies), something which appears to have become part of the official state religion in Dahomey. 

From 1728 the Oyo Empire, a Yoruba state based in western Nigeria, which had by this time expanded to cover most of Yorubaland (the area occupied by culturally Yoruba people) and several areas outside it, staged a series of invasions of Dahomey, finally incorporating it as a vassal state in 1748. During this time a Dahomean King named Agaja married a Voodún priestess named Hwanjile, about whom there are a variety of legends. Hwanjile may have been herself captured by the Dahomeans as a slave, before marrying the king, and may have brought two sons from a previous relationship with her, who then became stepsons of the king. It is also said that when the Oyo Emperor demanded a son of the King of Dahomey be sent to him as tribute, and all of the other wives of the king refused to send their sons, that Hwanjile sent one of hers. 

The Oyo Empire at its greatest extent in about 1780. Lovejoy (2014)/Wikimedia Commons.

Agaja was a successful military leader, who expanded to borders of Dahomey in several directions by taking over smaller states, but was unable to completely repel assaults by the Oyo Empire, a much bigger state with a large and highly effective cavalry. In 1740 Agaja died, probably in another battle with Oyo, he was not succeeded by his oldest son and official heir, Zinga, but by a younger son, Tegbessu, who was either a son of Hwanjile, or had made an alliance with her. 

Upon becoming king, Tegbessu appointed Hwanjile his 'Kpojito' a title which translates roughly as 'Queen Mother'. Similar roles exist in many traditional African kingdoms, and imply a female relative who co-rules with a king. This is not usually the king's spouse, and while it could be his mother, it might equally be an aunt or sister. In this role Hwanjile appears to have significantly re-arranged religious practices within Dahomey, introducing new deities, and (allegedly) the custom of human sacrifice. The name Hwanjile has itself become an important title, with a succession of priestesses descended from the original Hwnajile taking on the name and serving as chief priestess to this day. Tegbessu was eventually forced to submit to dominion by the Oyo Empire, but remained an important ruler, reigning over Dahomey until his death in 1774.

Tegbessu was followed as king by Kpengla (the relationship between the two is unclear), who ruled till 1789, with the throne passing to his son, Agonglo, who ruled till 1797, and was succeeded by his son, Adandozan. All three of these kings struggled against, but were ultimately unable to overthrow, dominion by the Oyo Empire. In 1818 Adandozan was overthrown in a coup by his brother (or son, depending on the version of the story), Ghézo, with the aid of a Brazilian slave trader called Francisco Félix de Sousa. With de Souza's help, Ghézo significantly restructured the Dahomean state, replacing a peasant-based agricultural system with plantation system manned by captive slaves, similar to that of Brazil, and using the freed Dahomean workforce to vastly expand the military. 

In 1823, the Dahomeans began raiding areas outside of Dahomey but considered to be under the protection of the Alaafin (Emperor) of Oyo. The Alaafin immediately ordered that this cease, and a large sum of reparations be paid. Instead, Ghézo sent de Souza as a messenger to Oyo, demanding that Oyo make peace. The Alaafin rejected this, and dispatched an expeditionary force to Dahomey, with the aim of putting down the rebellion, but this was defeated by Ghézo's reformed military, securing independence for the kingdom.

Portrait of King Ghézo of Dahomey, painted some time before 1851. From Frederick Forbe's Dahomey and the Dahomans. New York Public Library/Wikimedia Commons.

In theory, this should have left Dahomey well-placed, and independent nation with a strong military, no need to pay tribute to a larger state, and having good contacts with the market to which its primary commodity, slaves, was sold. Unfortunately, in the 1833 slavery was abolished throughout the British Empire, and the British Navy began imposing a blockade on the Atlantic coast of Africa intended to prevent everyone else stop trading in slaves too. This did not bring an immediate end to the slave trade (demand for slaves was growing in the Americas at this time, British merchants were still quite happy to trade in goods derived from slave labour, and those freed by the British Navy intercepting slave ships were more likely to end up in indentured servitude in a British colony somewhere), but it did present a significant problem for African states with economies reliant on the export of slaves. 

This lead to a slow weakening of Ghézo's power both at within his kingdom, where a poor economy undermined support for his reign, particularly following the death of de Souza in 1849, and in the wider region, where states such as Abeokuta and Badagry in southern Nigeria profited by forming close ties with the British. In 1851 Ghézo launched a disastrous invasion of Abeokuta, in which the armies of Dahomey were defeated with British support, and which led to a British naval blockade of all ports controlled by Dahomey. In 1852 Ghézo was forced to sign in a treaty promising to end the trade in slaves (although British officials in the region never believed he was completely sticking to this), and in 1853, in a second treaty, he agreed to end the sacrifice of captives taken in wars in the annual customs (convicted criminals could still be executed at these events). In 1857 Ghézo officially announced he was breaking these treaties and resuming the slave trade probably in response to internal pressures within his kingdom, and in 1858 launched a second war against Abeokuta. Unfortunately, the Dahomean army was defeated just as quickly on this occasion, and Ghézo was (probably) assassinated later that year, being replaced by his son Glele. Dahomey never regained its military strength, and in 1890 was invaded by France, being first made a protectorate, and then four years later officially part of French West Africa. 

Each of the Kings of Dahomey built their own palace complex, including a mixture of residential, administrative, and ceremonial buildings. In 1944, Charles André Maurice Assier de Pompignan, Lieutenant-Governor of the Dahomey Colony and Dependencies, converted these palaces into a museum, the Royal Palaces of Abomey, open to the public, something which they remain to this day, additionally becoming a UNESCO World Heritage Site in 1985. Between 2018 and 2022, a collaboration between the Musée du quai Branly - Jacques Chirac, the Université d'Abomey-Calavi, and the French Ministère de l'Europe et des Affaires étrangères, were allowed to carry out archaeological investigations at the Royal Palaces of Abomey.

