Taforalt or Grotte des Pigeons is a cave in the province of
Berkane
Berkane ( ber, translit=Berkane, ⴱⴻⵔⴽⴰⵏ) is a city in northeastern Morocco, in the tribal area of Aït Iznasen, limited by the Mediterranean to the north, the Kis river (Moroccan-Algerian border) and Oujda Province in the east, Nador ...
,
Aït Iznasen region,
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
, possibly the oldest
cemetery
A cemetery, burial ground, gravesite or graveyard is a place where the remains of dead people are buried or otherwise interred. The word ''cemetery'' (from Greek , "sleeping place") implies that the land is specifically designated as a buri ...
in North Africa (Humphrey ''et al.'' 2012). It contained at least 34
Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
adolescent and adult human skeletons, as well as younger ones, from the Upper Palaeolithic between 15,100 and 14,000
calendar
A calendar is a system of organizing days. This is done by giving names to periods of time, typically days, weeks, months and years. A date is the designation of a single and specific day within such a system. A calendar is also a physi ...
years ago. There is archaeological evidence for Iberomaurusian occupation at the site between 23,200 and 12,600 calendar years ago, as well as evidence for
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
occupation as old as 85,000 years.
Site description
La Grotte des Pigeons is a cave in eastern
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
near the village of Taforalt. Human occupation and natural processes in the cave have produced a thick sequence of archaeological layers dating between at least 85,000 and 10,000 years ago.
These occupation layers include pre-
Mousterian
The Mousterian (or Mode III) is an archaeological industry of stone tools, associated primarily with the Neanderthals in Europe, and to the earliest anatomically modern humans in North Africa and West Asia. The Mousterian largely defines the latt ...
,
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
, and
Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
lithic industries, plus an unusual non-Levallois industry between the Aterian and the Iberomaurusian dating to c. 24,500 cal BP. These industries date from the
Middle Stone Age
The Middle Stone Age (or MSA) was a period of African prehistory between the Early Stone Age and the Late Stone Age. It is generally considered to have begun around 280,000 years ago and ended around 50–25,000 years ago. The beginnings of pa ...
and the
Later Stone Age
The Later Stone Age (LSA) is a period in African prehistory that follows the Middle Stone Age.
The Later Stone Age is associated with the advent of modern human behavior in Africa, although definitions of this concept and means of studying it ar ...
.
Excavations of the Iberomaurusian layers dating from 15,100 to 14,000 years ago have recovered dozens of burials with some showing evidence of postmortem processing. Some show potential rituals with burials containing animal remains including horns, mandibles, a hoof, and a tooth.
The deep and highly stratified cave floor has yielded hearths, lithics, and shell beads, among a variety of artefacts of varying ages. The dryness of the cave has contributed to the notable level of preservation found among the remains and artefacts.
Geography
![Taforalt landscape](https://upload.wikimedia.org/wikipedia/commons/7/76/Taforalt_landscape.jpg)
The site is located around steep hills, rocky mountains, and the natural vegetation of the thermo-Mediterranean biozone including ''
Tetraclinis articulate'' and ''
Pinus halenpensis''. The area itself is located in the Eastern part of
Morocco
Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to ...
near the community of Taforalt (Tafoughalt) at (34°48′38″ N, 2°24′30″ W). The large mouth of the cave opens to the northeast and has an area > . Today the site lies around from the Mediterranean coast and at an altitude of above sea level.
Culture
The earliest layers of human habitation in the cave, dating from 85,000-82,000 years ago, contain evidence of a
pre-Mousterian
The Mousterian (or Mode III) is an archaeological industry of stone tools, associated primarily with the Neanderthals in Europe, and to the earliest anatomically modern humans in North Africa and West Asia. The Mousterian largely defines the lat ...
industry where no evidence of the
Levallois lithic technology is apparent. The following (newer) layers contain side scrapers, small radial Levallois cores, and thin, bifacially worked foliate points typical of the
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
technological industries.
