In an early post on this blog, I summarized my maternal-line ancestors and where and when they lived. In the last 6 years, there have been substantial revisions to estimates of the dates associated with these mitochondrial DNA (mtDNA) haplogroup founders, and revisions to the mtDNA haplogroup tree (deep-maternal-ancestry-and-mtdna) and this post provides an update. I am a member of mtDNA haplogroup U5, which is one of nine native European haplogroups stemming from haplogroup U which most likely arose in the Near East, and spread into Europe in a very early expansion. The presence of haplogroup U5 in Europe pre-dates the last ice age and the expansion of agriculture in Europe. Today, about 11% of modern Europeans are the direct maternal descendants of the founder U5 woman. They are particularly well represented in western Britain and Scandinavia. My more recent maternal ancestors were part of the population that tracked the retreat of ice sheets from Europe at the end of the last ice age and re-colonized Britain about 12,000 years ago.
The mtDNA sequence at the root of each haplogroup arose from one or more mutations in the mtDNA of just one woman, and the age of the associated haplogroup gives the time in the past when this specific woman lived. To emphasise that the maternal clan founders were real individuals, I have used the names given to them by Sykes  and Oppenheimer  and given my own names to the more recent subgroup founders. The Table below summarizes these founders, dates and locations and is followed by brief biographies. The haplogroups are identified by the labels used in Build 17 of the ISOGG mtDNA tree which can be accessed at http://phylotree.org/ . Dates in the table below have been updated using most recent available dating estimates as described in my previous post deep-maternal-ancestry-and-mtdna.
The migration path out of Africa into Europe of the “grandmothers” linking mitochondrial Eve through to Ursula (U5) is shown on a map in my previous post deep-maternal-ancestry-and-mtdna. The subsequent migration from Europe to Britain is shown in the map below.
Updated biographies of my maternal haplogroup great* grandmothers follow below.
Mitochondrial Eve (haplogroup L1)
Mitochondrial Eve, my great*7,075th grandmother, lived approximately 177,300 years ago in southern Africa. Her haplogroup L1 is ancestral to all mitochondrial haplogroups found today. Mitochondrial Eve was not the first woman, nor was she the only woman alive at the time. Her contemporaries could still have descendants, whose line zig-zagged back and forth between males and females, but Eve was the only one who had an unbroken line of daughters, through thousands of generations, right down to the present time. She is the maternal ancestor of all humans alive today.
Mitochondrial Eve is estimated to have lived around 178,000 years ago (with 90% uncertainty range 159,400 to 196,600 years), when Homo sapiens sapiens (anatomically modern humans) were developing as a population distinct from other human sub-species in Africa (during the period from 250,000 to 150,000 years ago). She lived later than Homo heidelbergensis from whom Homo sapiens split around 250,000 (±54,000) years ago but earlier than the period when modern humans left Africa. She lived much later than the earlier “Eve” ancestral to both modern humans and Neanderthals, who split from Homo heidelbergensis around 550,000 (±54,000) years ago, probably in Europe.
A recent paper by Chan et al in Nature  identified Eve’s homeland as Makgadikgadi, a vast wetland in northern Botswana. I discussed this finding in more detail in a recent post (mitochondrial-eve-an-update). Haplogroup L1 is today found in West and Central sub-Saharan Africa. Haplogroup L1 arose with Mitochondrial Eve and haplogroup L0 is an offshoot. African haplogroups L2 and L3, the latter of which gave rise to all non-African haplogroups, are also descendants of L1.
Lara (Haplogroup L3)
Sykes  gave the name Lara to the ancestral mother of the L3 haplogroup. She lived approximately 71,900 years ago in Africa (with uncertainty range 64,600 to 79,100 years), which would make her my great*2,860th grandmother. L3 is the haplogroup from which the haplogroups M and N are believed to have arisen. These latter two haplogroups are ancestral to all haplogroups outside Africa and are believed to represent the initial successful migration by modern humans out of Africa.
Today, haplogroup L3 is confined to Africa and emigrant African populations. It is most common in East Africa and Lara likely lived in Ethiopia. It has been estimated that from a population of 2,000 to 5,000 individuals in Africa, only a small group, possibly as few as 150 to 1,000 people, crossed the Red Sea around 70,000 years ago, and these were all descendants of Lara. Today at the Bab-el-Mandeb straits, the Red Sea is about 20 kilometres wide, but 70,000 years ago sea levels were 70 meters lower (owing to glaciation) and the water was much narrower. Though the straits were never completely closed, there may have been islands in between which could be reached using simple rafts.
An earlier migration by humans out of Africa occurred around 125,000 years ago. During a warm, moist period of Earth’s history when the Sahara was green grasslands, the first migration of humans from Africa saw them reach the Levant, the region of the Middle East along the Eastern Mediterranean Sea. But the Earth’s climate again shifted and the Sahara and Levant became desert. This group of humans did not survive and they left no modern descendants.
Nasreen (Haplogroup N, Eurasia)
Of all the lineages present in Africa only the female descendants of one lineage, mtDNA haplogroup L3, are found outside Africa. Had there been several migrations one would expect descendants of more than one lineage to be found outside Africa. L3’s female descendants, the M and N haplogroup lineages, are found in very low frequencies in Africa and appear to be recent arrivals. The founder of the N haplogroup, Nasreen, lived in the Arabian Pensinsula within a few thousand years after the exodus from Africa. Nasreen has been dated to 62,800 (59,000-66,700 years) ago.