Entrance to the Palace of Ghézo at the Royal Palaces of Abomey. Karalyn Monteil/UNESCO.

Within the compound of Ghézo at the Royal Palaces of Dahomey is the tomb of the king himself. The mortar from which these buildings were made is reputed to have contained a mixture of ritual ingredients, including the blood of 41 sacrificed Human victims (the ritual inclusion of magical or sacred ingredients into the mortars from which traditional buildings were made is not unusual in West Africa, but including the blood of sacrificed Human victims certainly is).

The archaeologists were able to take two samples from an inside wall of King Ghézo's tomb at the Royal Palaces of Dahomey. These were then tested for their protein content, with the results of the study being published in May 2024 in a paper in the journal Proteomics, by Philippe Charlier of the Musée du quai Branly - Jacques Chirac, the Laboratory of Anthropology, Archaeology, and Biology at Université Paris-Saclay, and the Foundation of Anthropology, Archaeology, and Biology at the Institut de France, Virginie Bourdin, also of the Laboratory of Anthropology, Archaeology, and Biology at Paris-Saclay University, Didier N’Dah of the Département d’Histoire et d’Archéologie at the Université d’Abomey-Calavi, and Mélodie Kielbasa, Olivier Pible, and Jean Armengaud of the Département Médicaments et Technologiespour la Santé at Université Paris-Saclay, present the results of this study and discuss its implications. 

Proteomics, the study of proteins recovered from ancient materials, is a relatively new discipline, which has some distinct advantages over the more established discipline of palaeogenomics. Proteomics it targets ancient proteins rather than DNA. Since protein is much more stable than DNA, this enables the study of much older samples, as well as samples from environments where DNA is unlikely to be preserved well, such as the wet tropical climate of the Abomy Plateau. Moreover, while our DNA is identical in every cell, each tissue the body produces contains a number of unique proteins, which not only helps to determine the type of tissue being examined, but also helps to rule out contamination, as this is typically in the form of skin or hair cells, and can be excluded from a study such as that carried out by Charlier et al. which was looking primarily for traces of blood.

Illustration of the wall of King Ghezo’s tomb. Panel A shows a general view of the cenotaph wall. Panel B shows specific detail of the red wall buttressed by a wood beam. Philippe Charlier in Charlier et al. (2024).

Charlier et al. were able to identify 5866 different peptides (short chains of amino acids, which can be combined together to form proteins) in their sample, including 4208 from Bacterial proteins, 1240 from Eukaryotes, and 50 from Archaeans. Of the Eukaryotic peptides, 271 could be assigned to Chordates, including 49 from Bovids, 21 from Chickens, and 215 from Humans. Furthermore, while the Bovine peptides were largely associated with milk, many of the the Human and Chicken peptides were associated with blood, indicating that both Human and Chicken blood had been added to the mixture from which the mortar had been made, with Human blood making up a significantly larger portion.

Taxonomical results obtained for dataset B3. Five taxonomical ranks (Superkingdom, Phylum, Class, Order, Genus) are represented by rings (from the inner to the outer, respectively). The number of entities identified for each of the taxonomical ranks are separated by white lines,and their width corresponds to the abundance measured (TSMs ratio). The most abundant genera are indicated. Charlier et al. (2024).

Thus, Charlier et al.'s data strongly indicates that Human blood was a significant ingredient of the mixture from which the walls of the tomb of Ghézo were made. This in turn appears to support the more lurid stories of European, particularly British, visitors to the Dahomey from the eighteenth and nineteenth centuries, which emphasise the frequency and large scale of Human sacrifice within the kingdom. However, this is a study of a single tomb from a specific point in the history of Dahomey, and does not imply that this was the case throughout the kingdom's history, thus the earlier, largely Portuguese, accounts which do not mention these practices may equally be true, in which case the kingdom may have adopted these practices during its long involvement with the slave trade and constant wars with neighbouring states.

See also...

France returns skull of King Toera of Menabé to Madagascar.

The head of King Toera (or Itoera) of Menabé, a former kingdom in the west of Madagascar, betweem the Mangoky and Manambalo rivers, has been returned to the island by France. The former king was captured in battle following a rebellion against the French government of Madagascar in 1897, and was subsequently beheaded along with two of his supporters. The three heads were then taken to Paris, where they were placed in the collection of the Muséum national d'Histoire naturelle as 'anthropological specimens'.

Harea Georges Kamamy, the current King of Menabé, carrying his ancestors skull following a ceremony at the French Ministry of Culture on 27 August 2025. Stéphane du Sakutin/AFP.

The Kingdom of Menabé was formed in about 1540 by Adriamandazoala, who led a group of Sakalava people from the south of the island on a migration into the area (one of two kingdoms in the west of Madagascar formed by migrating Sakalava people at about this time, along with Boina, to the north of the Manambalo River. The kingdom grew under notable rulers such as Andriandahifotsy, (died 1685) and Andriandrainarivo (died 1727), but was incorporated into a unified Kingdom of Madagascar by Radama I in 1820 (although sporadic uprisings would occur until 1840 when Queen Ranavalona I placed garrisons in all the major towns of  Menabé and moved farmers into the region from other regions of Madagascar to dilute loyalty to the Menabé kings (who were nevertheless able to keep their titles, albeit as rulers of a subordinate kingdom).

