The Turkish domestication of the chickpea.

One of the neolithic farmers founder crops.

Chickpeas (Cicer arietinum) are large roundish legumes, that look rather like a large round pea with an interesting bumpy surface. A staple of Middle Eastern, African and Indian cuisines, domesticated chickpeas (also called garbanzo beans) come in two main groups called desi and kabuli, but you can also find varieties in 21 different colors and several shapes. The wild version of chickpeas (Cicer reticulatum) is only found in parts of what is today southeastern Turkey and adjacent Syria, and it is likely that it was first domesticated there, about 10,500 years ago. Chickpeas store really well, and are high in nutritive value, and were part of the farming culture that grew out of the Neolithic of the Fertile Crescent.

Domesticating Chickpeas
There are some interesting features about the domestication of chickpeas that were pointed out in a 2007 article by Zohar Kerem and colleagues at the Hebrew University of Jerusalem’s Faculty of Agricultural, Food and Environmental Quality Sciences. The wild form of chickpea ripens only in the winter, while the domesticated form can be sown during the spring for summer harvest. Chickpeas grow best in winter when there is adequate water available; but during the winters they are susceptible to Ascochyta blight, a devastating disease which has been known to wipe out entire crops.

In addition, according to recent studies, the domesticated form of chickpea contains nearly twice the tryptophan of the wild form, an amino acid that has been connected with higher brain serotonin concentrations and higher birth rates and accelerated growth in humans and animals.

Chickpea Varieties and Archaeological Sites
The oldest variety of chickpea is the desi form; desi are small, angular, and variegated in color. Scholars believe desi originated in Turkey and was subsequently introduced into India where the most common form of chickpea is the kabuli. Kabuli have large beige beaked seeds.

Domesticated chickpeas have been found at several archaeological sites, including Tell el-Kerkh in Syria; Cayönü (7250-6750 BC), Hacilar (ca 6700 BC), and Akarçay Tepe (7280-8700 BP) in Turkey; and Jericho (8350 BC to 7370 BC) in the West Bank. The earliest to date is Tell el-Kerkh, in the late 10th millennium BC, and scholars suspect that since el-Kerkh is a considerable distance from the native lands of the wild chickpea, the domestication took place somewhat earlier than that.

So, the  domestication seems to be Turkish, about 10,000 plus years ago.

The Turkish/Northern Syrian origin of lentils.

Identification of the lentil’s wild genetic stock

  The origin of lentil from the taxon Lens culinaris subsp. orientalis has been proved by morphological evidence and breeding experiments. This wild form exhibits variation in many characters and is distributed over a vast area from the Middle-East to central Asia. Characters that are polymorphic in the wild progenitor but monomorphic in the cultigen can be utilized for better identification of the genetic stock which gave rise to the domesticated lentil. Three characters of that kind have been identified in lentil: chromosomal architecture, crossability potential and restriction pattern of chloroplast DNA. Nearly all accessions of the cultivated lentil tested to these three characters have been found monomorphic, but considerable polymorphism exists in the wild accessions. Three subsp. orientalis accessions have been shown to share the above characters with the cultigen and hence can be regarded as members of the genetic stock from which lentil was domesticated. These three accessions originated from eastern Turkey and northern Syria.

Also, the oldest lentils found were 11,000 years old from a Greek cave. Since the lentil is not native to Greece, it’s not a stretch to figure out these must have been cultivated. This would mean the growing of lentils predates cereals in Greece, meaning farming started earlier than believed in Europe (by about two thousand years) and that cultivation of lentils predates the cultivation of cereals.

Hallan Cemi Tepe, home of the first pork chop

Some Preliminary Observations Concerning Early Neolithic Subsistence Behaviors in Eastern Anatolia

Michael Hosenberg, .Clark Nesbitt, Richard K Redding, Thornas F Strasser

In 1991 a salvage excavation was begun at Hallan Cemi Tepesi, a largely aceramic (so some pottery) site in the Taurus foothills of eastern Turkey.’ The results of the 1991 through 1993 field seasons permitted some preliminary observations concerning the material culture of the site’s early Neolithic inhabitants. Of particular note was the relatively high degree of cultural complexity implied by that material culture (see Rosenberg and Davis 1992; Rosenberg 1994). Also of note was the evidence suggesting that, at its earliest stages, the Neolithic tradition in eastern Anatolia evolved with only minimal influence from the contemporaneous Levantine complex.

Excavations at Hallan Cemiare ongoing and the results of the 1994 field season make it necessary to once again modify some of the tentative conclusions concerning the site’s stratigraphy. More importantly, the ongoing analyses of the botanical and fauna1 remains, as well as of relevant aspects of the artifact assemblage, now make it possible to begin making some preliminary observations about the subsistence behaviors of the site’s inhabitants. The picture that is emerging from these ongoing analyses is often at odds with prior expectations. For example, though sedentism is indicated, it was apparently not based on the exploitation of cereals. The site’s inhabitants also appear to have been experimenting with animal domestication. In all. the Hallan Cemi data promise to significantly alter our understanding of the origins of food production and animal husbandry in south-western Asia.
The Botanical Assemblage Carbonized plant remains are consistently well preserved in the Hallan Cemi depos- its. Collection was largely by means of flotation involvinga sample of the site’s deposits, though individual seeds, nuts, etc. were also collected by hand in some instances. What follows is for the most part based on the formal analysis of a limited number of flotation samples from the 1992 season  as well as more preliminary analysis of samples from other contexts. Analysis of the sample balance is ongoing.

In the samples analyzed to date, relatively few seeds of wild grasses were found and most were in fragmentary condition. None have yet been identified as belongingto the cereal grasses. Compared to other sites of this period in Iraq and the Levant, this relative paucity of wild grasses is surprising It is, however, consistent with the dearth of sickle blades in the Hallan Cerni chipped stone assemblage (Rosenberg 1994: 128).

In contrast, pulses are common. They are mostly fragmentary and thus cannot be identified beyond I Viciailarlgrzis. However, identifiable examples of both lentils (Lens sp.) and bitter vetch (l’icia enilia) were found. Nuts are also common. These include wild almond (Anq@alussp.), pistachio (Pisracia S-.) and another thin-walled nut that remains to be identified. During both the 1993 and 1994 seasons, deposits in several parts of the mound yielded concentrations of wild almond (Fig. 5). Wild almonds contain potential toxins, yet almonds were clearly of great economic importance at Hallan Cemi despite that latent toxicity. This suggests the existenc eof processes for mitigating that latent toxicity and rendering almonds into an edible product. Judgingfrom the concentrations of charred almonds encountered in 1993 and 1994, roasting seems to have played a part in the processing of almonds.

It is also perhaps noteworthy that small, shallow sand and gravel pits occur scattered about the site. Though it is not clear whether these shallow pits were used for food preparation and, if so, for what foods, it is conceivable that these sand pits also played a role in the processingof almonds. Also common in the botanical assemblage are the seeds of sea club-rush (Bolboschoems ntaririnzus), a species of Po&gonzim, and Gundelia tournefortii. The presence of Guidelia totirizefortii is particularly interesting, as it is not often reported to be found at sites of this period. Gztndelia is a perennial tumbleweed belonging to the daisy family (Composirae). Though typically native to steppe habitats, it does occur in open woodland, such as appears to have then existed in the vicinity of Hallan e m i .The fruit consists of a woody and fibrous capitulum enclosing a single waxy achene (weight ca. 0.03 gm) in the single fertile floret at the center of the capitulum. The achene, as its waxy appearance suggests, is rich in fatty oils.

‘ According to Rouena Gale, who graciously provided these data, Fraxinus, Quercus, Prunus, Pistacia, and Salix or Popuftrs are represented in the wood charcoal from the site. Questionably, buckthorn (cf. Frangula alnus) is also present. The SalidPopults charcoal probably indicates the proximity of riverine forests to the site, ivhilc the olhrr specics are consistent \vith a mixed oak forest. 

 Collectingthe fruit simply involves shaking the plant upside down, as this causes the fruits to drop out. The achene, however, is tightly enclosed at the base of the capitulum and cannot be extracted without breaking open the fruit. One widely used method for extract- ing fat-rich seeds from tough shells in nuts is roasting. It is, therefore, noteworthy that, in addition to being found as scattered single fruits, a 5 cm thick lens, consisting of hundreds of more or less intact charred Grtndelia fruits, was found in the central open area. Perhaps a batch was being roasted and, for whatever reason, the fruits were burnt too completely for consumption of the seed, resulting in them being discarded as a unit.

The Fauna1 Assemblage

Animal exploitation was an important subsistence activity at Hallan Cemi, as attested to by the more than 2 tons of animal bone thus far unearthed. Of the 22,000+ bones (in- cluding small fragments) examined to date, 2,097 could be assigned to mammalian taxa and 91 1 to non-mammalian taxa. The bones and horn cores of sheep (Ovis sp.) and goats (Capm sp.) are the single most numerous mammalian component of the fauna1 assem- blage, comprising ca. 43% of all mammalian bone. Sheep outnumber goats at approxi- mately 6: 1. Red deer (Cervus elephlrs) follow at ca. 27% of all mammalian bone, followed in turn by canids (including two species of fox – L’zrlpes vrrlpes and Ciirlpes corsac – and either dog or jackal) at ca. 13%, pig (Sus sp.) at ca. 12%, brown bear (Ursus arclos) at ca. 3%, cape hare (Lepus capensis) at 2%. Stone marten (Martesjiona), wild cat (Fefis catzrs), beaver (Castor jiber), and European hedgehog (Erinacerrs ezrropaeus) also occur, but at less than 1% each. The remains of wild cattle (Bos primigenizrs) were not present in the samples analyzed so far, but are known to be present at the site (see Rosenberg 1994:Fig. 10). Non-mammalian taxa include two types of fish (catfish and a species of cyprinid), lizards, turtles of the genus Marrremjs and birds. Of these, turtle bones are by far the most numerous at 84% of the non-mammalian bone, followed by bird (10%), fish (6%), and lizard.

 Morphologically, the sheep and goats are wild. Moreover, approximately 66% of the sheep-goat remains (for which an age could be determined) come from individuals that survived to at least 42 months of age. This is a pattern consistent with the results of the hunting of a wild population (cf. Hesse 1982).

 In the case of pigs, the sample analyzed to date contains two measurable lower third molars and one measurable upper second molar. The two lower third molars measure 38.4 and 40.0 mm in length, which places them in the area of overlap between wild and domes- tic taxa. The upper second molar, however, measures 21.8 mm in length, within the range for domestic pig (cf. Flannery 1982). While this sample is obviously small, other lines of evidence are consistent with incipient pig domestication. The survivorship curve for pigs is in marked contrast to that for sheep-goats (Fig. 6). At least 10% of the individuals were less than 6 months of age when consumed, 29% never reached the age of 12 months, and only 3 1% survived to the age of 36 months. This pattern of consumption is similar to that found by one of the authors (Redding) at sites in Egypt, Iraq, and the Levant that yielded domestic pigs.

The present day economic importance of sheep and goats in the Near East has tended to foster the implicit presumption that they were the earliest animal domesticates in that area. However, the possible early domestication of pigs is not surprising when one considers certain facts. As Redding (n.d.) has noted: 1) the fecundity and growth rate of pigs make them superior producers of protein relative to all other native Near Eastern domesticates; 2) the labor required for pig maintenance is lower than for other Near Eastern domesticates; 3) young pigs tame readily and will imprint on humans; and, 4) juvenile or neonate pigs are relatiely easy to obtain. These qualities make the pig an ideal candidate for early experiments with animal domestication.

However, as also noted, pigs are more difficult to control or herd than sheep or goats. This makes pigs a poor choice of domesticate (relative to sheep and goats) in situations where intensified production of animals is desired. Pigs are also competitors with humans for cereals. This makes pigs a poor choice of domesticate (relative to sheep and goats) in contexts where cereal Ygrass exploitation is a significant component of the human subsis- tenceeconomy. However, in situationswhere, for whatever reason, cereals were not a significant component of the human subsistence economy (as was apparently the case at Hallan Cemi), pigs would seem superior to sheep and goats at the early stages of animal domestication.

Lastly, it should be noted that domesticated pigs are present at Cayonii(Lawrence 1980) and pigs, in general, are particularly common (relative to sheep and goats) in the lower levels of that site (Lawrence 1982). Whether domesticated pigs precede domesti cated sheep and goats at that site is not made clear in the published reports.