Rohani and Rian (Haplogroup R, West Eurasia)
Oppenheimer  named the South Asian ancestral mother of the R haplogroup as Rohani who lived 60,300 (56,900-63,700) years ago. The period from 74,000 to 58,000 years ago was a period of intense cold in the Northern Hemisphere and the Europeans’ ancestors were unable to move back into the Levant from Asia for over 10,000 years. During this period, a later West Eurasian ancestral mother for haplogroup R has been identified by Soares et al . She has not been given a name by Sykes or Oppenheimer, and I am calling her Rian (after the great-grandmother of Elrond). She lived 59,100 (47,100-71,400) years ago, which makes her my great*2,350th grandmother. Although Rian is not the founder of the R haplogroup per se, she is the most recent common ancestor for all descendents of the R haplogroup in Europe.
Both the N (‘Nasreen’) and Rohani haplogroups are unknown except in South Asia, where Nasreen root types are found at low rates and Rohani is found in great variety. Most Rohani types in India are found nowhere else, although two very old subtypes have been found in China. While recent analyses place Rohani in West Asia (perhaps Pakistan), it is possible that she lived further south in India (South Asia).
Europa (Haplogroup U)
Haplogroup U originated some 50,000 years ago in the Middle East and gave rise to all the native European haplogroups. For this reason, Oppenheimer gave the name Europa to the ancestral mother of haplogroup U. Europa lived around 49,500 (44,100-54,900) years ago, making her my great*1,970th grandmother.
Dramatic warming of the climate in the period 58,000 to 48,000 years ago meant groups were finally able to move north up the Fertile Crescent returning to the Levant. From there they moved into Europe via the Bosporus from 50,000 years ago. Given the uncertainties involved in dating the mtDNA mutations, the move into Europe can be more accurately dated by looking at the climatic opportunities .
From 56,000 to 50,000 years ago, there followed in quick succession a run of four warm and wet periods (see Figure 2). Apart from the opening of the Fertile Crescent corridor, dry areas of the Levant such as the Negev Desert became potentially habitable. Europa probably lived in or near Iran and her earliest daughter lines entered Europe 48,000 years ago. Haplogroup U appears to have lived during a period of rapid population growth and expansion because it has nine major surviving daughter groups, U1 through U9, which are now found among people who have ancestral origins throughout Europe, Asia, and Africa.
Several lineages followed the great herds of buffalo, antelope, woolly mammoths, and other animals from the Fertile Crescent region through what is now modern-day Iran to the vast steppes of Central Asia. These semi-arid grass-covered plains formed an ancient “superhighway” stretching from eastern France to Korea. Having migrated north out of Africa into the Middle East, these people then traveled both east and west along this Central Asian route. Others traveled westward into Europe. These Upper Paleolithic people, the Cro-Magnon, dominated the human expansion into Europe that eventually lead to the demise of the native Neanderthal population by around 30,000 years ago.
Recently, analysis of Neanderthal DNA has suggested they are closer to non-African than African anatomically modern human populations, which is probably due to interbreeding between Neanderthals and the ancestors of the Eurasians between roughly 70,000 and 50,000 years ago, shortly after (or perhaps before) they emigrated from Africa and while they were still one population. This resulted in 1–4% of the genome of people from Eurasia having been contributed by Neanderthals.
From around 45,000 years ago until 43,000 years ago, the climate became much colder, causing a mini-Ice Age in Europe. Beginning about 40,000 years ago the Earth’s climate shifted and became colder and more arid. This marked the beginning of the most recent Ice Age (the Würm glaciation of Switzerland and the Alps, and the Weichselian glaciation of northern Europe and Scandinavia) which lasted over 20,000 years. Drought hit Africa reverting the grasslands of Northern Africa to desert. For the next 20,000 years the Saharan Gateway between Africa and the Middle East was effectively closed.
Europa’s descendents, who had entered Europe in the window around 50,000 years ago, moved from Turkey into Bulgaria, and thence into Europe. These people developed a more sophisticated stone tool technology and their culture is referred to as Aurignacian. The earliest Aurignacian stone tools are found in Bulgaria and dated to around 45,000 years ago. The new style of stone tools moved up the Danube into Hungary then Austria. By 40,000 years ago, Aurignacian people had also reached Spain. These people were hunter-gatherers and moved into Europe long before the expansion of agriculture into Europe.
Ursula (Haplogroup U5)
The clan of Ursula (Latin for she-bear) is the oldest of the seven native European clans (the seven U haplogroups found in Europe, whose founders were dubbed the seven daughters of Eve by Sykes). Ursula is estimated to have lived 30,200 (21,500-39,000) years ago, making her my great*1200th grandmother. Ursula lived in Europe during the relatively warmer period of the Denekamp interstadial (32,000 to 28,000 years ago) . Summer temperatures were somewhat lower than the present day, but winters were markedly colder, with snow cover on the plains of Europe for three to six months of the year and much greater climate variability than today.