In 1883 France invaded Madagascar, with fighting continuing till 1886, when a treaty was signed granting the French control of the port of Diego-Suarez (now Antsiranana) as well as the right to determine Malagasy foreign policy, making the island an effective (but not official) French protectorate. In 1894 France launched a second invasion, marching upon the capital, Antananarivo, which fell in September 1895, following a brief shelling by French artillery.  On 1 October 1895 Queen Ranavalona III signed a treaty making Madagascar an official protectorate of France. 

In 1897, following an uprising in central Madagascar, the Malagasy monarchy was abolished and Queen Ranavalona III was exiled to Réunion, never to return. The French military commander, Joseph Gallieni. King Toera did not take part in this initial rebellion, but when direct rule by France was declared, he joined the rebellion, putting together an army of 10 000 warriors armed with riffles. An expedition to Menabé was launched under the French General Augustin Gérard, who reached the Menabé capitol, Ambiky on 30 August 1987, which, like Antananarivo before it, fell after a short artillery bombardment. 

King Toera was captured and beheaded along with two of his senior advisers. Their heads were subsequently shipped to France and placed in the anthropological collection of the Muséum national d'Histoire naturelle. During the entire campaign only two French casualties were recorded, both soldiers from the Tirailleurs Sénégalais (a West African regiment that did not recruit exclusively from Senegal).

More than 20 000 sets of Human remains are still held in the collection of the Muséum national d'Histoire naturelle, including hundreds more from Madagascar, with collections also held by other French museums and universities. Similar collections are held by institutes in other countries in Europe, as well as other parts of the world. 

In 2023 France changed its laws on the ownership of such remains, in order to allow their return to countries from which they were obtained, this having become a sticking point in relations with many nations around the world. The return of the skulls of King Toera and his advisers is the first such repatriation under the new law.

DNA tests carried out at the Muséum national d'Histoire naturelle proved unable to determine which of the skulls belonged to King Toera, although they were able to determine that they did belong to people of Sakalava origin, so an identification was carried out by a Sakalava spirit medium. The three skulls were handed over in a ceremony in Paris on 27 August 2025, and reached Antananarivo on 1 September. From here they are being taken overland to Menabé, where they will be buried.

The three skulls arriving in Antananarivo on 1 September 2025. Mamyrael/AFP.

See also...

Little Foot.

In the late nineteen thirties, palaeontologist Robert Broom discovered a series of Hominin skeletons in a cave system in Sterkfontain, South Africa. These, he assigned to the genus Australopithecus, which had been described in 1925 by Raymond Dart, suggesting that he had found a second species in the same genus. At the time, these discoveries did not receive a great deal of attention from the international scientific community. The cave deposits at Sterkfontain were estimated by Broom to be Late Pleistocene in age. This made Broom's material much younger than Dart's and contemporary with modern Human remains from many parts of the world, which meant that, while interesting, Broom's new 'Ape-man' could not be ancestral to modern Humans.

More locally, however, the discoveries caused considerable interest, leading many amateur fossil hunters to begin exploring the Sterkfontain cave system. One of these was Helmut Kurt Silberberg, a Johannesburg art dealer and antiquarian with an interest in prehistory. In 1942, Silberberg's explorations led to the discovery of a small cave with numerous bones embedded in the walls. Unfortunately, the rock in which the bones were embedded was extremely hard (more so than the bone) making excavation difficult. Silberberg paid a local man to hammer free a chunk of rock, during which process several bones were broken, displeasing Silberberg. When the bones in the rock were examined, they were found to belong to a Hyena and a Baboon. 

Discouraged by these findings, Silberberg lost interest in the work, giving his specimens to Broom, and moving onto other projects. While the fossils were not interesting to Silberberg, the material proved to be much more useful to Broom, who recognised the Hyena as a member of the genus Lycyaena, which was otherwise known from the Early Pliocene of Europe and India. Even if this South African specimen was younger than other fossil Hyenas assigned to the genus, it was unlikely to be Late Pleistocene in age. This caused him to re-evaluate the age of the cave system using Mammal fossils to establish an age sequence, which proved that his Australopithecus fossils were much older, and therefore much more significant, than he had originally realised. Broom named the Hyena fossil Lycyaena silberbergi (a name which is no longer used) and the cave where it was found was eventually named Silberberg Grotto.

In 1966, palaeoanthropologists Philip Tobias and Alun Hughs began a new series of excavations at the Sterkfontein Caves, working methodically through the cave system from west to east, and uncovering several new specimens, albeit mostly fragmentary. In 1978 they revisited Silberberg Grotto, removing large sample of breccia, which was cleaned by technicians at the site to recover a larger amount of fossil material, most of which was moved to the University of the Witwatersrand, while some remained in storage at Sterkfontein.

From left to right, Raymond Dart, Alun Hughes, and Phillip Tobias at Witwatersrand University in the mid-1970s. Clarke & Kramer (2012).

In 1992 Tobias' team returned to Silberberg Grotto, this time employing two mining engineers, John Cruise and Dusty van Rooyen, to blast away a section of the fossil-rich wall of the cave. Analysis of the material recovered by this process in 1994 by palaeoanthropologist Ronald Clarke revealed something surprising. The rocks had produced numerous bones associated with Carnivores and Monkeys, but for some reason almost no Bovids (usually the most abundant Mammals in South African deposits). This led him to return to the material stored at Witwatersrand, which he also analysed for the proportions of different Mammal groups. 