These
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
layers were dated to come from approximately 32,000 to >40,000 years ago,
though other research has found a non-Levallois industry continuing at the site until 25,000 years ago.
By about 21,000 years ago, the
Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
industry marked by
microlith
A microlith is a small Rock (geology), stone tool usually made of flint or chert and typically a centimetre or so in length and half a centimetre wide. They were made by humans from around 35,000 to 3,000 years ago, across Europe, Africa, Asia an ...
ic backed bladelets became the dominant archaeological material, which has been found at the site. These Iberomaurusian layers contain microlithics, ostrich egg shells, potentially ritualized primary and secondary burials, and a notable increase in land snail remains indicating a shift in dietary practices.
Excavation history
The cave was discovered in 1908 and was excavated in 1944–1947, 1950–1955, 1969-1977, and 2003-2018. Much of the field records from the early excavations have been lost.
In 1951, Roche's team discovered human remains associated with the
Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
.
The Roche excavation encountered 10 metres of archaeological deposits with the
Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
occupying the top . This same stratification has been encountered in the subsequent excavations in other parts of the cave. Because of the dozens of skeletons located by Roche in the 1950s and the burials located during the Bouzouggar, Barton, and Humphrey excavations taking place since 2003, Grotte des Pigeons represents what is likely the earliest and most extensively used known prehistoric cemetery in
North Africa
North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
.
Stratigraphy
![Taforalt stratigraphy](https://upload.wikimedia.org/wikipedia/commons/e/ed/Taforalt_stratigraphy.jpg)
The
stratigraphy
Stratigraphy is a branch of geology concerned with the study of rock (geology), rock layers (Stratum, strata) and layering (stratification). It is primarily used in the study of sedimentary rock, sedimentary and layered volcanic rocks.
Stratigrap ...
in Grotte des Pigeons, going as deep as as in the case of Roche’s excavations, differs slightly throughout the cave but follows a simple pattern based on their colour: the Grey Series overlies the Yellow Series.
The Yellow series goes from the beginning of the occupation of the cave about 85,000 years ago to c. 15,000 cal BP. The overlying Grey Series dates from c. 15,000 to 12,500 cal BP ago, and hence accumulated rapidly in some 2500 years. The Grey Series, associated with the later Iberomaurusian, is characterized by extensive hearths and charcoal deposits (hence its colour), along with all of the site's burials. The Yellow Series is associated with the earlier Iberomaurusian, as well as with
Levallois artefacts of the
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
industry. The increased density of artefacts and evidence of food production in the Grey Series is seen as a sign of year-round occupation at the site whereas the Yellow Series is seen as evidence of seasonal habitation with occasional periods without humans.
There is a theorized 2,000 year gap of habitation between 18,000 and 20,000 uncal BP with this sterile layer being noted in Sector 8 of Barton’s excavations,
though other excavations near the mouth of the cave challenge this finding.
Dating
With 67 radiocarbon dates, Taforalt is the most extensively dated site of the North African Later Stone Age. Starting in the 1960s, it has been dated with both conventional and
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Educational institutions
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radiocarbon dating
Radiocarbon dating (also referred to as carbon dating or carbon-14 dating) is a method for determining the age of an object containing organic material by using the properties of radiocarbon, a radioactive isotope of carbon.
The method was dev ...
,
OSL,
TL, and
U-series.
Looking at all dates recovered from excavations, the habitation dates in this cave stretch from 12,500 cal BP ago to 85,000 years ago with a shift to
sedentary
Sedentary lifestyle is a lifestyle type, in which one is physically inactive and does little or no physical movement and or exercise. A person living a sedentary lifestyle is often sitting or lying down while engaged in an activity like soci ...
habitation about 15,000 cal BP.