DNA tests on ancient skeletons have shown that U5 was the principal mitochondrial haplogroup of Paleolithic and Mesolithic hunter-gatherers in Northern Europe. U5 has been found in human remains dating from the Mesolithic in England, Germany, Lithuania, Poland, Portugal, Russia, Sweden, and France. Fu et al.  found two U5 individuals at the Dolni Vestonice burial site in the Czech Republic that has been dated to 31,155 years ago. A third person from the same burial was identified as haplogroup U8. The Dolni Vestonice samples have only two of the five mutations ( C16192T and C16270T) that are found in the present day U5 population. This indicates that the U5-(C16192T and C16270T) mtDNA sequence is ancestral to the present day U5 population that includes the additional three mutations T3197C, G9477A and T13617C .
Because present-day U5 is distinguished from U by five additional mutations, it must have experienced a long period of very slow population growth or a population bottleneck in Europe. Ancient DNA tests conducted in Britain, Germany and Scandinavia indicate that the frequency of U5 has progressively declined over time through the Neolithic, Bronze Age, Iron Age and Middle Ages. Nowadays it remains most common in the far north of Europe, where the Mesolithic population has been least affected by subsequent migrations.
According to Bryan Sykes , Ursula was likely born in the mountains of Greece near Delphi, close to the beginning of the Ice Age. At the time he was writing, Ursula was dated to around 45,000 years ago, the time when modern humans first moved into Europe. With the more recent dating to around 30,000 years ago it is likely that Ursula lived further west, perhaps even as far west as Spain. In any case, the world that Ursula was born into was a lot colder than it is today, and would get colder still in the following millennia. She was part of a semi-nomadic band who followed the deer or bison herds between their summer and winter grazing lands. Ursula and other women spent a lot of time gathering food: fruit, berries, nuts, roots, as well as small animals and bird eggs. Life was extremely hard, with high risk of injuries in hunting, and of being taken by predators such as lions and leopards, as well as the ever-present risk of starvation in bad years. The average life expectancy at that time is estimated to be around 35 years. However, life expectancy was lower for women than men, with few women surviving into their thirties.
Between around 30,000 and 26,000 years ago, modern humans reached Britain only to retreat again from the advancing ice. Because so much of the Earth’s water was trapped in ice, the sea’s level was about 127 m (417 ft.) lower than it is today. Consequently, Britain was joined to Ireland by an exposed “land bridge,” making transit between those regions more practical as boats were no longer needed for the journey. The lowered sea level also joined Britain to Continental Europe by an area of dry land, known today as Doggerland (which extended also to Norway). This meant that humans retreating from the ice would have been able to walk south into Europe. It is possible that Ursula lived in Britain or north-westerm Europe, but I have followed Rob Spencer  in showing her likely location as in the Balkans.Ursula probably gave birth to her first daughter around age 15 and she would have nursed the baby and carried it on her back while she foraged in the forest. Three or more years later, she had another baby daughter, and both daughters grew up strong and healthy and gave Ursula grand-daughters. Ursula probably died in her 30s, perhaps because she lost too many teeth to chew the tough food that was the staple between animal kills, or perhaps because she was unable to escape a predator, or in childbirth. The oldest known cave paintings in Europe date from the time of Ursula, the Aurignacian period (38,000 to 30,000 years ago), and these paintings often depict dangerous animals such as lions, bears, hyaenas, and woolly rhinoceroses, as well as humans, horses, and other food animals. Ursula’s time was in the period when humans developed art, music and personal decoration.
Ursula lived around the time that Neanderthals became extinct. Most recent remains found for Neanderthals in Spain are dated to 24 to 33,000 years ago . Possibly Ursula lived in the same region, and may have encountered some of the last remaining Neanderthals.
Ursula and her family likely encountered Neanderthals. Her people were slightly taller than Neanderthals and much slimmer, a trait that helped her ancestors adapt better to their previous homelands of the Middle East and Africa. Neanderthals were shorter and stockier, with large broad noses, built better for the colder weather of Europe where they resided for the past quarter million years. Eventually, with their new and sophisticated type of stone tools, and possibly better communication skills and social organization, they edged the Neanderthals into extinction.
Udala (Haplogroup U5a)
Haplogroup U5a is a lineage that branched off from U5 approximately 22,400 years ago (14,400—30,500) and is mostly distributed in southern Europe today. Sykes did not name the U5a founder, I have chosen the Basque name Udala and she would be approximately my great*880th grandmother. Not long before her time, humans intentionally began producing sculptures for the first time. The Venus of Willendorf (illustrated left) dates from around 24,000 years ago.
Udala lived during the period of the Last Glacial Maximum (LGM) when much of northern Europe was covered in thick ice sheets, and much of Europe south of the ice sheets was permafrost and unsuitable for human occupation. This period lasted from around 25,000 years ago to 13,000 years ago. During this period, the U5a and U5b subclades of U5 were forced south into ice age refugia in southern Europe, and perhaps Ukraine and the Near East.
Haplogroup U5 and its subclades U5a and U5b form the highest population concentrations in the far north, in Sami (Laplanders), Finns, and Estonians, but it is spread widely at lower levels throughout Europe. For example, Cheddar Man, the oldest remains of anatomically modern humans in Britain, was in Haplogroup U5a. The Cheddar Man is the nickname for the ancient human remains found in Cheddar Gorge; his approximate date of death was 7150 BCE (9162 years ago).