While examining this material, Clarke discovered what appeared to be the talus, navicular, medial cuneiform and proximal half of the first metatarsal of an unknown Hominin. This was surprising for two reasons. Firstly, the material had been examined previously by specialists palaeoanthropologists looking for exactly this sort of material, and secondly because the Silberberg Grotto had by this time been identified as somewhat older than the previous Hominin-producing deposits at Sterkfontein, implying any fossil Hominin found here could be the oldest yet identified specimen from the cave system. In addition to these four bones, Clarke found what appeared to be a badly damaged lateral cuneiform from the right foot, and a fragment of a possible Hominid calcaneum.

This material suggested a small Hominin, capable of standing upright like a Human, but still retaining an disposable big toe like an Ape, which suggested that at least some of its time was still spent within the trees. This specimen was given the official designation StW 573, but quickly became known as 'Little Foot'.

In 1997 Clarke discovered a box labelled 'D18 Cercopithecoids' (Old World Monkeys), which when inspected was found to also contain material labelled 'D20 Cercopithecoids'. Since D20 implied the layer of material exposed at Silberberg Grotto, and Clarke was interested in the fauna of this cave, he took down this box to inspect its contents. Within this material he found a Hominid intermediate cuneiform, which he found fit with the lateral cuneiform associated with Little Foot. Along with this was the left lateral cuneiform, the proximal end of the left second metatarsal and the left distal fibula. A subsequent search of related material uncovered the disto-medial portion of a Hominid tibia. This did not fit exactly with the Little Foot talus, but was close enough that, once the damage to the material was taken into account, Clarke considered it likely to have come from the same individual.

This tibia had a break across its shaft consistent with the sort of damage caused by mine blasting, Clarke next turned to the material stored at Sterkfontein. Here, in a bag of material labelled as Bovid tibiae, he found another Hominid tibia, which this time fit perfectly with the left talus of StW 573, leading him to realise that the previously discovered tibia was in fact from the right leg. Further inspection of this material also produced a second piece of cuneiform, which fit together with the one from the original Little Foot material, giving a complete bone. This had a bowl-like surface on its articular surface with the cuboid bone, indicating that these bones still articulated (could be moved relative to one-another), as in Chimpanzees, and unlike the condition seen in Modern Humans, where the two are immobilised by a keel-and-socket arrangement, which gives more stability when walking and running upright.

This gave Clarke a collection of twelve bones from the feet and lower legs of a single Hominin, including the left tibia and fibula, which joined to an articulated set of eight foot and ankle bones, and the distal fragment of a right tibia and right lateral cuneiform.

Convinced that more material might still be found in Silberberg Grotto, Clarke prepared a cast of the distal fragment of the right tibia of StW 573, which he gave to two perpetrators based at Sterkfontein, Nkwane Molefe and Stephen Motsumi. Molefe and Motsumi then searched the area exposed by blasting within Silberberg Grotto, looking for an exposed cross section of bone which would fit this cast. 

Clarke was not optimistic that this approach would work. The various excavations had turned the grotto into a cavern of some size, with exposed breccia on the walls, floor and ceiling, all of which might contain the matching bone fragment - if it was there at all. As if this was not difficult enough, there was neither natural light nor a rigged lighting system within the cave, forcing the two investigators to search using hand-held torches. 

After two days Molefe and Motsumi reported finding a bone embedded in a talus slope at the western end of the grotto, which was a perfect match for the cast given to them by Clarke. This was unexpected, as this was not the area where Cruise and van Rooyen had carried out their blasting, but an older area of excavations where blasting had been carried out by commercial miners extracting limestone for use in the construction industry, an operation which had stopped 65 years before. Furthermore, a second section of bone could be seen to the left of this bone, which turned out to be a perfect match for the broken left tibia.

Ronald Clarke, Stephen Motsumi and Nkwane Molefe. Kathleen Kuman/Mail & Guardian.

Following this discovery, Clarke, Molefe, and Motsumi began excavating the site using hand tools, taking almost a year to uncover the left tibia shaft and the distal ends with lower shafts of the left and right femurs. After which, however, the material apparently stopped, until Motsumi hammered away a section of travertine (dense limestone deposited by hot springs) higher up the slope exposing more breccia, and the distal and of a Hominid humerus and mandible. The skeleton had apparently been entombed in rock, broken in half by an ancient rockfall, and covered by travertine from an ancient hot spring, before eventually being exposed by limestone miners some time in the early twentieth century.

In the following months, and eventually years, the skull, left forearm, left hand, pelvis, ribs, vertebrae, humerus, and most of the lower limb bones were extracted. The final skeleton was 90% complete, the most complete non-Human Hominin skeleton ever recovered. For comparison, the famous Lucy (or Dink'inesh) skeleton AL 288-1, considered to be the most complete specimen of Australopithecus afarensis, is only 40% complete.

The complete skeleton of StW 573, Little Foot.  Paul John Myburgh/ University of the Witwatersrand.

The Sterkfontein cave system, now part of the Maropeng Cradle of Humankind World Heritage Site, are a karstic cave system on the south side of the Bloubank River, about 50 km to the northwest of Johannesburg. Karstic cave systems form as water percolates through limestones, dissolving some of the mineral to create voids. This is a complex process, as cave systems form, are enlarge by the hollowing-out of water erosion, then collapse bringing sediments from above down to lower levels and often redirecting the water flow which caused the hollowing out. Furthermore, the limestone dissolved in one place is typically re-deposited in another, as travertines or flowstones. All of this makes establishing timelines for the karstic deposits extremely complicated.

Sterkfontein Caves began to form about six million years ago, towards the end of the Miocene. Throughout much of the intervening time the local environment has been a mosaic of grasslands, woodlands, and lakes, albeit with warmer, wetter phases, and cooler, dryer phases. At least five different Hominin species have been found within the caves, some of whom appear to have sought shelter within cave systems, others to have been dragged their for consumption by cave-dwelling carnivores, while others appear to have fallen into sinkholes.