The Ground Stone Assemblage

Ground stone tools of types generally thought to be subsistencerelated constitute the next largest tools artifact assemblage after chipped stone tools. Sandstone of varying types appears to have been the most commonly used raw material for both mobile (i e., hand stones, pestles) and stationary (i.e., querns, mortars) types. Limestone and various kinds of metamorphic rocks were also used. While much of this assemblageremains to be analyzed in detail, it is now possible to make some preliminary observationsabout the assemblage as a whole.

The handstonesare typically ovate to sub-rectangular in form – having often been purposehlly ground or pecked to shape – with either one or two flat to slightly convex workingsurfaces. They rarely exceed 15 cm on the longest dimension. In many cases, one or both of the horizontal surfaces were reused as what are sometimes informally called ‘nuttingstones’.’ Such ‘nuttingstones’ also occur on simple water-worn pebbles and stones ‘ As a rule ‘nutting stones’ are characterized by relatively small, very shallow, irregularly circular deprcssi- ons produced by batteringthat appear to have been shaped into a variety of configurations. Pestles are less common than handstones.

 They are typically cylindrical to slightly conical in form and circular to slightly squared in section. They rarely exceed 30 cm in length and are usually less well shaped than are the handstones. The querns are of both trough and basin type and range up to 50+ cm in overall length on the intact examples. The exteriors of these are also often pecked or ground to shape, with ovate and sub-rectangular forms the most common (Fig. 7). Bowl mortars are less common than querns and they range up to almost 20 cm in depth. The most common forms are ovate and sub-rectangular/squared, but the evidence for purposehlexterior shaping is less clear than for the querns.

 It has been suggested (Moore 1985; Goring-Morris 1987) that a prevalence of quernsover mortars in an assemblage implies an emphasis on the exploitation of seeds, as op- posed to nuts. Though Wright (1994:241) notes that the ethnographic record provides cause to question such a strict correlation, she does go on to note (1994:242-243) that grinding (as opposed to pounding) is most beneficial in the processing of cereals. In view of the preliminary botanical evidence (see above) suggesting that at Hallan Cemi grasses played a smaller dietary role than did nuts and pulses, the higher frequency of quernsover mortars in the ground stone assemblage is puzzling.

 Lastly, it was earlier suggested (Rosenberg and Davis 1992) that many of the querns and mortars were purposehllyrendered useless through intentional perforation of the bottoms. At that time, this conclusion was based solely on the fact that the perforationswere very often relatively large and that their edges were thick and not convergent with the base (see Fig. 7). This conclusion has now been supported by two new lines of evidence. First, during the 1994 season, we recognized for the first time four intact bases that had been punched out of stationary grinding stones by a (presumably) heavy blow to the interior working surface. Second, during the 1994 season, several (intact) perforated grindingstones were found that had apparently been spirally scored near the base of the interior surface. Such scoring would no doubt facilitate breakage and may have been carried out for precisely that purpose. No unperforated grinding stones exhibit this scarring. Why these grinding stones were intentionally rendered useless remains unknown. However, destruction associated with human death is an obvious, albeit untestable, possibility that is brought  to mind by a similar destruction pattern for prehistoric Mimbres ceramic vessels in the American southwest (e.g., see Fiedel 1987:213).

Concluding Comments

The subsistencepatterns emerging from the Hallan Cemidata are significant for two reasons. First, they are the first clear indication that we have for the existence of subsis- tence systems in southwestern Asia that did not revolve around reliance on the exploitation of grasses. Hallan Cemiwas, nevertheless, occupied year-round. This would appear to challenge theories that place cereal grass exploitation at the center of explanations (e.g., cr cdri Henry 1989) for the increased sedentism we see in southwestern Asia at the end of the Pleistocene. Second, the Hallan Cemi data suggest that pigs were the earliest animal domesticate, at least in eastern Anatolia. The data from Cayonii have long obliquely hinted at this. However, the consistently greater economic importance of ovicaprids in southwestern Asia aceramic sites has tended to foster the presumption that the earliest attempts at animal domestication would focus on these economically more important animals. The fauna1 data from Hallan Cemi are consistent with the data from these other sites, in that ovicapridswere here too much more intensively exploited than were pigs. It would appear, though, that factors other than economic importance (see above) were paramount in the selection of the earliest food animal domesticate. It is perhaps only with subsequent changes in plant food subsistence (to the exploitation of grasses), or the subsequent need to further intensify food animal production, that the knowledge gained in working with pigs was applied to ovicaprids.

 

The appalling spelling, the authors, not mine!

So, it looks like lentils preceded grains. Were these grains domesticated or wild?

First farmers ate bacon before bread.

From the NY Times

DIGGING at the ruins of a village in southeastern Turkey, where people lived more than 10,000 years ago, archeologists expected to turn up the usual traces of a society on the verge of the agriculture revolution. There should be leftover grains of wild wheat and barley and perhaps the bones of butchered sheep and goats in some early stage of domestication.

The archeologists found nothing of the kind. Instead, to their complete surprise, they dug up the ample remains of pig bones.

The discovery, they said, strongly suggests that the pig was the earliest animal that people domesticated for food. The diminished size of the molars was one of several clues that the transformation of wild boars into pigs was under way at that time. Radiocarbon analysis put the date at 10,000 to 10,400 years ago.

So in the foothills of the Taurus Mountains at a site known as Hallan Cemi, the domestication of the pig appeared to have occurred 2,000 years earlier than once thought — and 1,000 years before the taming and herding of sheep and goats.

Much earlier, at least 12,000 years ago, wolves more or less invited domestication as the dog, developing a symbiotic relationship with people. They became camp followers, sentinels and “best friend.” Only in a few cultures later on were dogs served as food.

A broader significance, archeologists said, was the absence of any sign of wheat or barley at the settlement. The prevailing assumption, based mainly on research to the south in Syria and the Jordan River Valley, has been that with the end of the last ice age, wild grains were abundant in the more temperate climate over the entire Middle East. People settled down to harvest them, and this led to agriculture, animal husbandry and eventually the rise of cities and civilization.

“All early agricultural models are predicated on the assumption that people gathered wild wheat and other grains,” said Dr. Michael Rosenberg, an archeologist at the University of Delaware and director of the Hallan Cemi excavations. “But this is the earliest settlement site so far north, and it has no cereals. So another resource must have made it possible to settle down.”

In a report at a recent meeting of the Society for American Archeologists, Dr. Richard W. Redding, a University of Michigan archeologist and member of the discovery team, said that a heavy reliance on data from southern sites in the Levant might have resulted “in a very narrow view of the origin of food production.” There may have been a variety of ways by which people made the transition from foraging to farming, and some of them did not include the intensive use of wild cereals as a crucial first step.

Dr. Patricia Wattenmaker, a University of Virginia archeologist with wide experience in Middle East excavations, said the new findings were among the first from this part of the Turkish highlands in prehistory and were certain to force a serious rethinking of theories regarding human subsistence patterns leading up to agriculture. Roles of Environment and Culture

“It looks as if the pattern varies from place to place,” Dr. Wattenmaker said. “This takes the punch out of arguments that environmental factors” over a wider area triggered the transition toward agriculture, she added, and suggested that “cultural factors were really the key.”

Dr. Robert J. Braidwood, a professor emeritus of archeology at the University of Chicago who is a specialist in research on early agriculture, praised the Hallan Cemi excavations for providing a much-needed examination of pre-agricultural cultures beyond the Levant. But the Turkish village of no more than 150 inhabitants was extremely small, he cautioned, and evidence from three or four more sites in the area might be necessary before drawing any sweeping conclusions.

The Hallan Cemi site is scheduled to be flooded next year by a new dam on the Batman River.

In three years of excavations at Hallan Cemi, though, archeologists have established that people there had left the wandering life of hunting and gathering for a more sedentary village existence. Ruins of small stone houses and stone sculptures indicated a permanent settlement, and the growth pattern in fresh water clam shells at the site revealed year-round occupation. Evidence of long-distance trade in obsidian, copper and Mediterranean shells reflected the expansion of economic horizons by an increasingly complex society.

All this was happening at the time of the Natufians, people in the Jordan Valley who were probably the first to adopt settling down as a permanent way of life. But if wild cereals were critical to the Natufians’ transition, the people at Hallan Cemi apparently depended on gathering nuts and seeds, hunting wild sheep and deer and raising pigs. The absence of any wild grains at the site was determined by Dr. Mark Nesbitt, a paleobotanist at University College, London.

No single piece of the pig evidence is conclusive, Dr. Redding reported, but all the clues together “are congruent with the early phases of the domestication of pigs.” Clue on Domestication

Not only are the bones plentiful and the molars smaller, he said, but they show that the people appeared to favor young male pigs more than would be expected if they were hunting wild animals. A preponderance of the bones were of male pigs under one year of age. If they were raising pigs, they would spare most of the young females for breeding. Survivorship patterns of hunted animals reveal a more normal age distribution.

Pigs may have been the villagers’ insurance against famine caused by any sudden shortage of nuts and fruits and wild game. In a pre-agricultural sedentary culture, Dr. Rosenberg said, such shortages posed a greater risk because the people had a more limited foraging and hunting range.

“We think they fiddled around with maintaining animals to decrease that risk,” he said, “and pigs make sense if they are not gathering and growing grains.”

For one thing, young pigs are easily obtained and tamed. They require little labor to control since they can be left to forage for themselves throughout the community. And they are the most efficient domesticated animal, Dr. Redding said, in that they convert 35 percent of food energy into meat, compared with 13 percent for sheep or a mere 6.5 percent for cattle.

Pigs, the archeologist concluded, may have represented one more transitional step in some pre-agricultural societies; the pattern was not always a direct progression from settling down to growing cereals to raising animals. Perhaps the subsistent strategy of the highland villagers was to supplement their diets of nuts, fruits and grasses with pigs until cereal production was adopted. In time, Dr. Redding said, the highlanders took up grain cultivation, probably as an innovation borrowed from the south. Decline of Pigs

In any case, the archeologists said, as soon as the people of Hallan Cemi began growing grain, there was a sharp decline in domestic pigs, which were gradually replaced by domestic sheep and goats. It was a necessity. Pigs compete with people for cereals. They could no longer be left to forage unattended near the village and fields, and they are not as easily herded as sheep and goats.

Although some of the interpretations are tentative and more research is required, Dr. Rosenberg and Dr. Redding said they were increasingly confident in their evidence for the early domestication of the pig. And contrary to previous findings in the Levant, they said, there could be sedentary village life without an abundance of grains, wild or cultivated.

“Hallan Cemi is almost a mirror image of what’s going on at this time in the Levant,” Dr. Redding said. “We will have to rethink all the models we’ve been developing about early food production.”

As a side comment to this article, the oldest domesticated wheat has been found just south of the Turkish border, but originates in Turkey.  It didn’t seem to spread into the Natufian culture particularly quickly, as 2,000 years later some were still eating  wild wheat. Also originating in Turkey is the wild chickpea, another neolithic staple crop. I’m really starting to doubt the Natufians were the originators of the farming revolution. More so, in the light that the farmers of the Neolithic expansion didn’t seem to look like them(Natufians were part Negroid), they seem to resemble to Anatolians more.

Also, the temple at Gobekli Tepe predates farming in most of the Natufian Levant by a few hundred, years, and that kind of stone temple building is not entry level civilisation (11,500 years ago).

It seems to me that the south east of Turkey needs to be thoroughly dug up so we can track down the original sites!

Early upper paleaolithic Europeans.

Nicked straight from Dienekes!

Some of the discordance Van Vark et al. see between genetic and morphometric results may be attributable to their methodological choices. It is clear that the affiliation expressed by a given skull is not independent of the number of measurements taken from it. From their Table 3, it is evident that those skulls expressing Norse affinity are the most complete and have the highest number of measurements ( = 50.8), while those expressing affinity to African populations (Bushman or Zulu) are the most incomplete, averaging just 16.8 measurements per skull. Use of highly incomplete or reconstructed crania may not yield a good estimate of their morphometric affinities. When one considers only those crania with 40 or more measurements, a majority express European affinity.