Malyarchuk et al.  reconstructed a detailed phylogenetic tree for U5 and concluded that expansions of U5 subclusters started earlier in central and southern Europe, than in eastern Europe. Although U5 is now ubiquitous in Europe, the oldest Europa great-granddaughter, U5a, is commonest in the Basque country of northern Spain. One of the few European refuges during the last ice age, the Basque region managed to preserve more of its original genetic diversity than did other parts of Western Europe. We can thus assume that Udala likely lived in the Basque region of South-West Europe or her descendents were forced into that region as Europe entered the Last Glacial Maximum period from around 23,000 to 16,500 years ago.
Urd (Haplogroup U5a1)
During the last ice age, Northern Europe was depopulated, with isolated surviving groups locked in southern refuges. Urd lived in the Basque refuge around 16,900 (12,300-21,600) years ago. Neither Sykes or Oppenheimer named the founders of U5a subclades, I have chosen the name Urd. Urd is one of the three Norns (the Fates of Scandinavian mythology), the Norn of the past. Urd is my great*940th grandmother.
The Last Glacial Maximum (LGM) covered much of Europe in thick ice sheets from 23,000 to 16,500 years ago. Much of Europe south of the ice sheets was permafrost and unsuitable for human occupation. There were three main refuge areas of Southern Europe to which the Palaeolithic peoples of Europe retreated. From west to east, the first consisted of parts of France and Spain either side of the Pyrenees, in the Basque country, characterized by the finely knapped stone ‘leaf points’ of the Solutrean culture (named after a French village called Solutre). Perhaps receiving their technologies from north-west Europe, this south-western refuge was culturally distinct from other southern refuges, whose stone technology is described more generally as Gravettian. The second refuge area was Italy, with more or less continuous local occupation. The third was the Ukraine, a large area north of the Black Sea defined by two great rivers, the Dnepr and the Don, and separated from the rest of Southern Europe by the Carpathian Mountains, which were partially glaciated at the LGM.
During the LGM, permafrost extended down to southern France, just north of Bordeaux and into the uplands of northern Provence. The areas just to the south of the main ice sheets had little or no vegetation. Only in the Dordogne region of southwest France and the foothills of the Pyrenees, where modern humans had lived for millennia before the LGM, is there widespread evidence of their having stuck it out during the LGM . This region was arid semi-desert with little thick woody vegetation. The main refugia for deciduous trees and pines were on the western side of the mountains in Greece.
The sudden changes in climate during this period were unlike anything we have experienced in recorded history. The sudden jumps associated with centuries-long fluctuations were typically in the range of 5 to 10 C in average temperatures. But decade to decade variations were much greater. These wild swings in climate would have required an extraordinarily adaptable and migratory lifestyle, as well as making any form of agriculture impossible.
While Neanderthals had obtained almost all their dietary protein from animal sources, the diet of modern humans had become extremely diverse by the end of the LGM. Their prey were principally gazelle, deer, fox and hare as well as many species of birds, and lots of fish. Fish and mollusks became an important component of their diet, and many people must have survived the ice age by living close to the seashore. Their diet also included a huge range of edible plants, nuts, berries and other fruits. More than 100 species of edible plant have been identified. There was increased use of milling tools after the LGM, and some evidence of storage facilities and preserved plant parts.
In short, Urd lived in the Basque region and probably was part of a group who moved seasonally between the sea shore in the winter and inland areas where game animals and edible plants were plentiful in the summer. Urd’s descendents in the Pyrenees over the next two or three thousand years would have been part of the Magdalenian culture in the foothills of the Pyrenees who produced the stunning cave paintings at sites such as Roc-de Sers, Lascaux, and Niaux. I have visited the Grotte de Niaux twice, once in 1992 and again in 2011 (grotte-de-niaux). It is one of the few caves with Paleolithic paintings that can still be visited.
The galleries with the paintings are 800 metres inside the cave. Visitors are provided with hand-held torches and walk through the horizontal cave system until they reach the gallery with the paintings. On both occasions that I went into the cave, there were gasps of amazement as the guide shone his torch onto the Paleolithic paintings of bison, horses, deer and other animals. Now, through radiocarbon dating of the charcoal, we know the rock art is at least 14,000 years old, possibly from the time of Urd.
The end of the ice age in Europe
The ice covering northern Europe began to retreat about 15,000 years ago, and haplogroup U5 was among the first people to repopulate central and northern Europe. The dominant U5 and its sister group U4 represented about 90% of the earliest Mesolithic hunter-gatherers. Fu et al.  found haplogroup U5 in both pre-ice age Paleolithic remains and post-ice age Mesolithic remains, and they concluded: “Because the majority of late Paleolithic and Mesolithic mtDNAs analyzed to date fall on one of the branches of U5, our data provide some support for maternal genetic continuity between the pre- and post-ice age European hunter-gatherers from the time of first settlement to the onset of the Neolithic.”