Skeleton StW 573, or Little Foot, was found closely associated with a large Baboon (an animal quite capable of predating a small Hominin), but does not show any signs of having been consumed. Rather, it seems to have fallen face-down into a shallow puddle of water within a sinkhole. Possibly the Hominin and Baboon fell together following a struggle, but this is guesswork we will never be able to prove. The chamber where it was found contained Pliocene Hyenas, and was directly below the chamber which produced Bloom's most famous Australopithecus specimen, Mrs. Ples (STS 5), and therefore must be older than that specimen.

In 1978, geomorphologist Timothy Cooper was part of the team led by Tobias and Hughes which re-examined the Silberberg Grotto. He established that the chamber had formed as a cone-shaped void, the walls of which had then been covered by flowstone (dissolved and then re-deposited limestone), before an opening had reached the surface, forming a sinkhole into which surface sediments, and various Animals now preserved as fossils, had fallen, filling in the cone. These sediments were then covered by a second flowstone deposit. The sediments within this cone were thought to be about 2.8 million years old (Late Pliocene) on the basis of the fossil fauna they contained, implying a similar age for Little Foot. 

However, Molefe and Motsumi had established that Little Foot had in fact come from below the lower flowstone deposit, with only a small portion displaced into the accessible sediments by an ancient rockfall and the actions of early twentieth century miners. Various attempts have been made to date this flowstone deposit, including cosmogenic uranium-lead series nuclide dating, palaeomagnatism. These studies produced dates between 4 million years and 1.07 million years ago, although most fell into the range between 2.8 and 2.2 million years ago. However, subsequent studies also established that the flowstone itself contained pockets of younger, redeposited material, making all dating of this layer suspect.

Cosmogenic aluminium²⁶ and beryllium¹⁰ dating works because both aluminium²⁶ and beryllium¹⁰ are produced in quartz exposed to cosmic rays (i.e. on the surface) at a steady rate, producing a ration of 6.75 aluminium²⁶ to 1.0 beryllium¹⁰. Once the quartz is buried no more of either radioisotope is formed. Since the half-life of aluminium²⁶ is roughly half that of beryllium¹⁰, over time the ratio between the two radioisotopes will change, enabling the precise measurement of both how long a piece of quartz was exposed on the surface, and how long ago this happened. In April 2015 a study led by Darryl Granger of Purdue University produced a date for the sediments in which StW 573 was directly lying, based upon cosmogenic aluminium²⁶ and beryllium¹⁰, yielding an age of 3.67 million years before the present (equivalent to the dates for early Australopithecus afarensis specimens from East Africa. 

Darryl Granger. Lena Kovalenko/Purdue University.

This also makes StW 573 the oldest Hominin fossil discovered in South Africa, as well as the most complete pre-Human Hominin fossil ever found. It also predates the oldest stone tools in South Africa by over a million years, and the oldest known stone tools anywhere by over 300 000 years, suggesting that it came from a population which did not yet have this technology. 

Studies of Little Foot have suggested that she was female, and stood about 1.20-1.30 m tall, with longer legs than arms (similar to modern Humans, but unlike Chimpanzees). She had an 'S' shaped collarbone, which is a more Human-like trait, and but the shape of her pectoral girdle and shoulder blades, suggest that she was comfortable moving in trees suspended by her arms. The structure of its foot suggests that it was able to walk in an erect manner, similar to modern Humans, but still had an opposable big toe, implying it was still a comfortable climber (Pliocene Hominins are in general thought to have been woodland species, with the preference for open grasslands emerging in the Pleistocene when a cooler, dryer climate led to the retreat of forests and expansion of grasslands). 

This fits well with the environment in which her remains were found. While the Silberberg Grotto produced the remains of some terrestrial Animals, such as Antelope and Hyenas, the majority of the fossils are of Animals associated with an arboreal lifestyle, such as small Monkeys and Felids. Such Animals might have climbed down into the sinkhole in order to obtain water and become trapped, or possibly have fallen directly from trees into the hole - in Little Foot's case, possibly while trying to escape from a large Baboon.

Surprisingly, despite the completeness and significance of Little Foot, she has never been assigned to a specific species. She was referred to by Ronald Clarke as possibly a specimen of Australopithecus prometheus. However, this species name is now considered to be invalid; it was used by Raymond Dart to refer to a partial occipital bone (part of the back of the skull) from Makapansgat, but this has now been assessed as a specimen of Australopithecus africanus, a species common in South Africa from about 3.3 to about 2.1 million years ago. However, Little Foot is not generally accepted as belonging to Australopithecus africanus, which it predates as well as differing from in its brain capacity (smaller in Little Foot) and shoulder arrangements (more Ape-like in Australopithecus africanus). Nor does she conform to Australopithecus afarensis, a species whose time range she overlaps, but which is not found in Southern Africa. 

Thus Little Foot appears to be an otherwise entirely unknown species of Australopithecus from South Africa, with an arboreal lifestyle and no use of stone tools. This is not completely surprising, as species such as Australopithecus africanus and Australopithecus afarensis are clearly already some way removed from their closest Ape relatives. and species which live in trees seldom have good (if any) fossil records.

See also...

The origin of the ancient Dembeni people of the Comoros Islands.