To examine this idea further, we use the eight Upper Paleolithic crania available from the test series of Howells ([1995]), all of which are complete. Our analysis of these eight, based on 55 measurements, is presented in Table 1. Using raw measurements, 6 of 8 express an affinity to Norse, and with the shape variables of Darroch and Mosimann ([1985]), 5 of 8 express a similarity to Norse. Using shape variables reduces the Mahalanobis distance, substantially in some cases. Typicality probabilities (Wilson, [1981]), particularly for the shape variables, show the crania to be fairly typical of recent populations. The results presented in Table 1 are consistent with the idea that Upper Paleolithic crania are, for the most part, larger and more generalized versions of recent Europeans. Howells ([1995]) reached a similar conclusion with respect to European Mesolithic crania.

…

Next, let us examine the issue of whether the EUP situation can be regarded as parallel to the Native American one. There are some obvious differences, principal among them the time frame. The European crania used by Van Vark et al. span 26,000 years, as against our North American sample that spans about 2,400 years. Their EUP series dates from 37,000 BP to about 9,000 BP, as against a maximum time frame for our North American sample of 9,400-7,000 BP (Jantz and Owsley, [2001]). The Upper Paleolithic time span is significantly older and more than 10 times longer than the American one, yet the EUP crania are not correspondingly further removed from the contemporary population. Given that European fossil crania are separated from their supposed descendants by greater temporal distance than is the case in America, one could easily accept that European fossil crania might be more loosely connected to the modern population. Yet, we observe just the opposite. The data in Van Vark et al. demonstrate a higher degree of affiliation with the supposed descendent modern population (16/35 = 46%) than we found in the American situation (1/11 = 9%).

American Journal of Physical Anthropology
Volume 121, Issue 2, Pages 185-188

Paleolithic and neolithic lineages in the European mitochondrial gene pool.

Paleolithic and neolithic lineages in the European mitochondrial gene pool.
M. Richards, H. Côrte-Real, P. Forster, V. Macaulay, H. Wilkinson-Herbots, A. Demaine, S. Papiha, R. Hedges, H. J. Bandelt, and B. Sykes
Department of Cellular Science, Institute of Molecular Medicine, University of Oxford, Oxford, UK.

.
Phylogenetic and diversity analysis of the mtDNA control region sequence variation of 821 individuals from Europe and the Middle East distinguishes five major lineage groups with different internal diversities and divergence times. Consideration of the diversities and geographic distribution of these groups within Europe and the Middle East leads to the conclusion that ancestors of the great majority of modern, extant lineages entered Europe during the Upper Paleolithic. A further set of lineages arrived from the Middle East much later, and their age and geographic distribution within Europe correlates well with archaeological evidence for two culturally and geographically distinct Neolithic colonization events that are associated with the spread of agriculture. It follows from this interpretation that the major extant lineages throughout Europe predate the Neolithic expansion and that the spread of agriculture was a substantially indigenous development accompanied by only a relatively minor component of contemporary Middle Eastern agriculturalists. There is no evidence of any surviving Neanderthal lineages among modern Europeans.

A commonly ‘misunderstood’ paper by Dr C. Loring Brace.

The amount of times I’ve read work that thinks this paper proves …

• The ancient Greeks were black
• All the moors were black
• etc
• That the original Europeans weren’t Caucasians (yes, some people are that  dumb).

It’s really quite entertaining. Dr Brace recently said of the Cro Magnons…

I was able to get just under 20 measurements on Cro Magnon of the two dozen data set I have used to compare populations in the world and the statistics showed convincingly that while  Cro Magnon does not tie in with the recent French, it does indeed tie closely with our English and Scandinavian samples. What we have been able to show is that the Upper Paleolithic and subsequent Mesolithic of northwest Europe simply developed there in situ out of Neanderthal precursors. We published some of this in Human Evolution 19(1):19-38 (2005) and in the Proceedings of the National Academy of Sciences 103(1):242-247 (2006). In the latter paper we showed that a picture of demic diffusion from the Middle East and subsequent absorption by the indigenous north and western Europeans can account for the appearance of living European form.

C. L. Brace

The conclusion reads that the Natufians in the Levant seemed to be a mix of Eurasian and Negroid, tending more to the Eurasian, and the African features had vanished into the population by the time of the Neolithic farming expansion. I’ll mark the most relevant quotes in bold.

So, here it is..

The questionable contribution of the Neolithic and the Bronze Age to European craniofacial form

C. Loring Brace,*† Noriko Seguchi,‡ Conrad B. Quintyn,§ Sherry C. Fox,¶ A. Russell Nelson, Sotiris K. Manolis,** and Pan Qifeng††Received September 20, 2005.

Many human craniofacial dimensions are largely of neutral adaptive significance, and an analysis of their variation can serve as an indication of the extent to which any given population is genetically related to or differs from any other. When 24 craniofacial measurements of a series of human populations are used to generate neighbor-joining dendrograms, it is no surprise that all modern European groups, ranging all of the way from Scandinavia to eastern Europe and throughout the Mediterranean to the Middle East, show that they are closely related to each other. The surprise is that the Neolithic peoples of Europe and their Bronze Age successors are not closely related to the modern inhabitants, although the prehistoric/modern ties are somewhat more apparent in southern Europe. It is a further surprise that the Epipalaeolithic Natufian of Israel from whom the Neolithic realm was assumed to arise! ( slightly incorrectly, as Turkey is now looking good for the origin of the Neolithic revolution) has a clear link to Sub-Saharan Africa. Basques and Canary Islanders (Guanches) are clearly associated with modern Europeans. When canonical variates are plotted, neither sample ties in with Cro-Magnon as was once suggested. The data treated here support the idea that the Neolithic moved out of the Near East into the circum-Mediterranean areas and Europe by a process of demic diffusion but that subsequently the in situ residents of those areas, derived from the Late Pleistocene inhabitants, absorbed both the agricultural life way and the people who had brought it.

Among those who deal with the background of European history, there is a generally accepted view that the foraging way of life in the post-Pleistocene Mesolithic was succeeded by the Neolithic farming way of life. With the addition of metallurgy, the Neolithic morphed into the Bronze Age, which was succeeded by the Iron Age and the more recent European civilization (1–4). Further there is a general acceptance of the assumption that the farming way of life of the Neolithic arose in the Middle East ≈11,000 years ago and spread to the western edge of Europe by about 6,500 years ago (Incorrect. The Neolithic farmers seem to have arrived in Europe about 8,000 years ago, and the oldest founddomesticated grains are 13,500 years old in Abu Hurerya, Northern Syria. The original grains seem to be domesticated from a wild race of Turkish einkhorn wheat) (5–10). Researchers have questioned whether that spread took place by cultural diffusion to in situ people (11) or whether it was a “wave of advance” or a matter of “demic diffusion,” the actual movement of groups of people (see refs. 1, 8, and 12–15). Some researchers have observed that, although the two possible modes of Neolithic spread need not be mutually exclusive (see refs. 9 and 12), principal components analysis of allele frequencies in living humans shows a southeast–northwest cline that favors the idea that the spread had been the result of actual demic movement rather than by diffusion of cultural elements to pre-existing populations (see refs. 11–15).

Previous assessments of the Neolithic spread from the Middle East westward have been based on a consideration of tools and pottery on the one hand and genetically controlled aspects of living human populations on the other (14, 15). Here we offer an assessment based on a comparison of a set of metric dimensions of both prehistoric and more recent human craniofacial morphology. Craniofacial analysis has been previously applied to this question, but the comparison to living populations was not done (16). It has already been shown that the quantitative treatment of craniofacial form produces a picture of the movement of human populations from Asia into the New World that is largely compatible with the picture produced by the molecular genetic comparison of nucleotide haplotypes (17, 18).

The underlying reason that such different approaches yield comparable results is that neither the nucleic acid components identified nor the particular craniofacial dimensions used have any obvious adaptive value. Both evidently behave in a manner compatible with what has been called the “neutral theory,” where the traits assessed are under genetic control and the differences between groups are principally the result of genetic drift (12–22). What they show, then, is the extent of genetically shared relationships between adjacent populations. Here we offer a comparable treatment of samples of recent and prehistoric human populations running from the Middle East to the western edge of the Eurasian continent, north to Crimea, east to Mongolia, and southward through Nubia and Somalia plus samples from North Africa and representatives of the Niger-Congo-speaking peoples of Sub-Saharan Africa (Table 1). Teeth and the tooth-bearing parts of facial skeletons of course do reflect differences in response to the forces of selection on different populations (23), but they were left out of our analysis.

 Table 1.
Samples and numbers used in the analysis

Sample	No.
1. Norway	40
2. Finn/Sami	21
3. Denmark	19
4. Iceland	34
5. England	39
6. France	67
7. Basque	22
8. Canary Islands	24
9. Switzerland	50
10. Germany	27
11. Czech	25
12. European Upper Palaeo.	8
13. France Mesolithic	4
14. Denmark Neolithic	40
15. England Neolithic	12
16. France Neolithic	44
17. Swiss Neolithic	22
18. German Neol. (Mühl.)	9
19. Ger. Neol. (Tauberbisch.)	7
20. England Bronze	26
21. Portugal Mesolithic	12
22. Portugal Neolithic	18
23. Italy	80
24. Sicily	9
25. Sardinia	15
26. Etruscan	38
27. Italy Eneolithic	32
28. Italy Bronze	7
29. Greece	22
30. Franchthi (Greek Mesolithic)	1
31. Nea Nikomedea (Greek Neolith.)	7
32. Greek Bronze	16
33. Middle East (Iran/Iraq)	16
34. Morocco	24
35. Algeria	25
36. Berber	15
37. Tunisia	12
38. Egypt	28
39. Israeli Fellaheen (farmers)	15
40. Taforalt/Afalou (Morocco)	10
41. Natufian	4
42. Algerian Neolithic	6
43. Egypt Bronze (Naqada)	52
44. Jericho Bronze	4
45. Kurgan Bronze (Crimea)	30
46. Mongolian Bronze (Chandman)	54
47. Somalia	30
48. Nubia	64
49. Nubia Bronze	15
50. Congo (Gabon)	36
51. Dahomey (Benin)	32
52. Haya (Tanzania)	36
Total	1,282

 

References Neighbor-Joining ComparisonsA battery of 24 craniofacial measurements (Table 2) was used to compare the similarities and differences of living human populations and their prehistoric predecessors where possible throughout the area in question. The significance of the difference between any pair of the total sample can be assessed from Mahalanobis D2 figures (24), and a graphic depiction of the similarities and distinctions of the various groups tested can be seen from the dendrogram produced by using the D2 figures as input for the neighbor-joining procedure (Fig. 1) (25). To compute the Mahalanobis distances, we used a pooled within-group covariance matrix derived from all groups and weighted by sex and group sample size. The neighbor-joining method can be used for discrete differences, as is done with molecular data, or it can be used on continuous data, as we have done here (25). Assessments can also be made with canonical variate plots, which have the added advantage that single individuals can be placed in relation to the other samples used (Fig. 2) (29–32).

 Table 2.
 Craniofacial measures used in the UMMA data set

Variable no.	Description
1	Nasal height
2	Nasal bone height
3	Piriform aperture height
4	Nasion prosthion length
5	Nasion basion
6	Basion prosthion
7	Superior nasal bone width
8	Simotic width
9	Inferior nasal bone width
10	Nasal breadth
11	Simotic subtense
12	Inferior simotic subtense
13	FOW subtense at nasion
14	MOW subtense at rhinion
15	Bizygomatic breadth
16	Glabella opisthocranion
17	Maximum cranial breadth
18	Basion bregma
19	Basion rhinion
20	Width at 13 (fmt fmt)
21	Width at 14
22	IOW subtense at nasion
23	Width at 22 (fmo fmo)
24	Minimum nasal tip elevation

Figure 1

Neighbor-joining dendrogram for a series of prehistoric and recent human populations running from the western edge of the Eurasian continent and North Africa to the Middle East and down East Africa as far as Somalia, plus a sampling of Niger-Congo-speaking people from Gabon, Benin, and Tanzania in Sub-Saharan Africa. The samples used and the number for each are spelled out in Table 1. The kinds of measurements used to generate the dendrogram are listed in Table 2.

 Fig. 2.
Placement of the samples used in Fig. 1 determined by the values of canonical variates 1 (30.0%) and 2 (16.2%).

It is no surprise to discover that individual samples of recent humans tie more closely with other samples of extant people from the same part of the world than with more distant peoples. What does come as a surprise is that the Neolithic samples tend to tie with Neolithic samples across the entire range from east to west but do not cluster with the living people in many of the areas tested. There is more of a link between the prehistoric and modern samples in southern Europe as opposed to the picture in central and northern Europe. Much the same is true for the Bronze Age samples, although these do tend to tie to the preceding Neolithic in the same part of the range tested.