The end of the last Ice Age can be identified with the sudden warming in the period known as the Bølling/Allerød interstadial, that commenced 14, 600 years ago (see Figure 1 above). This warm period is associated with a rise in sea level and substantial melting of the Finnoscandian ice sheet. During this interstadial, woodland cover extended further northwards and people spread from the southern refugia into Switzerland, southern Germany and Belgium.
Sites such as Gough’s Cave in Somerset dated at 14,000 years ago provide evidence suggesting that humans returned to Britain towards the end of this ice age, in a warm period known as the Dimlington interstadial, although further extremes of cold right before the final thaw probably caused them to leave again and then return repeatedly. The environment during this ice age period would have been a largely treeless tundra, eventually replaced by a gradually warmer climate, perhaps reaching 17° Celsius in summer, encouraging the expansion of birch trees as well as shrub and grasses.
Two colder periods interrupted the Bølling/Allerød interstadial. Firstly, the Older Dryas occurred at 14,100 years ago and then the very dramatic reversal of the Younger Dryas occurred during the period 12,900 to 11,600 years ago (see Figure 1). During the Younger Dryas there was a temporary disappearance of the woodland cover that had previously extended over much of Europe. When the Younger Dryas ended, the change for northern Europe was dramatic. Annual average temperatures rose by about 15 °C with a midsummer rise of 5 °C and in midwinter by over 20 °C . It was towards the end of the Younger Dryas that humans returned to Britain, this time permanently, with perhaps Una among them.
Una (Haplogroup U5a1a)
Una probably lived 12,100 years ago (8,800-15,400) at the time people re-inhabited Britain, this time permanently. Una is approximately my great*470th grandmother. Haplogroup U5a1a is a lineage within Haplogroup U5 that arose in Europe around 12,000 years ago and is mainly found in northwest and north-central Europe. The modern distribution of Haplogroup U5a1a suggests that individuals bearing this Haplogroup were part of the populations that had tracked the retreat of ice sheets from Europe.
For Una and more recent haplogroup grandmothers, its not yet possible to pin down their locations based on ancient mtDNA distributions. Rather, geographic origins are identified based on the present day geographic distribution of data in the FTDNA database and GenBank . However, it should be noted that the sample size among FTDNA customers is largest from northwestern Europe, especially Scandinavia, the UK, Ireland, and central Europe, including Germany and Poland. Estimates of geographic origins may change as more test results from under sampled regions become available. It should also be noted that the place of greatest frequency in the present day may not indicate the place of origin of a subclade. Rather, the place with the greatest diversity in test results is a better indicator of the place of origin .
FTDNA has only four samples for U5A1A as most people alive today are from descendant haplogroups with additional mutations in the mtDNA. Three of the four samples are from Scotland and the fourth from Latvia. These numbers are too small to make any inference as to location, but are consistent with Una being part of the migration back into northern Europe and Britain around 12,000 years ago. At this time, Ireland and Britain were still joined to continental Europe by Doggerland, and so the reoccupation of Britain would have been on foot. I like to imagine that Una was one of the first humans to return to Britain after the ice.
As mentioned earlier, during the last ice age the sea level was substantially lower due to the amount of water locked up in the ice sheets. Britain was connected to Europe by an area of land known as Doggerland. This stretched from Britain’s present east coast to present-day Netherlands, the western coast of Germany and the peninsula of Jutland. It was probably a rich habitat with human habitation during the Mesolithic period, and was not just a path for humans to reach Britain after the ice receded. Fishing vessels have dragged up remains of mammoths, lions and other animals, and a few prehistoric tools and weapons. Doggerland was flooded by rising sea levels around 8500-8200 years ago. Hill et al.  have presented evidence that the final inundation of islands resulting from the partial inundation of Doggerland was due to a massive tidal wave 8,150 years ago due to an earthquake off the coast of Norway. The resulting tidal wave of up to 10-20 metres height could have wiped out the last remaining human settlements in Doggerland.
Although Una may have lived in what is now Europe, or may have been one of the first humans to reach Britain, I’ve chosen to show her location as in Doggerland on the migration map. At 11,600 years ago, dramatic global warming killed off the bitterly cold winters and resulted in the thickest covering by woodland that Europe had known in 100,000 years . Many animal species also returned to inhabit the land.
Ulfa (Haplogroup U5a1a1)
Haplogroup U5a1a1 originated around 6,800 years ago (5,000 – 8,700). I have given the name Ulfa (Old Norse for female wolf) to the ancestral mother of U5a1a1 and she was approximately my 260th great grandmother.
By 9,500 years ago, the rising sea levels caused by the melting glaciers cut Britain off from Ireland and, by around 6500 to 6000 BC, the plains of Doggerland were submerged and continental Europe was cut off for the last time. The warmer climate changed the Arctic environment to one of pine, birch and alder forest; this less open landscape was less conducive to the large herds of reindeer and wild horse that had previously sustained humans. Those animals were replaced in people’s diets by pig and less social animals such as elk, red deer, roe deer, wild boar and aurochs (wild cattle), which would have required different hunting techniques. Tools changed to incorporate barbs which could snag the flesh of an animal, making it harder for it to escape alive. Tiny microliths were developed for hafting onto harpoons and spears. Woodworking tools such as adzes appear in the archaeological record, although some flint blade types remained similar to their Palaeolithic predecessors. The dog was domesticated because of its benefits during hunting, and the wetland environments created by the warmer weather would have been a rich source of fish and game. It is likely that these environmental changes were accompanied by social changes. Humans spread and reached the far north of Scotland during this period.