The local environmental data helps establish the
seasonality
In time series data, seasonality is the presence of variations that occur at specific regular intervals less than a year, such as weekly, monthly, or quarterly. Seasonality may be caused by various factors, such as weather, vacation, and holidays a ...
of the site as much of the modern vegetation was utilized by the prehistoric population and follows a set seasonal process of food production. The presence of plant remains that would have been harvested in spring indicate that the cave or nearby environs were inhabited during that season. Proxies for environmental conditions during the phases of cave occupation are available from both wood charcoal and small mammal evidence. A feature of considerable interest in the charcoal record concerns the fluctuating presence of cedar in the C–F sequence. ''
Cedrus
''Cedrus'', common English name cedar, is a genus of coniferous trees in the plant family Pinaceae (subfamily Abietoideae). They are native plant, native to the mountains of the western Himalayas and the Mediterranean region, occurring at altit ...
'' currently grows in Morocco only from ≈1,300–2,600 m in the
Rif
The Rif or Riff (, ), also called Rif Mountains, is a geographic region in northern Morocco. This mountainous and fertile area is bordered by Cape Spartel and Tangier to the west, by Berkane and the Moulouya River to the east, by the Mediterrane ...
, the
Middle Atlas
The Middle Atlas (Amazigh: ⴰⵟⵍⴰⵙ ⴰⵏⴰⵎⵎⴰⵙ, ''Atlas Anammas'', Arabic: الأطلس المتوسط, ''al-Aṭlas al-Mutawassiṭ'') is a mountain range in Morocco. It is part of the Atlas mountain range, a mountainous region ...
, and
Eastern High Atlas, and its presence throughout the Taforalt record highlights a significant vegetation shift since the
Holocene
The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
. In particular, the vegetation excavated by Barton in Group E is dominated by the presence of ''
Cedrus atlantica
''Cedrus atlantica'', the Atlas cedar, is a species of tree in the pine family Pinaceae, native to the Rif and Atlas Mountains of Morocco (Middle Atlas, High Atlas), and to the Tell Atlas in Algeria.Gaussen, H. (1964). Genre ''Cedrus''. Les F ...
'' and deciduous ''
Quercus
An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...
'', with the latter declining at the expense of ''
Cedrus
''Cedrus'', common English name cedar, is a genus of coniferous trees in the plant family Pinaceae (subfamily Abietoideae). They are native plant, native to the mountains of the western Himalayas and the Mediterranean region, occurring at altit ...
''. This is consistent with environmental cooling and drying that comes with a change to a
montane
Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures fall as elevation increases, causing the ecosystem to stratify. This stratification is a crucial f ...
climate. This climatic shift coincides with the dates recovered from Group E and validates the dates recovered there.
Archaeological finds
Artefacts
The lithic collections recovered from the excavations at Grotte des Pigeons reflect a wide range of technologies and include unretouched and retouched flakes and bladelets, single and opposed platform bladelet cores, river cobbles,
microburin
A microburin is a characteristic waste product from manufacture of lithic tools — sometimes confused with an authentic burin — which is characteristic of the Mesolithic, but which has been recorded from the end of the Upper Paleolithic until ...
s, La Mouillah points, backed bladelets, Ouchtata bladelets, obtuse-ended backed bladelets, side scrapers, large bifacial tools, shell beads associated with bifacial foliates and tanged tools associated with the
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
culture, and potential rock
palettes.
Faunal remains
Animal remains found at the site largely appear to be
food waste
Food loss and waste is food that is not eaten. The causes of food waste or loss are numerous and occur throughout the food system, during production, processing, distribution, retail and food service sales, and consumption. Overall, about o ...
though excavations in the 1950s and 2000s, 2010s have revealed burials associated with
antelope
The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia.
Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mammals ...
horns,
bovine
Bovines (subfamily Bovinae) comprise a diverse group of 10 genera of medium to large-sized ungulates, including cattle, bison, African buffalo, water buffalos, and the four-horned and spiral-horned antelopes. The evolutionary relationship betwee ...
horns, and at least one
horse
The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million y ...
tooth.
The more sedentary Grey Series phase includes a substantial amount of land
Mollusca
Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is esti ...
remains in conjunction with hearths indicating extensive land snail collection and cooking.
The earliest layers from approximately 80,000 years ago contain shell beads of the ''N. gibbosulus'' however analysis of these shells indicate that they were collected along the Mediterranean shore after they had been dead.