Unlike the Neolithic, Bronze Age, and modern samples, the Palaeolithic samples are not from single sites. There is no single European Upper Palaeolithic sample large enough to run as a single twig in a dendrogram. Instead, we had to use Cro-Magnon 1, La Ronde du Barry, Abri Pataud, Saint Germain-La Rivière, and Le Placard, all from southwestern France, plus Obercassel 1 from western Germany, and Předmostí 3 and 4 from the Czech Republic. Measurements of the latter two specimens were taken on casts because the originals had been destroyed by retreating Germans near the end of World War II (33). The same kind of problem of finding more than one individual in a burial site also tended to be true for some of the available Mesolithic of Europe. Individual specimens from Brittany to Monaco (Gramat, Rastel, Recheril and Téviec) were lumped together to make the European Mesolithic sample. There are larger Mesolithic samples, but we were not able to get permission to work on them. The North African Epipalaeolithic sample was made on the basis of specimens from Afalou in Algeria and Taforalt in Morocco. The Natufian sample from Israel is also problematic because it is so small, being constituted of three males and one female from the Late Pleistocene Epipalaeolithic (34) of Israel, and there was no usable Neolithic sample for the Near East.

The difficulty in making comparisons with Neolithic and Palaeolithic samples is the result of the very different treatment of the deceased. Neolithic communities established cemeteries where the remains of the departed accumulated in some numbers. Most Upper Palaeolithic peoples tended to bury the dead singly and in widely separated locations. Furthermore, Neolithic pottery became fractured with considerable frequency, leaving potsherds in quantity at Neolithic sites. Consequently there may well have been a tendency to overestimate the size of Neolithic populations vis-à-vis the contemporary surviving foragers (6, 35, 36). Despite the small numbers and scattered locations of the Late Pleistocene specimens, they tend to cluster with each other rather than with any groups of more recent date.

In dendrograms such as Fig. 1, the little Natufian sample clusters with the Mesolithic of France, the North African Epipalaeolithic, and the European Upper Palaeolithic, but the lengths of each of these twigs show that the relationships are comparatively remote. These are all Late Pleistocene or very early post-Pleistocene groups, and they are also noticeably more robust than more recent human groups. The three Niger-Congo-speaking groups (the Congo from Gabon, the Dahomey from Benin, and the Haya from Tanzania) cluster together away from most of the other samples. They do show a somewhat more distant link to the Nubians and the Nubian Bronze Age, who are so close to each other that they were combined for subsequent analyses.

When the samples used in Fig. 1 are compared by the use of canonical variate plots as in Fig. 2, the separateness of the Niger-Congo speakers is again quite clear. Interestingly enough, however, the small Natufian sample falls between the Niger-Congo group and the other samples used. Fig. 2 shows the plot produced by the first two canonical variates, but the same thing happens when canonical variates 1 and 3 (not shown here) are used. This placement suggests that there may have been a Sub-Saharan African element in the make-up of the Natufians (the putative ancestors of the subsequent Neolithic), although in this particular test there is no such evident presence in the North African or Egyptian samples. As shown in Fig. 1, the Somalis and the Egyptian Bronze Age sample from Naqada may also have a hint of a Sub-Saharan African component. That was not borne out in the canonical variate plot (Fig. 2), and there was no evidence of such an involvement in the Algerian Neolithic (Gambetta) sample.

Conclusions
References Combining SamplesWhen groups that are close to each other in the dendrogram in Fig. 1 are combined to make a single dendrogram twig, the picture is simplified, but much the same conclusion is supported. Czech, Denmark, England, Etruscan, Finn/Sami, France, Germany, Iceland, Norway, Sardinia, and Swiss samples are combined to make a sample designated as “Modern Europe.” Algeria, Berber, Greece, Iran/Iraq, Italy, Morocco, Sicily, and Tunisia samples were combined to generate a “Modern Mediterranean” twig, and the Algerian Neolithic was run as a separate twig. Next the Congo, Dahomey, and Haya samples were run as a “Niger-Congo” twig. Then Neolithic samples from Denmark, England, France, Germany, and Portugal were combined with Bronze Age samples from England, Jericho, and Mongolia to make a “Late Prehistoric Eurasia” sample. Mongolia is a long way east of any of the other samples used, but it has previously been shown that the Mongolian Bronze Age sample is unrelated to modern Mongols and has more in common with prehistoric Europeans and the Native Americans of the United States–Canada border (17).

Next the Portuguese Mesolithic, Greek Neolithic, Italy Eneolithic, and Swiss Neolithic samples and the Italian and Greek Bronze Age samples were combined to make a “Prehistoric Mediterranean” twig. Then Naqada Bronze Age Egyptian, the Nubian, Nubia Bronze Age, Israeli Fellaheen (Arabic farmers), and Somali samples were lumped as “Prehistoric/Recent Northeast Africa.” The Natufians and the Algerian Neolithic samples were run as separate twigs, and there were separate twigs for Basques and Canary Islanders. Figure 3 shows the results of running all of these twigs in a single neighbor-joining dendrogram. Only 18 of the 24 variables were used to construct Fig. 3, allowing us to add the Basque sample. When the Basques are left out and all 24 variables are used, the main twigs in the resulting dendrogram relate to each other in exactly the same way as those in the 18-variable version shown in Fig. 3. The D2 figures that were used in the construction of Fig. 3 are printed in Table 3.

 Fig. 3.
Neighbor-joining dendrogram of combined adjacent groups from Fig. 1.

Mahalanobis distance figures for the twigs in Fig. 3

	1	2	3	4	5	6	7	8	9	10
1. Modern Europe	—
2. Modern Mediterranean	3.34	—
3. Niger-Congo	16.42	16.26	—
4. Late Prehistoric Eurasia	1.87	2.52	12.15	—
5. Prehistoric Mediterranean	4.19	3.90	15.60	2.65	—
6. Prehist/Recent NE Africa	5.16	5.22	6.67	4.54	5.78	—
7. Canary Islands	3.58	7.22	19.16	4.68	5.90	7.01	—
8. Basques	7.16	8.81	30.77	10.98	14.31	11.82	7.94	—
9. Natufian	21.00	19.93	14.66	14.00	16.59	15.31	20.62	33.97	—
10. Algerian Neolithic	8.20	7.62	12.84	6.71	5.71	5.14	6.47	14.98	17.60	—

 There are some generalizations that are apparent from the picture presented in both the greater individual numbers of twigs shown in Fig. 1 and the combined pattern shown in Fig. 3. When the maximum number of twigs is plotted, despite the very small numbers involved, the Late Pleistocene samples from Israel, Europe, and North Aftica tend to link to each other before they tie to the modern representatives of each of the areas in question, as shown in Fig. 1. In that run, the Natufian of Israel ties to the French Mesolithic and then to the Afalou/Taforalt sample from North Africa. These then link with the European Upper Palaeolithic sample and, somewhat surprisingly, with the Chandman (the Mongolian Bronze Age sample) and finally, at the next step, with the Danish Neolithic. One of the things that these geographically diverse groups clearly have in common is a degree of robustness that sets them apart from the recent inhabitants of the areas in which they are found.

Apart from the quantitative relationships shown in Figs. 1, 2, 3, 4, most of the Neolithic samples in Europe share nonmetric features of the lateral edge of the orbit, the shape of the gonial angle of the mandible, and the configuration of menton that are present even when degrees of size and robustness vary between the regions represented. These nonmetric attributes all support the view that most of the Neolithic inhabitants of Europe tie more closely together with each other than with the living representatives of the areas in question. The principal exception to this generalization is one of the two small samples of the German Neolithic, the Mühlhausen sample, which ties closer metrically to the living inhabitants of the Middle East and North Africa. Metrically the other German Neolithic sample, Tauberbischofsheim, links with the living Central European samples. Nonmetrically, those two small German Neolithic samples also appear strikingly different from each other.

 Fig. 4.
Canonical variates 1 (58.1%) and 2 (16.2%) for the same groups represented in Fig. 3.

The Niger-Congo speakers (Congo, Dahomey, and Haya) cluster closely with each other and a bit less closely with the Nubian sample (both the recent and the Bronze Age Nubians) and more remotely with the Naqada Bronze Age sample of Egypt, the modern Somalis, and the Arabic-speaking Fellaheen (farmers) of Israel. When those samples are separated and run in a single analysis as in Fig. 1, there clearly is a tie between them that is diluted the farther one gets from Sub-Saharan Africa. The other obvious matter shown in Fig. 3 is the separate identity of the northern Europeans. This matter is treated in the next section.
 
The Basque language is a linguistic isolate unrelated to any other language (37), and there is a long-held idea that the Basques may represent a modern survival of the Pleistocene human inhabitants of western Europe (38). Our measurements were made on the sample gathered from the French side of the French/Spanish frontier that runs through Basque country in southwestern France. These specimens were stored in the Broca collection at the Musée de l’Homme in Paris. Paul Broca himself had promoted the view that the Basques represent the continuing existence of the kind of Upper Paleolithic population excavated at the Cro-Magnon rock shelter in the village of Les Eyzies in the Dordogne region of southwestern France in 1868 (38–41). Shortly thereafter the “old man” (“le vieillard”) found in that rock shelter was elevated to the status of typifying a whole “Cro-Magnon race” regarded as ancestral to not only the Basques but also the aboriginal inhabitants of the Canary Islands (38, 42–45).

When the Basques are run with the other samples used in Fig. 1, they link with Germany and more remotely with the Canary Islands. They are clearly European, although the length of their twig indicates that they have a distinction all their own. It is clear, however, that they do not represent a survival of the kind of craniofacial form indicated by Cro-Magnon any more than do the Canary Islanders, nor does either sample tie in with the Berbers of North Africa as has previously been claimed (38, 45–46). This is particularly well documented when the 18 variables are used to generate a plot of the first two canonical variates as shown in Fig. 4. In this figure, one can see a clear link between the Niger-Congo sample and the Natufians. The Prehistoric/Recent Northeast African sample also has a subsequent link to the Niger-Congo sample in Fig. 3. Yet the D2 values in Table 3 show that it is slightly closer to Late Prehistoric Eurasia than to the Algerian Neolithic, Modern Europe, and Modern Mediterranean and that it is farthest from the Niger-Congo, the Natufians, and the Basques. Although the Algerian Neolithic sample has an even more residual link to this cluster, the D2 figures in Table 3 show that it is almost as far from the Niger-Congo twig as from the Basques and Natufians. The generally high D2 values for the Natufian sample in Table 3 are almost certainly a reflection of the very small sample size.

To test the analysis shown in Fig. 3, Cro-Magnon (Fig. 4, ×) was removed from the European Upper Palaeolithic sample and run as a single individual. Interestingly enough, Cro-Magnon is not close to any more recent sample. Clearly, Cro-Magnon is not the same as the Basque or Canary Island samples. Fig. 4 plots the first and second canonical variates against each other, but that conclusion is even more strongly supported when canonical variate 3 (not shown here) is plotted with variate 1. The probabilities of Cro-Magnon’s ties to any of the groups in Figs. 3 and 4 are shown in Table 4. If this analysis shows nothing else, it demonstrates that the oft-repeated European feeling that the Cro-Magnons are “us” (47) is more a product of anthropological folklore than the result of the metric data available from the skeletal remains.

 Table 4.
Probabilities and squared Mahalanobis distances between Cro-Magnon 1 and reference samples 

   Probabilities and squared Mahalanobis distances between Cro-Magnon 1 and reference samples

	ModEur	ModMed	NigCon	LPEurasia	PrehMed	P/RNEAfr	CanIsl	Basq	Natuf	AlgNe
Cro-Magnon
Posterior probability	0.49	0.01	0.00	0.39	0.03	0.01	0.07	0.01	0.00	0.00
Typicality probability (F distribution)	0.26	0.04	0.01	0.25	0.10	0.04	0.19	0.09	0.07	0.04
Squared Mahalanobis distance	21.72	30.53	36.35	22.15	26.80	30.10	24.42	28.30	35.00	36.00

 

Conclusions
The assessment of prehistoric and recent human craniofacial dimensions supports the picture documented by genetics that the extension of Neolithic agriculture from the Near East westward to Europe and across North Africa was accomplished by a process of demic diffusion (11–15). If the Late Pleistocene Natufian sample from Israel is the source from which that Neolithic spread was derived, then there was clearly a SubSaharan African element present of almost equal importance as the Late Prehistoric Eurasian element. At the same time, the failure of the Neolithic and Bronze Age samples in central and northern Europe to tie to the modern inhabitants supports the suggestion that, while a farming mode of subsistence was spread westward and also north to Crimea and east to Mongolia by actual movement of communities of farmers, the indigenous foragers in each of those areas ultimately absorbed both the agricultural subsistence strategy and also the people who had brought it. The interbreeding of the incoming Neolithic people with the in situ foragers diluted the Sub-Saharan traces that may have come with the Neolithic spread so that no discoverable element of that remained. This picture of a mixture between the incoming farmers and the in situ foragers had originally been supported by the archaeological record alone (6, 9, 33, 34, 48, 49), but this view is now reinforced by the analysis of the skeletal morphology of the people of those areas where prehistoric and recent remains can be metrically compared.
 