The older view of Mesolithic Britons as nomadic is now being replaced with a more complex picture of seasonal occupation or, in some cases, permanent occupation. Travel distances seem to have become shorter, typically with movement between high and low ground.
Urwen (Subgroup U5a1a1-T16362C)
Urwen, the maternal founder of this subgroup, has not been dated. This subgroup is identified by a single additional mutation (T16362C) to U5a1a1, and the following haplogroup U5a1a1d by two further mutations, so I estimated the age of Urwen as 5,900 years ago (3,900-8,200) assuming a constant rate of mutation. Given the wide uncertainty, I somewhat arbitrarily adjusted this to 5,300 years ago, the time when British took up farming. She is named after the second child of Húrin and Morwen, and is my great*200th great grandmother.
Six of the ten samples in the FTDNA database with haplogroup U5a1a1-T16362C come from Britain. For the preceding haplogroup U5a1a1, 55% of the samples are from Britain and another 33% from Ireland. For the following haplogroup U5a1a1d, 61% are from Britain and 26% from Ireland. These percentages assume samples from people in former British colonies can be attributed to Britain or Ireland also. I provisionally locate Urwen in England 5,300 years ago, as at least 2/3 of the British samples are from England.
Ten thousand years ago, at the end of the last Ice Age, the human population numbered only a few millions and all their food came from wild plants and animals. Foragers moved seasonally in small groups to obtain their food supplies and population densities remained low for many millennia. Then around 10,000 years ago, people in West Asia began to domesticate some of those species, and settled down and occupied favourable sites year-round. This profound change in human behaviour led to the beginnings of agriculture, enabling more people to be supported on a given area of land – although at the cost of the greater effort needed to cultivate crops and raise domestic animals. The effects of settling down, population increase, and growing dependence on agriculture led to increases in the number and size of settlements, to the development of more complex, less egalitarian societies, and, eventually, to urban life and civilization.
Agriculture began independently in various parts of the world, West Asia, China, South Asia, South America, Africa and the Pacific. From around 9500 BC, the eight Neolithic founder crops – emmer wheat, einkorn wheat, hulled barley, peas, lentils, bitter vetch, chick peas, and flax – were cultivated in the Levant (West Asia). Agriculture spread west across Europe and reached Britain around 5,300 years ago.
There has been a longstanding debate about whether the farmers of western and northern Europe were the descendants of local hunter-gatherers, or migrants from the Levant. The recent published genetic evidence seems to suggest that the earliest European farmers were migrants from the east, and that there was substantial migration into western Europe with some interbreeding between the migrants and the hunter-gatherers [17, 18]. As the farming culture became more established, its likely that hunter-gatherer populations also adopted a farming lifestyle. Haplogroup U5 is a marker of the Mesolithic hunter-gatherer population of Europe that pre-dated and survived through the last ice age, and the substantial migration of farmers into Europe after the ice age is the reason that overall, U5 now only represents about 9% of European mtDNA and is concentrated in Scandinavia and the British Isles.
Urwen lived in the neolithic age at the beginning of agriculture in Britain about 3300 BCE. The Neolithic period in the British Isles was characterised by the adoption of agriculture and sedentary living, leading to the gradual decline of the hunter-gatherer lifestyle. To make room for the new farmland, these early agricultural communities undertook mass deforestation across the islands, dramatically and permanently transforming the landscape. At the same time, new types of stone tools requiring more skill began to be produced; new technologies included polishing. the Neolithic peoples of the British Isles grew cereal grains such as wheat and barley, and these therefore played a part in their diet. Nonetheless, this was supplemented at times with wild, un-domesticated plant foods such as hazelnuts.
The Neolithic also saw the construction of a wide variety of monuments in the landscape, many of which were megalithic in nature. The earliest of these are the chambered tombs of the Early Neolithic. Wayland’s Smithy is a Neolithic long barrow and chamber tomb site located near the Uffington White Horse in Oxfordshire, which I visited in 1992 and again in 2019 (waylands-smithy). This barrow was built in two phases, a timber chambered oval barrow built around 5700 years ago and the second stone chambered long barrow around 5400 years ago, the time of Urwen. In the Late Neolithic, this form of monumentalization was replaced by the construction of stone circles, a trend that would continue into the following Bronze Age.
Ulrica (Haplogroup U5a1a1d)
U5a1a1d has an age estimate of about 4000 years ago and is generally northern European, mostly the British Isles (which accounts for 87% of samples in the FTDNA database). The founder of this haplogroup, Ulrica, lived in the early years of the Bronze Age in Britain around 2000 BCE and is my 150th great grandmother. The Neolithic age was followed by the Bronze Age in Britain, lasting from around 2500 BCE to 800 BCE. The skill of refining metal is thought to have been of Iberian origin, brought to Britain as part of the Beaker culture around 4500 years ago. At first, items were made from copper but from around 2150 BCE smiths had discovered how to make bronze (which is much harder than copper) by mixing copper with a small amount of tin. With this discovery, the Bronze Age began in Britain. Over the next thousand years, bronze gradually replaced stone as the main material for tool and weapon making.