Ash lenses from the Aterian levels around 80,000 BP contain large ''
Otala punctate'' indicating small scale exploitation of land snails prior to the Grey Series.
Floral remains
The vegetation species found inside the cave provide an idea what the environment was like during periods of human habitation with the charred remains of Holm oak (''
Quercus ilex
''Quercus ilex'', the evergreen oak, holly oak or holm oak is a large evergreen oak native to the Mediterranean region. It is a member of the ''Ilex'' section of the genus, with acorns that mature in a single summer.
Description
An evergreen tr ...
'' L.) acorns, Maritime pine (''
Pinus pinaster
''Pinus pinaster'', the maritime pine or cluster pine, is a pine native to the south Atlantic Europe region and parts of the western Mediterranean. It is a hard, fast growing pine bearing small seeds with large wings.
Description
''Pinus pinast ...
'' Aiton) pine nuts, Juniper (''
Juniperus phoenicea
''Juniperus phoenicea'', the Phoenicean juniper or Arâr, is a juniper found throughout the Mediterranean region.
Description
''Juniperus phoenicea'' is a large shrub or small tree reaching tall, with a trunk up to in diameter and a rounded o ...
'' L.), Terebinth pistachio (''
Pistacia terebinthus
''Pistacia terebinthus'' also called the terebinth and the turpentine tree, is a deciduous tree species of the genus ''Pistacia'', native to the Mediterranean region from the western regions of Morocco and Portugal to Greece and western and s ...
'' L.), and wild oat (''
Avena
''Avena'' is a genus of Eurasian and African plants in the grass family. Collectively known as the oats, they include some species which have been cultivated for thousands of years as a food source for humans and livestock. They are widesprea ...
'' sp.) being recovered after likely being collected and processed by the previous inhabitants.
Human remains
Sector 10, excavated by Humphrey, and the burial deposits excavated by Roche in the 1950s, form a contiguous and spatially demarcated collective burial area with dozens of closely spaced burials. The presence of both articulated and disarticulated bones indicates extensive use and reuse of the burial area with evidence of secondary burial and selective bone removal being practiced, often disturbing or truncating earlier burials. Some burials were covered by large stones preventing future disturbances by burials.
The Roche excavations originally estimated that they had recovered the remains of approximately 180 individuals, but subsequent research adjusted this estimate to between 35 and 40 individuals.
These remains were not directly dated by Roche but based on the
stratigraphy
Stratigraphy is a branch of geology concerned with the study of rock (geology), rock layers (Stratum, strata) and layering (stratification). It is primarily used in the study of sedimentary rock, sedimentary and layered volcanic rocks.
Stratigrap ...
they were from a greater depth, and therefore greater age, than those in Sector 10. The recent excavations taking place in Sector 10 have recovered thirteen partially articulated skeletons along with a sample of disarticulated bones. Seven bone samples from Sector 10 yielded age estimates between approximately 15,077 and 13,892 years ago, corresponding to the base of the Grey Series deposits seen in Sector 8 excavations.
Burials situated toward the front of the cave and those higher within the deposits are likely to be progressively younger, and hence contemporary with higher levels in the Grey Series deposits recorded in Sector 8. A range of funerary practices is apparent based on the grave excavations that have taken place. Some remains appear to have been primary
inhumations
Burial, also known as interment or inhumation, is a method of final disposition whereby a dead body is placed into the ground, sometimes with objects. This is usually accomplished by excavating a pit or trench, placing the deceased and objec ...
while others appear to have sustained secondary
inhumation
Burial, also known as interment or inhumation, is a method of final disposition whereby a dead body is placed into the ground, sometimes with objects. This is usually accomplished by excavating a pit or trench, placing the deceased and objec ...
after removal for potentially ritual practices.