How this actually works in plain English… The Natufians were slightly more of Eurasian ancestry than African, and by the time the Neolithic farming expansion started, any Negroid features had been diluted to invisibility, and you are left with with an essentially Eurasian population. The African Niger Congo (included only as an outlier) never comes anywhere near the measurements of stone age/bronze age Europeans or bronze age North African and near East .

The origin and spread of Y chromosomes E and J.

Origin, Diffusion, and Differentiation of Y-Chromosome Haplogroups E and J: Inferences on the Neolithization of Europe and Later Migratory Events in the Mediterranean Area

Ornella Semino,1 Chiara Magri,1 Giorgia Benuzzi,1 Alice A. Lin,2 Nadia Al-Zahery,1,4 Vincenza Battaglia,1 Liliana Maccioni,5 Costas Triantaphyllidis,6 Peidong Shen,7 Peter J. Oefner,7 Lev A. Zhivotovsky,8 Roy King,3 Antonio Torroni,1 L. Luca Cavalli-Sforza,2 Peter A. Underhill,2 and A. Silvana Santachiara-Benerecetti1, 2003; 
 
The phylogeography of Y-chromosome haplogroups E (Hg E) and J (Hg J) was investigated in >2,400 subjects from 29 populations, mainly from Europe and the Mediterranean area but also from Africa and Asia. The observed 501 Hg E and 445 Hg J samples were subtyped using 36 binary markers and eight microsatellite loci. Spatial patterns reveal that (1) the two sister clades, J-M267 and J-M172, are distributed differentially within the Near East, North Africa, and Europe; (2) J-M267 was spread by two temporally distinct migratory episodes, the most recent one probably associated with the diffusion of Arab people; (3) E-M81 is typical of Berbers, and its presence in Iberia and Sicily is due to recent gene flow from North Africa; (4) J-M172(xM12) distribution is consistent with a Levantine/Anatolian dispersal route to southeastern Europe and may reflect the spread of Anatolian farmers; and (5) E-M78 (for which microsatellite data suggest an eastern African origin) and, to a lesser extent, J-M12(M102) lineages would trace the subsequent diffusion of people from the southern Balkans to the west. A 7%–22% contribution of Y chromosomes from Greece to southern Italy was estimated by admixture analysis.

References  It has been proposed that the observed decreasing frequency gradients of Y-chromosome superhaplogroups E (Hg E) (defined by the SRY4064 mutation) and J (Hg J) (characterized by the 12f2a-8kb allele) (Semino et al. 1996; Hammer et al. 1998; Rosser et al. 2000) reached southwestern Europe as a result of demic expansions of Neolithic agriculturalists from the Middle East (Semino et al. 1996; Hammer et al. 1998). The spatial frequency patterns of Hg E and Hg J, at this level of molecular resolution, accommodate both infiltrations of Neolithic agriculturalists into southwestern Europe and cultural adaptations in western and northern Europe by indigenous Mesolithic peoples. This is consistent with the Neolithic migration hypothesis (Ammerman and Cavalli-Sforza 1984; Cavalli-Sforza 2002). However, this first-order level of molecular resolution does not readily reflect apparent complexities in regional and local archaeological sequences. The archaeological records suggest that the large-scale clinal patterns of Hg E and Hg J reflect a mosaic of numerous small-scale, more regional population movements, replacements, and subsequent expansions overlying previous ranges. The recent findings of many biallelic markers, which subdivide these two haplogroups, give us the opportunity to investigate the contribution of different population movements that have spread Hg E and Hg J. Through analysis of the Alu insertion (YAP), the M174 and SRY4064 mutations, and the 12f2a deletion, we identified haplogroups D (YAP/M174), E (YAP/SRY4064), and J (12f2a) Y chromosomes in >2,400 males from 29 populations, mainly from Europe and the Mediterranean area but also from Africa and Asia. No subject belonged to the recently reported paragroup DE* (Weale at al. 2003), and only 6 belonged to the Asian-specific Hg D, whereas 501 were members of Hg E and 445 of Hg J. The survey of 36 biallelic markers in the Hg E and Hg J Y chromosomes allowed us to define the phylogenetic relationships of their numerous subclades (figs. 1 and 2) and to analyze their distributions in the various geographic areas (tables 1 and 2). In addition, the survey of eight microsatellites (figs. 3 and 4) in a subset of these samples allowed investigation of the relative dating of different subclades.

 

Figure 1
Phylogeny and frequency distributions of Hg E and its main subclades (panels A–G). The numbering of mutations is according to the Y Chromosome Consortium (YCC) (YCC ²⁰⁰²; Jobling and Tyler-Smith ²⁰⁰³). To the left of the phylogeny, the ages (in 1,000 years) of the boxed mutations are reported, with their SEs (Zhivotovsky et al. ²⁰⁰⁴). Because the procedure used is based on STR data, it actually estimates the ages of STR variation observed within the corresponding haplogroup in the studied populations. With the exception of the value relative to SRY₄₀₆₄ mutation, which as been calculated as T_D (with V₀=0) between the sister clades Hg E-P2 and Hg E-M33, the other values were estimated as the average squared difference (ASD) in the number of repeats between all current chromosomes of a sample and the founder haplotype, which has an expected value μt for single-step mutations (Thomas et al. ¹⁹⁹⁸) and wt for a general mutation scheme, where w is an average effective mutation rate at the loci, taken as 6.9×10^-4 per 25 years (Zhivotovsky et al. ²⁰⁰⁴) (microsatellite data available on request). In some cases, because of small sample sizes or long time passed since the occurrence of the mutation, the founder haplotype could not be reliably estimated as a modal haplotype. Therefore, we constructed it from modal alleles at single loci, although this can underestimate the age if the candidate founder haplotype differs from the real one. To make the computation of the P2 and M35 ages independent from those of their most-represented subclades, the STR variation observed at only the “asterisk” lineages (e.g., E-P2*) has been used. The M35 estimate is in agreement with those of Bosch et al. (²⁰⁰¹) and Cruciani et al. (²⁰⁰⁴ [in this issue]), obtained with different methods. The YAP insertion was studied as an amplified fragment-length polymorphism (Hammer and Horai ¹⁹⁹⁵). The other mutations were investigated in a hierarchical order by use of the denaturing high-performance liquid chromatography (DHPLC) methodology (Underhill et al. ²⁰⁰¹). Subhaplogroups observed in this study are illustrated by continuous lines, whereas subhaplogroups discussed elsewhere are indicated by dotted lines. For simplicity, the prefix “M” was omitted from the name of the marker mutations. Haplogroup-frequency surfaces were graphically computer reconstructed following the Kringing procedure (Delfiner ¹⁹⁷⁶) by use of the Surfer System (Golden Software) and the data reported in table 1.

 Figure 2
Phylogeny and frequency distributions of Hg J and its main subclades (panels A–F). The numbering of mutations is according to the YCC (YCC ²⁰⁰²; Jobling and Tyler-Smith ²⁰⁰³). To the left of the phylogeny, the ages (in 1,000 years) of the boxed mutations are reported, with their SEs (Zhivotovsky et al. ²⁰⁰⁴). With the exception of the age relative to the 12f2 mutation, which has been estimated as T_D (with V₀=0) between the combined data of the two sister clades Hg J-M267 and Hg J-M172, the other values have been determined as ASD, as described in figure 1. The 12f2a marker was examined as an RFLP by Southern blotting (Passarino et al. ¹⁹⁹⁸); the other mutations were investigated in hierarchical order by use of DHPLC methodology (Underhill et al. ²⁰⁰¹). Three new mutations, M327, M280, and M390, were found in this study. M327 is a T→C transition at np 404 within the STS containing mutation M92, M280 is a G→A transition at np 330 within the STS containing the mutation M67, and M390 is an A insertion after nt 175 in the STS containing the M365 mutation. Conventions used are the same as for figure 1. The frequency surfaces were drawn using the data reported in table 2 and, for Hg J (panel A), also the data from Rosser et al. (²⁰⁰⁰), Quintana-Murci et al. (²⁰⁰¹), and Scozzari et al. (²⁰⁰¹

 Table 1
Population Frequencies of Hg E and Its Subclades

 

	Hg E		Frequency of E Subhaplogroupb													Hg D
Population/Regiona	No.	%	2*c	58	191	154	P2*	329	35*	123	78	81	281	33	75	No.	%
Arab (Morocco)d (49)	37	75.5									42.9	32.6
Arab (Morocco)e (44)	32	72.7	6.8						2.3		11.4	52.3
Berber (Morocco)d (64)	55	85.9	4.7								10.9	68.7		1.6
Berber (north-central Morocco)e (63)	55	87.3	9.5						7.9		1.6	65.1		3.2
Berber (southern Morocco)e (40)	35	87.5	2.5						7.5		12.5	65.0
Saharawish (North Africa)e (29)	24	82.7	3.4									75.9		3.4
Algerian (32)	21	65.6							3.1	3.1	6.3	53.1
Tunisian (58)	32	55.2	3.4						3.4	5.2	15.5	27.6
Malif (44)	37	84.1	20.5									29.5		34.1
Burkina Fasod (106)	105	99.1	67.9	1.9	13.2				.9					3.8	11.3
North Cameroond (152)	69	45.4	20.3		12.5						1.3			7.9	3.3
South Cameroond (89)	83	93.3	43.8		40.4	9.0
Senegaleseg (139)	136	97.8	80.6		.7		2.9		5.0		.7	.7		5.0	2.9
Bantu (South Africa)f (53)	44	83.0	54.7	5.7		3.8	1.9		1.9						15.1
Khoisan (South Africa)d (90)	59	65.6	31.1		11.1	1.1			16.7						5.6
Sudanf (40)	12	30.0									17.5	5.0		2.5	5.0
Ethiopian (Oromo)g (78)	62	79.5					12.8	2.6	19.2	5.1	35.9		2.6		1.3
Ethiopian (Amhara)g (48)	22	45.8					10.4		10.4	2.1	22.9
Iraqi (218)	20	9.2	.9							2.8	5.5
Lebanese (42)	8	19.0								4.8	11.9	2.4
Ashkenazim Jewish (77)	14	18.2							1.3	11.7	5.2
Sephardim Jewish (40)	12	30.0							2.5	10.0	12.5	5.0
Turkish (Istanbul) (46)	6	13.0								2.2	8.7	2.2
Turkish (Konya) (117)	17	14.5								1.7	12.8					1	.9
Georgian (41)	0	.0
Balkarian (southern Caucasus) (39)	1	2.6									2.6
Northern Greek (Macedonia) (59)	12	20.3								1.7	18.6
Greek (84)	20	23.8								2.4	21.4
Albanian (44)	11	25.0									25.0
Croatian (57)	5	8.8								1.8	7.0
Hungarian (53)	5	9.4								1.9	7.5
Ukrainian (93)	8	8.6								1.1	7.5
Polish (99)	4	4.0									4.0
Italian (north-central Italy) (56)	6	10.7									10.7
Italian (Calabria 1) (80)	18	22.5							1.3	2.5	16.3	1.3		1.3
Italian (Calabria 2)h (68)	16	23.5							1.5	13.2	5.9			2.9
Italian (Apulia) (86)	12	13.9								2.3	11.6
Italian (Sicily) (55)	15	27.3							5.5	3.6	12.7	5.5
Italian (Sardinia) (139)	7	5.0							.7	1.4	2.9
Dutch (34)	0	.0
Bearnais (27)	1	3.7									3.7
French Basque (45)	0	.0
Spanish Basque (48)	1	2.1										2.1
Catalan (33)	2	6.1									3.0	3.0
Andalusian (76)	7	9.2									3.9	5.3
Andalusiane (37)	4	10.8							2.7		2.7	5.4
Hindu (India) (47)	0	.0
Tharu (Nepal) (98)	0	.0														4	4.1
Chinese (65)	0	.0														1	1.5