Britain had large reserves of tin in the areas of Cornwall and Devon in what is now Southwest England, and thus tin mining began. By around 1600 BCE, the Southwest of Britain was experiencing a trade boom as British tin was exported across Europe. The rich Wessex culture developed in Southern Britain at this time. The weather, previously warm and dry, became much wetter as the Bronze Age continued, forcing the population away from easily-defended sites in the hills and into the fertile valleys. Large livestock farms developed in the lowlands which appear to have contributed to economic growth and inspired increasing forest clearances. Higher ranking dead were buried under barrows and many of these can still be seen in Wiltshire.
During the British Bronze Age, large megalithic monuments similar to those from the Late Neolithic continued to be constructed or modified, including such sites as Avebury, Stonehenge and Silbury Hill. This has been described as a time when elaborate ceremonial practices emerged among some communities of subsistence agriculturalists of western Europe. Avebury was built during the period 3000 – 2600 BCE some hundreds of years before Ulrica lived. Construction of Stonehenge commenced around 3000 BCE and ended around 2200 BCE, not long before the time when Ulrica probably lived. I have somewhat arbitrarily chosen her location as Wiltshire, England.
The British Iron Age (750 BCE – 43 CE)
In around 750 BCE iron working techniques reached Britain from Southern Europe. Iron was stronger and more plentiful than bronze, and its introduction marks the beginning of the Iron Age. Iron working revolutionized many aspects of life, most importantly agriculture. Iron tipped ploughs could churn up land far more quickly and deeply than older wooden or bronze ones, and iron axes could clear forest land far more efficiently for agriculture. There was a landscape of arable, pasture and managed woodland. There were many enclosed settlements and land ownership was important.
It is generally thought that by 500 BCE most people inhabiting the British Isles were speaking Common Brythonic (or British), a Celtic language descended from Proto-Celtic. By the sixth century CE, British had produced four separate languages: Welsh, Breton, Cornish, and Cumbric. Evidence from Welsh shows a great influence from Latin on British during the Roman period, and especially so in terms related to the Church and Christianity, which are nearly all Latin derivatives. British was later replaced in most of Scotland by Gaelic and south of the Firth of Forth also by Old English (which later developed into the Scots language). British survived into the Middle Ages in Southern Scotland and Cumbria. British was gradually replaced by English throughout England; in the north, Cumbric disappeared as late as the 13th century and, in the south, Cornish was effectively a dead language by the 19th century.
Iron Age Britons lived in organized tribal groups, ruled by a chieftain. As people became more numerous, wars broke out between opposing tribes. This was the reason for the building of many hill forts; there are over 2,000 Iron Age hillforts known in Britain (see my earlier post). By about 350 BCE many hillforts went out of use and the remaining ones were reinforced. Pytheas was quoted as writing that the Britons were renowned wheat farmers. Large farmsteads produced food in industrial quantities and Roman sources note that Britain exported hunting dogs, animal skins and slaves.
Population estimates vary but the number of people in Iron Age Great Britain could have been three or four million by the first century BCE, with most concentrated densely in the agricultural lands of the South. The average life expectancy at birth would have been around 25, but at the age of five it would have been around 30. These figures would be slightly lower for women, and slightly higher for men throughout the Middle Iron Age in most areas, on account of the high mortality rate of young women during childbirth; however, the average age for the two sexes would be roughly equal for the Late Iron Age.
The end of the Iron Age extends into the early Roman Empire, but in parts of Britain that were not Romanized, such as Scotland, the period is extended a little longer, say to the 5th century.
The first extensive Roman campaigns in Britain were by the armies of Julius Caesar in 55 and in 54 BCE, when my great*75th grandmother was alive. But the first significant campaign of conquest did not begin until 43 CE, in the reign of the Emperor Claudius. Roman Britain was the part of the island of Great Britain controlled by the Roman Empire from 43 CE until ca. 410 CE.
Following the conquest of the native Britons, a distinctive Romano-British culture emerged under provincial government, which, despite steadily extended territorial control northwards, was never able to exert definite control over Caledonia. The Romans demarcated the northern border of Britannia with Hadrian’s Wall, completed around the year 128. Fourteen years later, in 142, the Romans extended the Britannic frontier northwards, to the Forth-Clyde line, where they constructed the Antonine Wall, but, after approximately twenty years, they then retreated to the border of Hadrian’s Wall. Most Romans departed from Britain around the year 410, which began the post-Roman period (5th–6th century CE), but the legacy of the Roman Empire was felt for centuries in Britain. The departure of the Romans took place in my great*58th grandmother’s time.
Viviane (Haplogroup U5a1a1*)
Viviane is the last of my maternal ancestors currently identifiable through the mitochondrial DNA analysis of my haplogroup. A very approximate estimate of the period she lived, based on average rate of mtDNA mutation, is 410 CE. This is the year that the Romans left Britain. However, the uncertainty range is quite wide at 20 CE to 1020 CE, and it is just as likely that she lived 100 years later around 500 CE. Why does that interest me? If King Arthur was not just a legend, but based on a real British leader, he most likely lived in the late 5th and early 6th century and led the defence of Britain against Saxon invaders in the early post-Roman period. So running low on Celtic/British names that start with U, I’ve chosen to call the founder of this final haplogroup Viviane, after the Lady of Lake who gave King Arthur his sword Excalibur, and later imprisoned Merlin in a cave. Viviane is also known as Nimüe in many of the legends.