Evidence of deliberate
post-mortem
An autopsy (post-mortem examination, obduction, necropsy, or autopsia cadaverum) is a surgical procedure that consists of a thorough examination of a corpse by dissection to determine the cause, mode, and manner of death or to evaluate any dis ...
modification include cut marks that are not indicative of cannibalism and extensive
ochre
Ochre ( ; , ), or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colours produced ...
colouring with one grave, Grave XII, containing Individual 1 with both cut marks and
ochre
Ochre ( ; , ), or ocher in American English, is a natural clay earth pigment, a mixture of ferric oxide and varying amounts of clay and sand. It ranges in colour from yellow to deep orange or brown. It is also the name of the colours produced ...
colouring present on the majority of the nearly intact skeleton.
Roche’s excavations in the 1950s yielded a single
mandible
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
from the
Aterian
The Aterian is a Middle Stone Age (or Middle Palaeolithic) stone tool industry centered in North Africa, from Mauritania to Egypt, but also possibly found in Oman and the Thar Desert. The earliest Aterian dates to c. 150,000 years ago, at the sit ...
levels.
A 2003 analysis
of
masticatory
Chewing or mastication is the process by which food is crushed and ground by teeth. It is the first step of digestion, and it increases the surface area of foods to allow a more efficient break down by enzymes. During the mastication process, th ...
and non-masticatory dental modifications among the remains recovered in the 1950s reflected a very high rate (90%)
of
avulsion of the upper central
incisor
Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
s which subsequently led to increased usage of the proximal teeth. Ritual tooth removal is known elsewhere in this region at other points in prehistory and history and likely took place during the entrance to adulthood. The food processing tasks of the teeth are reflected in the heavy chipping, perhaps indicative of a gritty diet involving bone and shell. Half of the surviving teeth (51.2%) exhibited
carious lesions while archaeological hunter-gatherers are expected to range between 0% – 14.3% and agriculturalists range between 2.2% - 48.1%. These numbers are likely a result of the acorns and pine nuts which would have been collected and processed, resulting in fermentable carbohydrates. The women in the population do not reflect the same proximal tooth wear as their upper central incisors were typically not removed.
A 2000 analysis of non-metric dental traits indicated genetic continuity from the terminal
Pleistocene
The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
onward in the Iberomaurusian and
Capsian
The Capsian culture was a Mesolithic and Neolithic culture centered in the Maghreb that lasted from about 8,000 to 2,700 BC. It was named after the town of Gafsa in Tunisia, which was known as Capsa in Roman times.
Capsian industry was concentra ...
areas.
Occupation site utility
The inhabitants of Grotte des Pigeons were hunter-gathers equipped with the knowledge of harvesting plants and animals as the archaeological context suggests some of the burials contained evidence of baskets and grind stones which were used for food preparation.
Some of the foods harvested from their local environment included acorns, pine nuts, and land molluscs.
The site exhibits evidence that the people that lived in this area used the cave year round by the Grey Series while staying there seasonally during the Yellow Series.
The perforated marine shells present from the 85,000 – 82,000 year old level at Grotte des Pigeons and other sites in the nearby Maghreb dated from that period reflect an exchange network that likely existed in order to provide shells to communities 40 km from the coast (Taforalt) and further.
While the meaning behind the beads cannot be discerned, the presence of an apparently widespread exchange network to facilitate their transport as well as their being worked for apparent ornamentation indicate some significance behind them.
World Heritage status
This site was added to the
UNESCO
The United Nations Educational, Scientific and Cultural Organization is a specialized agency of the United Nations (UN) aimed at promoting world peace and security through international cooperation in education, arts, sciences and culture. It ...
World Heritage
A World Heritage Site is a landmark or area with legal protection by an international convention administered by the United Nations Educational, Scientific and Cultural Organization (UNESCO). World Heritage Sites are designated by UNESCO for h ...
Tentative List on July 1, 1995 in the Cultural category under the name "Grotte de Taforalt".