 

Table 2

Population Frequencies of Hg J and Its Subclades

			Frequency of J subhaplogroupb
	Hg J		M172											M267c
Population/Regiona	No.	%	172*	158	12*	102*	280	47	67*	92*	327	68	Total %	267*	62	365	390
Arab (Morocco)d (49)	20	20.4	10.2										10.2	10.2
Arab (Morocco)e (44)	7	15.9											2.3	13.6
Berber (Morocco)d (64)	4	6.3												6.3
Berber (Morocco)e (103)	11	10.7											2.9	7.8
Saharawish (North Africa)e (29)	5	17.2												17.2
Algerian (20)	7	35.0												35.0
Tunisian (73)	25	34.2	1.4		1.4	1.4							4.1	30.1
Sudanf (40)	0	.0
Ethiopian (Amhara) (48)	17	35.4							2.1				2.1	33.3
Ethiopian (Oromo) (78)	3	3.8				1.3							1.3	2.6
Iraqi (156)	79	50.6	10.2			2.6		2.6	4.5	1.3		1.3	22.4	28.2
Lebanese (40)	15	37.5	20.0						2.5	2.5			25.0	10.0			2.5
Muslim Kurdg (95)	38	40.0											28.4	11.6
Palestinian Arabg (143)	79	55.2											16.8	38.4
Bedouing (32)	21	65.6											3.1	62.5
Ashkenazim Jewish (82)	31	37.8	12.2			1.2			4.9	4.9			23.2	14.6
Sephardim Jewish (42)	17	40.5	23.8			2.4			2.4				28.6	11.9
Turkish (Istanbul) (73)	18	24.7	11.0						2.7	4.1			17.8	5.5		1.4
Turkish (Konya) (129)	41	31.8	17.8			.8		.8	3.1	4.6	.8		27.9	3.1		.8
Georgian (45)	15	33.3	8.9					2.2	13.3	2.2			26.7	4.4		2.2
Balkarian (southern Caucasus) (16)	4	25.0	12.5			6.3			6.3				25.0
Northern Greek (Macedonia) (56)	8	14.3	3.6			5.4			3.6				12.5	1.8
Greek (92)	21	22.8	4.3			6.5	2.2		4.3	3.3			20.6	2.2
Albanian (56)	13	23.2				14.3			3.6	1.8			19.6	3.6
Croatian (48)	3	6.2				6.2							6.2
Hungarian (49)	1	2.0							2.0				2.0
Ukrainian (82)	6	7.3	2.4			2.4			1.2	1.2			7.3
Polish (97)	1	1.0				1.0							1.0
Italian (north-central Italy) (52)	14	26.9	5.8			9.6			9.6	1.9			26.9
Italian (Calabria 1) (57)	14	24.6	14.0			1.8			3.5	3.5			22.8	1.8
Italian (Calabria 2)h (45)	9	20.0	4.4						8.9	6.6			20.0
Italian (Apulia) (86)	27	31.4	16.3			3.5			2.3	7.0			29.1	2.3
Italian (Sicily) (42)	10	23.8	11.9						2.4	2.4			16.7	7.1
Italian (Sardinia) (144)	18	12.5	2.8			2.1			2.8	2.1			9.7	2.8
Dutch (34)	0	.0
Bearnais (26)	2	7.7	3.8			3.8							7.7
French Basque (44)	6	13.6	13.6										13.6
Spanish Basque (48)	0	.0
Catalan (28)	1	3.6							3.6				3.6
Andalusian (93)	8	8.6	2.2			1.1			3.2	1.1			7.5	1.1
Hunza (Pakistan)f (38)	5	13.2	2.6			7.9							10.5	2.6
Pakistan-Indiaf (88)	21	23.9	3.4	1.1	2.3	3.4		1.1		4.5			15.9	7.9
Hindu (India) (76)	4	5.3	2.6			1.3						1.3	5.3
Tharu (Nepal) (50)	7	14.0	8.0			6.0							14.0
Central Asiaf (184)	40	21.7	6.5	.5		2.2		.5	1.1	.5		.5	11.9	9.2	.5

 Figure 3

Networks of the STR haplotypes of the main subhaplogroups of Hg E. These networks were obtained by the analysis of a subset of the samples for the following microsatellites: YCAIIa, YCAIIb (Mathias et al. ¹⁹⁹⁴), DYS19, DYS389, DYS390, DYS391, and DYS392 (Roewer et al. ¹⁹⁹⁶). The phylogenetic relationships between the microsatellite haplotypes were determined using the program NETWORK 2.0b (Fluxus Engineering). Networks were calculated by the median-joining method (=0) (Bandelt et al. ¹⁹⁹⁵), weighting the STR loci according to their relative variability in Hg E and, with the exception of E-M81, after having processed the data with the reduced-median method. Circles represent the microsatellite haplotypes. Unless otherwise indicated by a number on the pie chart, the area of the circles and the area of the sectors are proportional to the haplotype frequency in the haplogroup and in the geographic area indicated by the color. The smallest circle of each network corresponds to one Y chromosome. The shaded area in E-M78 indicates the branch characterized by the DYS392-12 allele.

 Figure 4

Network of the STR haplotypes of the main subhaplogroups of Hg J. These networks were obtained by the analysis of a subset of the samples for the following microsatellites: YCAIIa, YCAIIb (Mathias et al. ¹⁹⁹⁴), DYS388 (Thomas et al. ¹⁹⁹⁹), DYS19, DYS389, DYS390, DYS391, and DYS392 (Roewer et al. ¹⁹⁹⁶), by the same procedures used for Hg E (fig. 3). Apart from the YCAII system in Hg J-M267, which was considered as a stable marker in this haplogroup (see text), the STR loci were weighted according to their relative variability in Hg J. The most complex networks, J-M267* and J-M172*, were calculated by the median-joining method (=0) on the preprocessed data with the reduced-median method; the other networks were calculated by using only the reduced-median algorithm. The shaded area in J-M267* indicates the branch characterized by the YCAIIa-22/YCAIIb-22 motif. For the areas of the circles and the sectors, see figure 3. The expansion time of this branch was calculated using T_D (Zhivotovsky ²⁰⁰¹), which gives 8.7 and 4.3 ky, respectively, for the earliest and the latest bounds of the expansion time. The former estimate was calculated by using the variance in the number of repeats of the remaining six loci, assuming a variance at the beginning of population separation (V₀) equal to zero, and thus gives an upper bound for the T_D (Zhivotovsky ²⁰⁰¹). The latter assumes a linear approximation of the within-population variance in repeat scores as a function of time and takes a predicted value of V₀ prior to population split; because the linearity can be achieved in a case of infinite population size only and because each survived haplogroup started from one individual and could maintain small size for a long time, the linear approximation overestimates V₀ and thus might be considered as a lower bound for divergence times (L.A.Z., unpublished method).

Hg E (fig. 1A) is observed in Africa, Europe, and the Near East and includes the subhaplogroups E-M33, E-M75, and the most widespread subclade, E-P2. The latter includes three clusters, two of which, E-M2 and E-M35, are the most widespread. Haplogroups E-M33 (fig. 1B), E-M75 (fig. 1C), and the not-shown E-P2* and E-M2 are virtually absent in European populations and appear to be geographically restricted to sub-Saharan Africa. The E-P2* lineages were observed mainly in Ethiopians, whereas E-M2, which is considered a signature of the Bantu expansion (Hammer et al. 1998; Passarino et al. 1998; Scozzari et al. 1999), shows its highest frequency (>80%) in Senegal and has been sporadically observed in North Africa and Iraq. E-M35 (fig. 1D) has been found in Africa, the Near East, and Europe, where it is believed to have arrived in Neolithic times (Hammer et al. 1998; Semino et al. 2000). In particular, from among its subgroups, E-M78 (fig. 1E) is present in Europe, the Middle East, and North and East Africa. However, whereas no preferential YCAII microsatellite motif is observed in the Middle East, prevalent associations with YCAIIa21-YCAIIb19 in Europe and YCAIIa22-YCAIIb19 in Africa are found. E-M81 (fig. 1F) is almost absent in Europe (with the exception of Sicily and Iberia) and the Middle East but characterizes the majority of the Y chromosomes of populations from northwestern Africa. E-M123 (fig. 1G) is spread in the Near East and is also observed in North Africa and Europe but does not reach the western European regions. E-M281 and E-M329 are geographically restricted, having been seen only in Ethiopians (two subjects each). The remaining 37 E-M35* Y chromosomes were found mainly in Africa, with a high frequency in the Ethiopians and the Khoisan.

Both phylogeography and microsatellite variance suggest that E-P2 and its derivative, E-M35, probably originated in eastern Africa. This inference is further supported by the presence of additional Hg E lineal diversification and by the highest frequency of E-P2* and E-M35* in the same region. The distribution of E-P2* appears limited to eastern African peoples. The E-M35* lineage shows its highest frequency (19.2%) in the Ethiopian Oromo but with a wider distribution range than E-P2*. Indeed, it is also found at high frequency (16.7%) in the Khoisan of South Africa (Underhill et al. 2000; Cruciani et al. 2002) (suggesting, once again, their ancient relationship with Ethiopians) and observed in southern Europe (present study). It is interesting that both E-P2* and E-M35* and their derivatives, E-M78 and E-M123, exhibit in Ethiopians the 12-repeat allele at the DYS392 microsatellite locus, an allele scarcely seen (Y-Chromosome STR Database), especially in other haplogroups and other populations (A.S.S.-B., unpublished data). In addition, the Ethiopian DYS392-12 allele is usually associated with the unusually short DYS19-11 allele, which is typical of this area. These findings are not easily explained. One possible scenario is that an ancient differentiation of the E-P2 haplogroup occurred in loco (East Africa). However, this also implies a low mutability of the associated microsatellite motif (DYS392-12/DYS19-11). Alternatively, the microsatellite motif may be due to homoplasy.

The first scenario is more likely, since this unique microsatellite haplotype occurs in E-P2*, E-M35*, and E-M78 but is almost absent in all other haplogroups and populations. In addition, the high stability of the DYS392 locus (Brinkmann et al. 1998; Nebel et al. 2001) and of the shorter alleles of DYS19 (Carvalho-Silva et al. 1999) has been reported elsewhere. Moreover, the observation that the derivative E-M78 displays the DYS392-12/DYS19-11 haplotype suggests that it also arose in East Africa. This is illustrated by the microsatellite network (fig. 3, shaded area), which reveals that the Ethiopian branch harboring DYS392-12 is not shared with either Near Eastern or European populations. The very low frequency of E-M123 in Ethiopia does not allow any inferences about the origin of this clade. The network of E-M78 and that of E-M123 are in agreement with the hypothesis of their ancient presence in the Near East and their subsequent expansion into the southern Balkans. The divergence time (TD) (Zhivotovsky 2001) between the Near East and European lineages has been estimated to a range of 7–14 thousand years (ky) ago. Cinnioğlu et al. (2004) found a high degree of variance of E-M123 in Turkey, which has been interpreted as being due to multiple founders rather than a single early dispersal event that has remained geographically circumscribed. E-M81 has the lowest variance and a compact network (fig. 3), indicating either its relatively recent origin followed by expansion or its recent expansion after a bottleneck. In Europe, this clade is restricted to the southernmost regions, such as Iberia and Sicily, and the absence of microsatellite variation suggests a very recent arrival from North Africa, consistent with previous observations (Bosch et al. 2001). The frequency pattern and the microsatellite network of E-M2(xM191) (fig. 3) indicate a West African origin followed by expansion, a result that is in agreement with the findings of Cruciani et al. (2002).