When the first Europeans reached the Comoros Islands in the late fifteenth century, they found a group of small independent city-states forming part of the 'Swahili Corridor', a system of small island and port-city states which stretched up the east coast of Africa, also including cities on Madagascar and the coasts of what today are Mozambique, Tanzania, and Kenya. These states were bound together by a common language, Swahili, and a common religion, and traded slaves and African goods with the Middle East and other areas of the Indian Ocean rim. Local legend had it that the people of the Comoros were descended from King Solomon and the Queen of Sheba, and that upon hearing of the Profit Mohammed, an emissary was sent from the islands, who returned bringing with him the Islamic faith, and built the first mosque in the archipelago on the island of Ngazidja. European ethnologists concluded that the Swahili culture came about as Arab traders moved down the coast of East Africa, taking local women as wives (or, in more lurid accounts, concubines). 

The position of the Comoros Islands in the Mozambique Channel. The island of Ngazidja is now more commonly known as 'Grand Comoros', and is home to Moroni, the capital of the Union of the Comoros, which gained independence from France in 1979 as the 'Federal Islamic Republic of Comoros', adopting its current name in December 2001. The island of Moare remained a French possession, now forming the Department of Mayotte. Wright (1975).

In the 1970s and 80s a series of archaeological digs were carried out in the Comoros. The earliest culture uncovered by these excavations was identified as the 'Dembeni Phase', in reference to the location where it was first discovered, close to the village of Dembeni on the island of Moare (now the French department of Mayotte). Overlying this in places were layers assigned to the more widespread 'Hanyundru Phase', which can be clearly associated with medieval Swahili culture, with buildings made from stone or coral and cement, and imported Middle Eastern Glassware.

Six sites were associated with the Dembeni Phase, Dembeni itself plus Old Sima on the island of Ndzuwani (now more commonly known as Anjouan), M'Bachile, Dzindani, and M'Beni, on the island of Ngazidja (now usually referred to Grand Comoros), and M'Ro Deoua on Mwali (now usually Mohéli). The Dembeni people built small-to-medium sized villages, comprising houses built from wooden posts with a mud covering, on beaches or ridge-tops, always close to lagoons, and with access to freshwater streams. They consumed significant amounts of seafood (Fish, Shellfish, and Turtles), and kept Carrle and Goats, and farmed crops such as Rice, Millet, and Coconuts, which are still common across the region today.

The Dembini people produced distinctive pottery, with two distinct styles found on the islands and thought to have been made locally. The first of these is plain, with a smooth exterior and a burnished interior, sometimes embellished with a band of incisions or shell imprints. The second is covered with a red slip, either on the interior or the exterior, or both, and is typically burnished and decorated with graphite. In both cases the clay used was course and contained particles of red, grey, and white grit, although the former was typically courser. This courser, plainer style was used to make jars and pots, many of which show signs of being used for cooking. The finer, more decorated, style was used to make shallow bowls, possibly for serving food. A single bead made from local pottery was found at Dembeni,

Examples of 'plain' Dembeni-ware pottery from (a) Dembeni, (b) Sima, (c) M'Ro Deoua, (d) Dembeni, (e) Sima, (f) Dembeni, (g) M'Beni, (h) Dembeni, (i) Dembeni, (j) Dembeni, (k) Dembeni, (l) M'Bachile, (m) M'Bachile, and (n) Sima. Carla Sinopoli in Wright (1984).

Dating of the Dembeni sites was difficult with the technology available in the 1970s and 80s. Attempts were made to obtain carbon¹⁴ dates from several items. A shell collected from a midden at Sima yielded a carbon¹⁴ date of 1550±70 years before the present. Since it was understood by this time that it was necessary to adjust dates from shells to take into account for the carbon¹⁴ content of the atmosphere and the ocean in which the shell had formed, this was interpreted as indicating a date of about 910 AD, with a 96% probability of a date between 785 and 1040 AD. A piece of charcoal from M'Bachile yielded a carbon¹⁴ date of 1140±60 years before the present, with a 96% chance of a date between 470 and 650 AD, while a second piece of charcoal from Sima yielded a carbon¹⁴ date of 1960±70 years before the present, with a 96% chance of a date between 155 BC and 215 AD. At first sight, these dates seem incompatible, sine the Dembeni culture is not thought to have persisted for many centuries, but dates obtained from charcoal represent the age of the original wood from which the charcoal was made, and if this wood came from the interior of a tree, may be much older than the burning event.

Another way to determine the age of archaeological sites is by comparing goods from them to similar items from other sites with known chronologies. Although the majority of the recovered material from the Dembeni sites appears to have been local in origin, items from further afield were found at all locations, together representing about 4% of the recovered ceramics. 

Among the items found at M'Ro Deoua were several large basins and the neck of a large pithoi (large storage jar) made from an oxidised yellow-green clay. These items were made from a base-material which contained fragments of older pottery and fragments of igneous rock, and were decorated with chopped-grass impressions. Smaller fragments of the same material were also found at Dembeni and Sima. This pottery resembled similar material known from several sites in the Near East, at sites dated to between the eighth and tenth centuries AD, although this pottery had never been the subject of any detailed study.

A second type of non-local pottery, found at all sites except Dzindani, comprised fragments of large sandy-ware jars decorated with incisions and barbotine appliqué, with a blue-green glaze on the external surface and an speckled grey or grey-green glaze on the interior surface. Similar jars are known from Siraf on the Gulf Coast of Iran, where they first appear in the eighth century, and on Manda Island, Kenya, where they are found alongside ninth-to-tenth century Chinese ceramics. 

The most common form of imported pottery, found at all of the Dembeni sites, were small fine bowls with a white glaze, typically with a splash of coloured glaze on the interior; the colour of this varies, with the most common being cobalt, turquoise, or gold. Similar bowls are known at Susa in southwest Iran, where examples with cobalt on the interior appear around 720 AD, while turquoise does not appear until about 820 AD, as well as at Shiraf, where cobalt appears around 800 AD and turquoise around 850. This may offer some insight into the chronology of the Dembeni sites, as only white pottery is present at M'Ro Deoua, cobalt at Dembeni, and turquoise at M'Bachile and Sima.