The population of Britain may have decreased after the Roman period by between 1.5 and 3 million. This reduction may have been caused by environmental changes or by plague and smallpox (around 600 CE, the smallpox spread from India into Europe). The Plague of Justinian entered the Mediterranean world in the 6th century and first arrived in the British Isles in 544 or 545, when it reached Ireland. It is estimated that the Plague of Justinian killed as many as 100 million people across the world. It caused Europe’s population to drop by around 50% between 550 and 700.
Viviane was approximately my great*58th grandmother, or 55th if we take the later date of 500 CE. From the other end of the long line of maternal ancestors, the earliest I have been able to identify is my great-great-great-grandmother, Elizabeth Poulteck (1766-1848) who married George Bookmaster (1771-1845) and had a daughter Amelia Buckmaster who was my great-great-grandmother. The family lived in Hounslow, Middlesex, which is now part of greater London neighbouring Heathrow Airport. Amelia was born in Hounslow in 1809 and died in 1892 in Wallingford, Berkshire. Amelia’s daughter Anne married a man called James Cook (not the one who discovered Australia) and they migrated to Australia in 1872
I think its quite wonderful that I have been able to use genetic science to decode the history of my ancestors coded in my mitochondrial DNA and trace their migration path all the way back to mitochondrial Eve, who lived in northern Botswana around 178,000 years ago.
 Sykes B. The seven daughters of Eve. London, Bantam Press 2001.
 Oppenheimer, Stephen (2003). Journey of Mankind. Adapted from Out of Eden / The Real Eve, http://www.bradshawfoundation.com/journey/
 Chan, E.K.F., Timmermann, A., Baldi, B.F. et al. Human origins in a southern African palaeo-wetland and first migrations. Nature 575, 185–189 (2019) doi:10.1038/s41586-019-1714-1 https://www.nature.com/articles/s41586-019-1714-1
 Oppenheimer, Stephen (2006), The Origins of the British: A Genetic Detective Story, New York, Carroll and Graf.
 Soares P, Ermini L, Thomson N, Mormina M, Rito T, Röhl A, Salas A, Oppenheimer S, Macaulay V, Richards MB (2009). Correcting for purifying selection: an improved human mitochondrial molecular clock American Journal of Human Genetics, 84(6):740-59.
 Fu Q, Mittnik A, Johnson PLF, et al. A revised timescale for human evolution based on ancient mitochondrial genomes. Curr Biol. 2013;23(7):553–559. doi:10.1016/j.cub.2013.02.044 https://www.cell.com/current-biology/fulltext/S0960-9822(13)00215-7?code=cell-site
 Gail Tonnesen. The Subclades of U5. FTDNA Haplogroup U5 Project. July 15, 2012, revised July 18, 2014; June 6, 2015.
 Rob Spencer. SNP Tracker. Tracking Back: a website for genetic genealogy tools, experimentation, and discussion. Available at http://scaledinnovation.com/gg/snpTracker.html?fbclid=IwAR0irzIVQiqVzLsWodhJfnA8h-fbxYlLaPllOYf6kEQ146Ba002sW8jxYok (accessed 25 February 2020).
 Young O, Steffen W. (2009) The Earth System: Sustaining planetary life support systems. In: Principles of Ecosystem Stewardship: Resilience-Based Resource Natural Resource Management in a Changing World. F.S. Chapin III, G.P. Kofinas and C. Folke (eds), Springer-Verlag, New York, pp. 295-315
 Melissa Hogenboom, 29 Jan 2020. How did the last Neanderthals live? https://www.bbc.com/future/article/20200128-how-did-the-last-neanderthals-live (accessed 28 Feb 2020)
 Malyarchuk et al. (2010)  Malyarchuk B, Derenko M, Miroslava Grzybowski T, Perkova M, Rogalla U, Vanecek T, Tsybovsky I (2010). The Peopling of Europe from the Mitochondrial Haplogroup U5 Perspective. PLoS One. 2010 Apr 21;5(4):e10285.
 Burroughs WJ (2005). Climate change in prehistory: the end of the reign of chaos. Cambridge University Press: NY.
 Hill J, Collins GS, Avdis A, Kramer SC, Piggott MD. How does multiscale modelling and inclusion of realistic palaeobathymetry affect numerical simulation of the Storegga Slide tsunami? Ocean Modelling 83 (2014) 11–25.
 Mithen S (2003). After the ice: a global human history 20,000-5,000 BC. Phoenix: London.
 Barras C. The three ancestral tribes that founded Western civilization. New Scientist, 1 July 2015.
 Hofmanová Z, Kreutzer S, Hellenthal G, et al. Aegean origin of European Neolithic farmers. Proceedings of the National Academy of Sciences Jun 2016, 113 (25) 6886-6891; DOI: 10.1073/pnas.1523951113 Available at https://www.pnas.org/content/113/25/6886