[Grotte de Taforalt - UNESCO World Heritage Centre](_blank)
/ref>
Ancient DNA
In 2018, van de Loosdrecht et al. performed the first aDNA tests on the ancient Taforalt individuals, directly dated to between 15,100 and 13,900 cal BP. The Taforalt samples are the oldest human DNA samples from Africa yet recovered. DNA analysis was performed on seven individuals: six males and one female. Only five of individuals, including four of the males, with higher coverage genomes were used in the nuclear DNA analysis. Nuclear DNA analysis reveals that the Taforalt individuals were all closely related to each other, showing evidence of a population bottleneck
A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population due to environmental events such as famines, earthquakes, floods, fires, disease, and droughts; or human activities such as specicide, widespread violen ...
event in their past."
The Taforalt genomes were found to be composed of three major components: a Holocene
The Holocene ( ) is the current geological epoch. It began approximately 11,650 cal years Before Present (), after the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene togethe ...
West-Eurasian/Levant
The Levant () is an approximate historical geographical term referring to a large area in the Eastern Mediterranean region of Western Asia. In its narrowest sense, which is in use today in archaeology and other cultural contexts, it is eq ...
ine component, a Hadza hunter-gatherer component from Tanzania
Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
, and a West Africa
West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Mali, Maurit ...
n component. The Taforalt individuals shows the relative closest genetic affinity for ancient Epipaleolithic Natufian
The Natufian culture () is a Late Epipaleolithic archaeological culture of the Levant, dating to around 15,000 to 11,500 years ago. The culture was unusual in that it supported a sedentary or semi-sedentary population even before the introduction ...
individuals, with slightly greater affinity for the Natufians than later Neolithic
The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several parts ...
Levantines. A two-way admixture scenario using Holocene Levantines and modern West African samples as reference populations inferred that the Taforalt individuals bore 63.5% Levantine-related and 36.5% Sub-Saharan African-related ancestries, with no evidence for additional gene flow from the Epigravettian
The Epigravettian (Greek: ''epi'' "above, on top of", and Gravettian) was one of the last archaeological industries and cultures of the European Upper Paleolithic. It emerged after the Last Glacial Maximum around ~21,000 cal. BP or 19,050 BC, a ...
culture of Upper Paleolithic
The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coin ...
Europe. The Taforalt individuals also show evidence of limited Neanderthal
Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ago. While th ...
ancestry.
When compared against modern populations, the Taforalt individuals form a distinct cluster and do not cluster genetically with any modern population; however, they were found to cluster between Middle Easterners or modern North Africans and West/East Africans. The Taforalt individuals also exhibit higher levels of indigenous African ("Sub-Saharan African") ancestry than do modern North Africans. The Sub-Saharan African DNA in Taforalt individuals is best samplified, most of all, by modern West Africans (e.g., Yoruba, or Mende). Additionally, the Taforalt samples of the Iberomaurusian
The Iberomaurusian is a backed bladelet lithic industry found near the coasts of Morocco, Algeria, and Tunisia. It is also known from a single major site in Libya, the Haua Fteah, where the industry is locally known as the Eastern Oranian.The " ...
culture also shares partial similarities with the remnants of a more basal Sub-Saharan African lineage (e.g., basal to modern West and East African lineages), suggesting the African component derives from a population prior or close to the split between West and Eastern indigenous Africans.
A 2018 DNA analysis shows that the Taforalt individuals belonged to the mtDNA haplogroup
In human genetics, a human mitochondrial DNA haplogroup is a haplogroup defined by differences in human mitochondrial DNA. Haplogroups are used to represent the major branch points on the mitochondrial phylogenetic tree. Understanding the evolu ...
s U6a and M1b, as well as to the Y-DNA haplogroup
In human genetics, a human Y-chromosome DNA haplogroup is a haplogroup defined by mutations in the non- recombining portions of DNA from the male-specific Y chromosome (called Y-DNA). Many people within a haplogroup share similar numbers of sh ...