The 12f2a mutation, which characterizes haplogroup J, was observed in 445 subjects. Hg J harbors two main clades (see phylogeny in fig. 2), J-M267 (Cinnioğlu et al. 2004) and J-M172. J-M172 is the more frequent and currently differentiates into eight subhaplogroups defined by mutations M12/M102, M47, M67/M92, M68, M137, M158, M339, and M340, four of which occur at informative frequencies. The less-heterogeneous clade J-M267 includes all of the other 12f2a Y chromosomes that were reported elsewhere as Eu10 (Semino et al. 2000). Its current level of subdivision includes five scarcely represented subclusters defined by mutations M62, M365, M367/M368, and M369 (Cinnioğlu et al. 2004) and by the new mutation M390. Similar to Hg E, different geographic distributions are displayed by the various subhaplogroups of J (fig. 2). J-M172 (fig. 2C), which occurs as frequently as J-M267 (fig. 2B) in some Middle Eastern populations, is the more prevalent in Europe. Among its subclades, J-M137, J-M158, J-M339, and J-M340 were reported elsewhere as single observations (Underhill et al. 2000; Cinnioğlu et al. 2004) and have not been observed in this study. Likewise, J-M47 and J-M68 characterize very few Near Eastern and Asian samples. However, J-M12 and J-M67 and their derivatives are informative, being diffused in Europe and observed also in Asia. J-M12 is almost totally represented by its sublineage J-M102, which shows frequency peaks in both the southern Balkans and north-central Italy (fig. 2D). The history underlying this apparent affinity remains uncertain. J-M67 (fig. 2E) includes J-M67* lineages (not shown), which are most frequent in the Caucasus, and J-M92, which indicates affinity between Anatolia and southern Italy (fig. 2F). Finally, the J-M172* lineages display a decreasing frequency gradient from the Near East toward western Europe and strongly contribute to the overall gradient of Hg J. J-M267 is notable, since this haplogroup shows its highest frequencies in the Middle East, North Africa, and Ethiopia (fig. 2B) and its lowest in Europe, having been observed only in the Mediterranean area. Of its five subhaplogroups, only two have been observed: the J-M365 (in two Turks and one Georgian) and the new subclade J-M390 (in one Lebanese).

The extent of differentiation of Hg J, observed both with the biallelic and microsatellite markers, points to the Middle East as its likely homeland. In this area, J-M172 and J-M267 are equally represented and show the highest degree of internal variation, indicating that it is most likely that these subclades also arose in the Middle East. However, their different frequencies in different Middle Eastern countries and in Europe suggest distinct demography processes, possibly in population groups that underwent different temporal expansions. This is especially true for J-M172. The majority of its lineages are undifferentiated and thus potentially paraphyletic (fig. 4). Although J-M172* encompasses most of the M172 Y chromosomes in continental Europe and India (Kivisild et al. 2003; present study), their degree of affinity and shared history remain uncertain. The J-M67*, J-M92, and J-M102 representatives reflect more distinctive origins and dispersal patterns. Whereas J-M67* and J-M92 show higher frequencies and variances in Europe (0.40 and 0.32, respectively) and in Turkey (0.32 and 0.30, respectively [Cinnioğlu et al. 2004]) than in the Middle East (0.17 and 0.09, respectively), J-M12(M102) shows its maximum frequency in the Balkans. In spite of the relative high value of variance of this haplogroup in Turkey (Cinnioğlu et al. 2004)—which, however, could be due to multiple arrivals—the pattern of distribution and the network of J-M12(M102) (figs. 2 and 4) are consistent with its diffusion in Europe from the southern Balkans. On the contrary, J-M67* and J-M92 could have arrived in Europe from Anatolia via the Bosphorus isthmus, as well as by seafaring Neolithic populations who reached southern Italy. J-M67* and J-M92 could represent, at least in part, the Y-chromosome component that King and Underhill (2002) found to correlate with the distribution, from Anatolia toward Europe, of archaeological painted pottery and anthropomorphic figurines. On the other hand, J-M67– and J-M12–related lineages have been observed in Pakistan and India; thus, they probably have marked other migratory events, but the small number of J subclades in these regions (Underhill et al. 2000; Kivisild et al. 2003; present study) does not allow an evaluation of the mode and time of their arrival.

Southern Italy (Apulia and Calabria) contains sites of the early Neolithic period (Whitehouse 1968), but we know from history that these regions were subsequently colonized by the Greeks (Peloponnesians). To test the relative contribution of Greek colonists versus putative earlier Neolithic settlers, an admixture analysis (Bertorelle and Excoffier 1998) was performed, using E-M78 and J-M172(xM12) as signatures of Greek and Anatolian lineages, respectively. The Anatolian source population was based on 523 Turks, of whom 118 were J-M172(xM12) and 25 were E-M78 (Cinnioğlu et al. 2004). The Greek population comprised 36 Peloponnesian samples, 5 of which were J-M172(xM12) and 17 of which were E-M78 (R.K., unpublished data). In spite of the small Peloponnesian sample size, the high E-M78 frequency (47%) observed here is consistent with that (44%) independently found in the same region (Di Giacomo et al. 2003) for the YAP chromosomes harboring microsatellite haplotypes (A. Novelletto, personal communication) typical of Hg E-M78 (Cruciani et al. 2004 [in this issue]; present study). The admixture analysis yielded an admixture proportion from Greece of 0.07±0.15 for the Calabrian samples and of 0.22±0.15 for the Apulian samples. SD was determined by bootstrapping 1,000 replicates.

The TD of the two sister clades J-M267 and J-M172 was estimated, with V0=0, and turned out to be 31.7 ky (see phylogeny in fig. 2). This estimate, however, is not easily interpretable, because such old haplogroups are differently represented in different regions where they probably underwent multiple bottlenecks. The lower internal variance of J-M267 in the Middle East and North Africa, relative to Europe and Ethiopia, is suggestive of two different migrations. In the absence of additional binary polymorphisms allowing further informative subdivision of J-M267, the YCAII microsatellite system provides important insights. The majority of J-M267 Y chromosomes harbor the single-banded motif YCAIIa22-YCAIIb22 in the Middle East (>70%) and in North Africa (>90%), whereas this association is much less frequent in Ethiopia and only sporadically found in southern Europe. Considering the distribution of this YCAII single-banded pattern—which, besides the usual stepwise mutational mechanism, could be due to a stable mutational event (one locus deletion or a single-nucleotide mutation in the primer sequence)—we suggest that the motif YCAIIa22-YCAIIb22 potentially characterizes a monophyletic clade of J-M267. A comparable situation is observed within Hg I-M170, in which the single-banded haplotype YCAIIa21-YCAIIb21 parallels a biallelic marker (O.S., unpublished data).

According to this interpretation, the first migration, probably in Neolithic times, brought J-M267 to Ethiopia and Europe, whereas a second, more-recent migration diffused the clade harboring the microsatellite motif YCAIIa22-YCAIIb22 in the southern part of the Middle East and in North Africa. In this regard, it is worth noting that the median expansion time of the J-M267-YCAIIa22-YCAIIb22 clade was estimated to be 8.7–4.3 ky, by use of the TD approach (see fig. 4 legend), and that this clade includes the modal haplotype DYS19-14/DYS388-17/DYS390-23/DYS391-11/DYS392-11 of the Galilee (Nebel et al. 2000) and of Moroccan Arabs (Bosch et al. 2001). These results are consistent with the proposal that this haplotype was diffused in recent time by Arabs who, mainly from the 7th century a.d., expanded to northern Africa (Nebel et al. 2002).

In conclusion, high-resolution Y-chromosome haplotyping and particular microsatellite associations reveal regional population differentiations, an East Africa homeland for E-M78, and recent gene-flow episodes consistent with the Neolithic in Europe. In particular, the spatial distributions of J-M172*, J-M267, E-M78, and E-M123 indicate expansions from the Middle East toward Europe that most likely occurred during and after the Neolithic, that of J-M102 illustrates population expansions from the southern Balkans, and that of E-M81 reveals recent gene flow from North Africa. Distinct histories of J-M267* lineages are suggested: an expansion from the Middle East toward East Africa and Europe and a more-recent diffusion (marked by the YCAIIa-22/YCAIIb-22 motif) of Arab people from the southern part of the Middle East toward North Africa.

Another North African Y chromosome study, a little old now.

High-Resolution Analysis of Human Y-Chromosome Variation Shows a Sharp Discontinuity and Limited Gene Flow between Northwestern Africa and the Iberian Peninsula

 Elena Bosch,1,* Francesc Calafell,1 David Comas,1 Peter J. Oefner,2 Peter A. Underhill,3 and Jaume Bertranpetit1
In the present study we have analyzed 44 Y-chromosome biallelic polymorphisms in population samples from northwestern (NW) Africa and the Iberian Peninsula, which allowed us to place each chromosome unequivocally in a phylogenetic tree based on >150 polymorphisms. The most striking results are that contemporary NW African and Iberian populations were found to have originated from distinctly different patrilineages and that the Strait of Gibraltar seems to have acted as a strong (although not complete) barrier to gene flow. In NW African populations, an Upper Paleolithic colonization that probably had its origin in eastern Africa contributed 75% of the current gene pool. In comparison, ~78% of contemporary Iberian Y chromosomes originated in an Upper Paleolithic expansion from western Asia, along the northern rim of the Mediterranean basin. Smaller contributions to these gene pools (constituting 13% of Y chromosomes in NW Africa and 10% of Y chromosomes in Iberia) came from the Middle East during the Neolithic and, during subsequent gene flow, from Sub-Saharan to NW Africa. Finally, bidirectional gene flow across the Strait of Gibraltar has been detected: the genetic contribution of European Y chromosomes to the NW African gene pool is estimated at 4%, and NW African populations may have contributed 7% of Iberian Y chromosomes. The Islamic rule of Spain, which began in a.d. 711 and lasted almost 8 centuries, left only a minor contribution to the current Iberian Y-chromosome pool. The high-resolution analysis of the Y chromosome allows us to separate successive migratory components and to precisely quantify each historical layer.
The systematic search for polymorphisms in the human Y chromosome, both by conventional techniques and by denaturing high-performance liquid chromatography (DHPLC), is producing a large number of new markers (Underhill et al. 1997, 2000; Shen et al. 2000), overcoming the initial dearth of available polymorphisms on that chromosome (Dorit et al. 1995). Among all the different types of Y-chromosome polymorphisms, base substitutions and insertion/deletion polymorphisms have proved to be especially useful in the reconstruction of the phylogeny of the 30-Mb Y-chromosome nonrecombining region. Given their nature, these mutations have probably arisen only once in evolutionary history and have created biallelic polymorphisms. In the absence of recurrence, the typing of such markers in nonhuman primates allows us to determine which is the ancestral allele. The knowledge of the ancestral and derived states of these markers, together with the fact that most of the Y chromosome does not recombine, allows the direct application of parsimony criteria to obtain its phylogeny. Underhill et al. (2000) developed a new set of markers and typed a large set of samples from different worldwide population, providing a well-established structure for Y-chromosome phylogeny and a wide context of very detailed information on Y-chromosome variation, against which any particular new population can be evaluated. A specific analysis of Europe (Semino et al. 2000) has shown the possibilities of the application of this marker set to a continental framework. Furthermore, because of this well-established phylogeny, we are able to characterize new populations by means of a fast hierarchical approach, in which markers are successively typed from the top to the bottom (from the root toward the branch tips) of the phylogenetic tree, as needed. Given the fine degree of paternal-lineage dissection achieved, the proper knowledge of the worldwide distribution and of patterns of variation of the haplotypes that constitute this phylogeny will pave the way for the elucidation of the patterns of male migration and admixture, among present and past human populations.

 

In addition to the geographical proximity of northwestern (NW) Africa and the Iberian Peninsula, which are separated only by the 15-km-wide Strait of Gibraltar, both regions are linked by historical events involving population movement. During the Upper Paleolithic (known as the “Late Stone Age” in the study of African prehistory), the Ibero-Maurusian industry, spanning the time period of 22,000–9,500 years ago (ya) (Newman 1995), is found throughout northern Africa. The prefix “Ibero-” refers to the presumption that this culture extended into Iberia, although an origin in the Nile River valley is now widely accepted (Camps 1974). The Ibero-Maurusian culture was followed, in the NW African Mesolithic, by the Capsian industry (10,000–4,700 ya; Desanges 1990). The Capsian culture persists well into the Neolithic (which began ~5,500 ya), a fact that may indicate a persistence of the Mesolithic population and a cultural adoption of agriculture and husbandry with some Neolithic admixture, rather than a replacement by Neolithic populations originating in the Middle East. In general terms, prehistoric culture changes in NW Africa were quite independent of the change dynamics on the European shores of the Mediterranean. In Iberia, the first Upper Paleolithic settlements appear as early as 40,000 ya. Later local developments, until the spread of the Neolithic, followed traditions having European or northern Mediterranean distributions.