As well as pottery, glass fragments were found at Dembeni, Sima, M'Ro Deoua, and M'Bachile. These were greenish, yellowish, clear, or cobalt blue, and round or cylindrical in shape. All appeared to have been blown, and resembled glassware forms widespread in the Near East. A number of iron items, and slag from iron forging were also found. While it was unclear where these had come from, iron is not a locally available resource in the Comoros. 

All of this appeared to show that the Dembeni people occupied the Comoros between about 800 and 1000 AD, and that from the very earliest, they had strong trading connections with the Near East. This apparently supported the idea that they were migrants from this area, rather than the east coast of Africa. However, two grave sites were also found associated with one of the Dembeni sites, Ngazidia. These both had their heads facing to the south, at odds with Islamic custom, making it highly unlikely that they were Muslims, which would almost certainly have been the case with Near Eastern emigrees of the period.

Founded in 2008, the Sealinks Project seeks to study the movements of ancient peoples and development of trading routes around the Indian Ocean. One of the problems this has sought to resolve is the peopling of Madagascar by settlers from Southeast Asia, and the subsequent adoption of Southeast Asian crops, such as Rice, Coconuts, and Mung Beans into the cultures of the Swahili Corridor. Madagascar was settled in the first millennium BC by hunter-gatherers from Africa, but agriculture appeared to have arrived on the island much later, with settlers from Southeast Asia, and today about 10% of Madagascar's flora is of Southeast Asian origin, either crop plants of weeds introduced along with them.

Between 2010 and 2013, archaeologists from the Sealinks Project carried out excavations at early sites on Madagascar and the Comoros, and settlements of the Swahili Corridor on the coast and offshore islands of East Africa, looking for early examples of both African and Southeast Asian crops, which were then dated using modern radiocarbon methods. Unsurprisingly, Southeast Asian crops were almost absent from the African sites, with only small quantities found at the largest and most active ports. On Madagascar, Southeast Asian crops appear abruptly around 1000 AD, with no evidence of any agricultural activity before this point. Surprisingly, the Comoron sites associated with the Dembeni Culture were also overwhelmingly dominated by Southeast Asian crops (only Sima produced small amounts of African crops such as Millet), and these appeared around 750 AD, significantly earlier than on Madagascar.

(A) Map of eastern Africa, including the Comoros and Madagascar, showing the locations of sites included in the crop study. The relative proportions of African and Asian crops are shown for each site (percentages based on numbers of identified specimens per site). (B) Chronological summary of African vs. Asian crop patterns by site from north to south. Crowther et al. (2016).

This raised some surprising questions. The reliance of the Dembeni people on Southeast Asian, rather than African or Middle Eastern crops, suggests that they did not come from either of these areas, but were instead of Southeast Asian origin, potentially part of the great Austronesian migration of peoples from the Malaysian Peninsula through the Islands of Indonesia and on as far as Hawai'i and New Zealand, but also westward to Madagascar. With the Comoros having apparently been settled before Madagascar, it is possible that the larger island was secondarily settled by the Dembeni people, although it is also possible that older sites on Madagascar have not been located.

The modern people of the Comoros are not obviously Southeast Asian. They appear similar to the peoples of East Africa, and speak a language (Comorian) which belongs to the Sabaki language group, which is otherwise spoken along the Tana River of Kenya, and surrounding parts of the Kenyan coast. Their closest cultural links are with the Swahili culture of coastal Kenya, Tanzania, and Mozambique, itself thought to have derived from the Middle East through trade along the east coast of Africa. Possibly this could indicate that the Dembeni people, although the first settlers of the Comoros, did not survive contact with later settlers from East Africa or the Middle East.

With this in mind, a project led by scientists from the Laboratoire d’Anthropologie Moléculaire et Imagerie de Synthèse at the Université de Toulouse carried out a study in which the genomes of volunteers from the Comoros and coastal Swahili communities in Kenya were compared to an extant database, containing genomes from Africa, Madagascar, the Middle East, Southeast Asia, South Asia, East Asia, and Europe.

Surprisingly, despite long-held assumptions about their having Middle Eastern ancestry, the Kenyan Swahili populations were found to be of entirely sub-Saharan African ancestry. The Kenyan Swahili populations were found to have a homogeneous structure, all more closely related to one another than to any other group, and to be more closely related to Bantu-speaking populations than any other African group. Some of the Kenyan Swahili individuals had recent Somali ancestry, but otherwise the Swahili populations were strongly isolated from Somali and Ethiopian populations. The Comorian and Malagasy populations were more closely related to the Kenyan Swahili populations than to any other African population, although both had admixtures of Southeast Asian ancestry, which made up between 33 and 39% of the ancestry of the Malagasy populations, and 8-9% of the ancestry of the Comorians. Curiously, despite being further south than the Kenyan Swahili populations, the Comorians also had some Middle Eastern ancestry, which made up 6-7% of their genomes.

A statistical model of the ancestry of modern Comorians suggested that their common ancestry formed as a distinct population between 1197 and 792 years before the present, with the admixture of a Swahili and a Southeast Asian group. Different islands within the group have also had subsequent influxes of genetic material from Swahili, Middle Eastern, and Southeast Asian populations. The Madagascan populations showed a more recent origin, with an admixture between a Swahili Group and an Island Southeast Asian group between 876 and 742 years before the present (this population appears to have arrived in the area separately, and not to have migrated from the Comoros). 