E1b1b1a1 (M78), which is closely related to the E1b1b1b (M123) sublineage that has been observed in skeletal remains belonging to the Epipaleolithic Natufian
The Natufian culture () is a Late Epipaleolithic archaeological culture of the Levant, dating to around 15,000 to 11,500 years ago. The culture was unusual in that it supported a sedentary or semi-sedentary population even before the introduction ...
and Pre-Pottery Neolithic
The Pre-Pottery Neolithic (PPN) represents the early Neolithic in the Levantine and upper Mesopotamian region of the Fertile Crescent, dating to years ago, (10000 – 6500 BCE).Richard, Suzanne ''Near Eastern archaeology'' Eisenbrauns; il ...
cultures of the Levant
The Levant () is an approximate historical geographical term referring to a large area in the Eastern Mediterranean region of Western Asia. In its narrowest sense, which is in use today in archaeology and other cultural contexts, it is eq ...
, possibly suggesting geneflow.
Since the Natufian samples, which are chronologically younger than the Taforalt samples by several thousands of years, were inferred to lack substantial African ancestry, the researchers also hypothesized that a Maghreb center of evolution for the Natufian-related ancestry could only be plausible if the admixture that was inferred for the Taforalt individuals either occurred after the population ancestral to the Natufians had moved into the Levant or if that admixture event was a locally-confined phenomenon at Taforalt.
Phenotypic analysis was performed on four of the Taforalt individuals with higher genomic coverage. The Taforalt individuals tested did not carry either of the derived SLC24A5
SLC may refer to:
Places
* Salt Lake City, Utah
* Salt Lake City International Airport, IATA Airport Code
Education
* Sarah Lawrence College, NY
* School Leaving Certificate (Nepal)
* St. Lawrence College, Ontario, Canada
* Small Learning Comm ...
alleles associated with lighter skin color, the derived OCA2
P protein, also known as melanocyte-specific transporter protein or pink-eyed dilution protein homolog, is a protein that in humans is encoded by the oculocutaneous albinism II (''OCA2'') gene. The P protein is believed to be an integral membrane ...
allele associated with blue eye
Eye color is a polygenic phenotypic character determined by two distinct factors: the pigmentation of the eye's iris and the frequency-dependence of the scattering of light by the turbid medium in the stroma of the iris.
In humans, the pi ...
color, or the derived MCM6
DNA replication licensing factor MCM6 is a protein that in humans is encoded by the ''MCM6'' gene. MCM6 is one of the highly conserved mini-chromosome maintenance proteins ( MCM) that are essential for the initiation of eukaryotic genome replicati ...
allele associated with lactase persistence
Lactase persistence is the continued activity of the lactase enzyme in adulthood, allowing the digestion of lactose in milk. In most mammals, the activity of the enzyme is dramatically reduced after weaning. In some human populations, though, lact ...
. However, they were found to carry the ancestral SLC24A4
Sodium/potassium/calcium exchanger 4 also known as solute carrier family 24 member 4 is a protein that in humans is encoded by the ''SLC24A4'' gene.
Clinical effects
Mutations in SLC24A4 cause amelogenesis imperfecta.
References
{{reflist
...
allele associated with dark eye color, suggesting that the West-Eurasian migrant group may not have evolved light skin yet.
See also
*Ifri n'Amr or Moussa Ifri n'Amr Ou Moussa is an archaeological site discovered in 2005, located in the rural commune of Aït Siberne, Khémisset Province, in Western Morocco. This site has revealed burials associated with both Moroccan Early Neolithic and Bell Beaker ...
*Kelif el Boroud Kehf el Baroud, sometimes mistakenly spelled Kelif el Boroud, is an archaeological site in Morocco. It is located to the south of Rabat, near Dar es Soltan.
Genetics
examined the remains of 8 individuals buried at Kelif el Boroud c. 3780-3650 BC ...
*Mechta-Afalou
Mechta-Afalou (Mechtoid) or Paleo-Berber are a population that inhabited parts of North Africa during the late Paleolithic and Mesolithic. They are associated with the Iberomaurusian archaeological culture.
Mechtoids are believed to have been assi ...
References
External links
Grotte des Pigeons
Grotte de Taforalt - UNESCO World Heritage Centre
{{Navbox prehistoric caves
Archaeological sites in Morocco
Caves of Morocco