Admixture Scenario for Populations along the Swahili Corridor as Estimated by GLOBETROTTER and ADMIXTURE Dark red arrows represent Swahili gene flow; light green arrows represent Island Southeast Asian Banjar or Malay; a yellow arrow represents Middle Eastern gene flow; and the purple arrow represents gene flow from the Horn of Africa. The pink arrow represents gene flow from central and southern Bantu speakers. Dates refer to the last detectable admixture event; dates below pie charts refer to the admixture event between the Swahili and Banjar or Malay; dates in italics represent secondary gene flow. Sex-biased gene flows are represented by male and female symbols in the tip of the arrows; note that they are not presentin Malagasy Antemoro and Comorian Moheli. Brucato et al. (2018).

Thus, the most likely scenario is that the Dembeni people reached the Comoros Islands from Peninsula Malaysia or Island Southeast Asia around 750 AD. They brought with them staple crops from Southeast Asia, as well as Cattle and Goats, and fished in the islands' lagoons and surrounding seas. They exchanged trade goods with the Near East, probably through early East African Swahili intermediaries, with whom they may have intermarried. Despite this cultural exchange, the Dembeni appear to have remained a distinct culture till around 1000 AD, when they were abruptly replaced by the Hanyundru culture, which appears to have been more distinctly Swahili in nature. Modern Comorians derive about 8-9% of their genetic ancestry from this group, and still farm many Southeast Asian crops, but speak a language of apparent Kenyan origin, and are culturally close to the Swahili populations of East Africa.

See also...

The Sarcophagus of Ankhnesneferibra.

Sarcophagus of Ankhnesneferibra, daughter of Psamtik II, 26th Dynasty, circa 530 BC. The British Museum.

Ankhnesneferibra was daughter of Psamtik II, the third Pharaoh of the 26th Dynasty of Egypt, known as the 'Sais Kings' or 'Saites', after the city of Sais in the Nile Delta, from where they ruled. In 595 she was sent to Thebes, to become the adoptive daughter of Nicotris I, God's Wife of Amun (the leader of the Egyptian Gods)and ruler of the city, adopting the title of Divine Adoratrice of Amun (High Priestess of the Cult of Amun). Nictrotis was a daughter of  Psamtik I, first Pharaoh of the 26th Dynasty and Psamtik II's grandfather, and as God's Wife of Amun ruled Thebes from 655 BC until her death in 585 BC, when Ankhnesneferibra succeeded her, becoming both Divine Adoratrice and God's Wife of Amun, until 560 BC, when Nicotris II, the daughter of Amasis II (fifth pharaoh of the 26th Dynasty and grandson of Psamtik II), became Divine Adoratrice of Amun, and adoptive daughter of Ankhnesneferibra.

Ankhnesneferibra ruled in Thebes from 585 until 525 BC, when Egypt was Invaded by the Achaemenid Persians under the Emperor Cambyses II, bringing the 26th Dynasty to an end. Cambysis re-organised the administration of Egypt significantly, incorporating it into his empire as a province, and abolished the roles of Divine Adoratrice of Amun and God's Wife of Amun (unfortunate for Nicrotis II, who by this time had been waiting for the role for 35 years). 

Statue  of Ankhenesneferibre, God's Wife of Amun, in the collection of the Nubian Museum of Aswan. John Campana/Flikr/Wikimedia Commons.

The title of God's Wife is a curious one. It originally applied to the wives and mothers of Pharaoh's (who were seen as living embodiments of the gods), with no specific god being associated with the post. This appears to have been modified to God's Wife of Amun in the New Kingdom, with little initial alteration of the role. The role as understood in the time of Ankhnesneferibra appears to have been created by Ramasses VI, the fifth Pharaoh of the 20th Dynasty, who gave the titles of God's Wife of Amun and Divine Adoratrice of Amun to his daughter Iset, with the intention that she would remain an unmarried virgin and adopt the daughter of the next Pharoah as her daughter and heiress. 

The God's Wife of Amun became the effective ruler of Thebes (ancient capital of Egypt) under Shepenupet I, the daughter of the 23rd Dynasty Pharoah Osorkon III, who adopted Amenirdis I, daughter of the Kushite invader Kashta as her daughter and Divine Adoratrice, thus securing her own political power and Kashta's claim to be a legitimate Pharaoh, ruler of Egypt, and founder of the 25th Dynasty at the same time. Kashta faced a rival claim to the throne from Tefnakht I, founder of the 24th Dynasty, which was also based in Sais, after the Kushites had conquered most of Egypt, including the traditional capital, Thebes.

For the Pharaohs of the 26th Dynasty the role of God's Wife of Amun was clearly an important one, and signficant to their claim to be legitimate rulers. The first Pharaoh of the 26th Dynasty, Psamtik I, was installed as a client Pharaoh at Sais by the Assyrian King Ashurbanipal, following the Assyrian invasion of Egypt, but secured independence when the Assyrians were forced to withdraw to concentrate on defending the heartlands of their empire against the Babylonians. A capable military leader, Psamtik I was able to drive out the Kushite Pharaoh's of the 25th Dynasty, smashing many of the monuments associated with their reign to assert his status as legitimate Pharaoh of Egypt. 

At this time, Shepenupet II, daughter of the Kushite Pharaoh Piye (son of Kashta) was the God's Wife of Amun, and her niece Amenirdis II was Divine Adoratrice. Psamtik I deposed Amenirdis II, installing his daughter Nicotris I as adoptive daughter of Shepenupet II, Divine Adoratrice and heir to the title of God's Wife, but left Shepenupet II in place as God's Wife and ruler of Thebes, like Kashta before him using this act to legitimise his own reign.