NW Africa enters history with the Phoenicians, who, originating in the Middle East, founded Carthage in 814 b.c. and established commercial relations with the local populations, who were the ancestors of the current Berbers. The Roman geographers documented the native kingdoms of the Mauri, Numidae, Gaetali, and Libii, all of whom were subsequently conquered, in ~150 b.c.–a.d. 50, by the Romans, who established themselves, probably with a limited demographic impact, along a 100-km-wide strip along the Mediterranean coast. This has a parallel in the history of Iberia, where Phoenicians and Greeks established trading posts, and where the local populations (Iberians and Celts) were later conquered (starting ~200 b.c.) by the Romans. Iberia (“Hispania”) then became a province of the Roman Empire.

During the 7th century a.d., the Arabs conquered northern Africa from east to west and spread their language and religion throughout the native Berber population. Although the cultural and political impact of this invasion changed the history of NW Africa profoundly, a precise estimation of the demographic contribution of the Arabs to NW Africa is not available. In a.d. 711, Berber troops under Arab leadership (Hitti 1990) crossed to the Iberian Peninsula, which they subsequently conquered. That date marks the start of an 8-century period during which the Iberian Peninsula was divided into the Christian kingdoms to the north and the Islamic kingdoms in the south. The border between the two moved southward until 1492, when the last Islamic kingdom was conquered. The demographic contribution of NW African populations to Iberia is not known precisely; it may have been on the order of tens of thousands of individuals in a total Iberian population of a few million (McEvedy and Jones 1978). After the first conquest of Iberia, two main Berber invasions swept through the peninsula: the Almoravids (a.d. 1056–1147) and the Almohades (a.d. 1121–1269). The empire founded by the former group extended into Africa as far south as present-day Senegal and Mali (Kasule 1998).

After 1492, first Jews and then Moslems were forced by the Spanish rulers to either convert to Catholicism or leave the country. Most of those who were expelled took refuge in NW Africa. However, the population substrate of the Moslem group is not well known; the extent to which this group was composed of converted Iberians rather than of the descendants of the Islamic invaders is difficult to ascertain.

In the present study, we have typed 44 biallelic polymorphisms and 8 microsatellites (also known as “short tandem repeats,” or “STRs”), to define the main Y-chromosome lineages in NW Africa and the Iberian Peninsula, in a well-established phylogeographical frame (Underhill et al. 2000), as well as to attempt to estimate the dates of both ancient and recent events in the history of those populations. Several hypotheses regarding population history are tested, such as those concerning the extent to which the Paleolithic genetic background may still be present in both regions, as well as the impact of the Neolithic wave of advance; we also quantify any gene flow between these two regions and from external populations into these regions. The present results are contrasted with those of previous analyses of classical polymorphisms (i.e., blood groups and protein polymorphisms), Alu-insertion polymorphisms, mtDNA control-region sequences, and other Y-chromosome studies. Previous analyses, of a smaller set of Y-chromosome polymorphisms in populations from NW Africa and the Iberian Peninsula, have been published by Bosch et al. (1999), who emphasized gene genealogy rather than population history, and by Rosser et al. (2000), who typed 11 biallelic polymorphisms in a broad survey of western-Eurasian populations and found that a principal-component analysis of haplotype frequencies separated NW African populations from European and Middle Eastern populations.
 
Different autochthonous samples from NW Africa and the Iberian Peninsula were typed. The NW African sample included blood from 29 Saharawis, 40 southern Moroccan Berbers, 44 Moroccan Arabs, and 63 north-central Moroccan Berbers. Samples from the Iberian Peninsula included blood from 37 Andalusians, 16 Catalans, and 44 Basques; the Basque individuals were also included in the study by Underhill et al. (2000). Appropriate informed consent was obtained from all participants in this study, and information about the geographical origin of their four grandparents and about their first language was recorded. DNA was extracted from fresh blood by standard phenol-chloroform protocols.
Polymorphism Typing
All samples in this study were characterized by means of a top-down approach, in which the markers indicated in figure 1 were successively typed, in hierarchical order, according to their position in the genealogy given by Underhill et al. (2000). The typing methods in our analysis would allow us to identify almost all haplotypes described by Underhill et al. (2000). Thus, the original haplotype notation of Underhill et al. (2000) has been kept.
 DHPLC was used to type all biallelic markers, with the exception of YAP (also known as “M1”). Marker information such as primer sequences and PCR conditions for their amplification, whether alleles are ancestral or derived, as well as additional details for their typing conditions by DHPLC, have been provided by Underhill et al. (1997, 2000). YAP was assayed as described by Hammer and Horai (1995). It should be noted that a subset of the polymorphisms used in the present study has been typed in a number of European populations (Semino et al. 2000) and that a different notation has been given to those haplotypes: H22 is termed “Eu2”; H35, H36, and H38 are subsumed under “Eu4”; H52, H50, H58, and H71 are termed “Eu7,” “Eu8,” “Eu9,” and “Eu10,” respectively; H88 is termed “Eu15;” H101, H102, H103, and H104 are subsumed under “Eu18”; and, finally, H108 is termed “Eu19.”

Data for eight STRs (DYS388, DYS19, DYS390, DYS391, DYS392, DYS393, DYS389I, and DYS389II) were available for almost all chromosomes in the sample (Bosch et al. 1999, and additional typings reported here). Complete haplotypes (biallelic markers and STRs) are available from the authors.

Data Analysis
Haplotype-frequency differences among populations from NW Africa and the Iberian Peninsula were tested, by analysis of molecular variance (AMOVA), with the ARLEQUIN software package (Schneider et al. 2000). AMOVA was performed both separately, for NW African and Iberian populations, and as a joint analysis in which genetic variance was partitioned hierarchically as interregion (NW Africa vs. Iberia) variance, intraregion variance, and intrapopulation variance.
Coalescence analysis (Griffiths and Tavaré 1994) was used to test whether NW Africa and Iberia could be regarded as a panmictic unit, to estimate the amount of gene flow among the two regions, and to estimate the ages of M35, M78, and M81, under assumptions of both constant and exponential growth, by means of the Genetree program (available from the Genetree Web site). All biallelic polymorphisms constituting the haplotypes were given the same weight regardless of whether they were nucleotide substitutions or indels, given that they were all compatible with the infinite-sites model implemented in Genetree. First, the values of θ=Nμ (where N is effective population size and μ is mutation rate) that maximized the likelihood of the gene genealogy were obtained separately for the combined haplotype frequencies and for consideration of the two regions separately (in this case, maximum-likelihood estimates of the Nm migration parameter were also obtained, where m is migration rate per generation, were also obtained); next, the likelihood values obtained in the two scenarios were compared by a likelihood-ratio test, after application of the appropriate combinatorial factor (Bahlo and Griffiths 2000). Mutation-age estimates were obtained by use of the growth, θ, and migration parameters that maximized gene-genealogy likelihood and under the assumptions of an effective population size of 5,000 and a 20-year generation time. Genetree provides mutation-age estimates as multiples of θ; thus, either N or μ should be fixed a priori to transform ages in θ units to ages in generations. We fixed N at 5,000, which is close to the global value obtained by Goldstein et al. (1996). With our estimated θ values and with N set at 5,000, we obtained mutation rates of ~6×10-9 per nucleotide, a result that is consistent with the nuclear-genome average (Li et al. 1985). All Genetree program executions were run for 1,000,000 iterations.

Phylogenetic relations for STR haplotypes within the haplotypes defined by biallelic polymorphisms were depicted by means of reduced median networks (Bandelt et al. 1995), as implemented in the Network 2.0c program (available from www.fluxus-engineering.com).

Separation times, within Y-chromosome lineages, between NW African and Iberian chromosomes, were estimated from STR haplotypes, by means of the average square distance (ASD) (Thomas et al. 1998), by use of a mutation rate of 2.1×10-3 (Heyer et al. 1997; Jobling et al. 1999) and a generation time of 20 years.

 
Male-Lineage Structure of NW African and Iberian Populations
Haplotype frequencies in Moroccan Arabs, north-central Moroccan Berbers, southern Moroccan Berbers, and Saharawis are given in table 1. Haplotype-frequency differences among those populations were tested via AMOVA. Only 0.8% of the genetic variance was found to be due to haplotype-frequency differences among the populations (statistically not significantly different from 0; P=.169). H38, which, according to Underhill et al. (2000), belongs to haplotype group III, is the most common haplotype in NW Africa (64%), with its highest frequencies found within the Saharawis (76%). H71, which belongs to group VI, is the second-most-frequent haplotype (11%) in this area. Other haplotypes, found at lower frequencies, are H22 and H35 (6% each) and H36 (5%), all belonging to group III. The remaining haplotypes, which jointly represent 8% of the NW African Y chromosomes, are found at frequencies of <3%. The genetic homogeneity of NW African Y chromosomes points to a common origin, for all populations analyzed, independent of ethnicity or language (Arab or Berber). These data support the interpretation of the Arabization and Islamization of NW Africa, starting during the 7th century a.d., as cultural phenomena without extensive genetic replacement.
 

Haplotype frequencies for Basques, Catalans, and Andalusians are also given in table 1. AMOVA showed that 2% of the genetic variance was attributable to haplotype-frequency differences among them (statistically not significantly different from 0; P=.08). Pairwise population comparisons via AMOVA did not yield any values significantly different from 0. The most frequent haplotype in these populations is H104 (56%), which belongs to group IX. Haplotypes H102 and H103, which also belong to group IX, are found at frequencies of ~10%. The frequency of H71 (8%) is similar to that haplotype’s frequency in NW Africa. The proportion of haplotypes belonging to group VI (which includes H71) is slightly higher in Iberia (16%) than in NW Africa (14%). H35, H36, and H38, the only haplotypes found to belong to group III, constitute 5% of the Iberian Y chromosomes.

These results clearly show that the contemporary populations from both regions originated from different patrilineages: group III haplotypes prevail in NW Africa, whereas Iberian haplotypes belong mostly to group IX. The proportion of genetic variance that can be attributed to the difference between the NW African and Iberian populations is 35.2% (P=.024), the minimum possible value, given the number of populations and the permutation procedure employed to estimate statistical significance (Excoffier et al. 1992). Moreover, a coalescence analysis of the gene genealogy (Bahlo and Griffiths 2000), including haplotype frequencies in both regions, allowed us to reject the hypothesis that they behave jointly as a panmictic unit (χ2=271.69, 1 df, and P≈0, for constant population sizes; and χ2=266.47, 1 df, and P≈0, for expanding populations). The migration parameters that maximized gene-genealogy likelihood were Nm=1.25 from Iberia to NW Africa and Nm=2 from NW Africa to Iberia, which indicates that gene flow from NW Africa to Iberia may have been greater than that in the opposite direction. Other studies, which analyzed either classical genetic markers (Bosch et al. 1997; Kandil et al. 1999; Simoni et al. 1999), a set of up to 21 autosomal STRs (Bosch et al. 2000), or 11 polymorphic Alu insertions (Comas et al. 2000), showed important genetic differences between NW African and Iberian populations. Moreover, Bosch et al. (1997) and Simoni et al. (1999), analyzing, respectively, 13 and 20 populations from all around the Mediterranean basin, found that the sharpest genetic differences were between populations situated on either side of the Strait of Gibraltar. However, beyond the identification of differences in allele frequencies, the use of a system such as high-resolution biallelic-polymorphism Y-chromosome haplotypes, with a well-established gene genealogy and clear geographical structure, allows us to recognize patterns of origin and diffusion of haplotypes, which can then be used to quantify gene flow, as discussed below.

Neither the overall AMOVA nor any pairwise comparison among populations within either NW Africa or Iberia were significantly different from 0, implying that Y-chromosome biallelic haplotypes are highly homogeneous within each geographical region. Classical genetic markers, together with linguistic, paleoanthropological, and archaeological data, point to a Mesolithic (or older) origin of the Basques (Calafell and Bertranpetit 1994). However, this degree of differentiation is not reached by Y-chromosome polymorphisms (Hurles et al. 1999). For further discussion on how different kinds of genetic markers reflect the Basque differentiation, see the report by Comas et al. (2000).

 

Geographical and Historical Origins of Y-Chromosome Haplotypes in NW Africa and the Iberian Peninsula
Analysis of the worldwide distribution of Y-chromosome haplotypes may help to establish the putative origins of the haplotypes that contributed to the present NW African and Iberian populations. Figure 2 shows the detailed frequencies of haplotypes H22, H35, H36, H38, H58, H71, H102, H103, and H104, for the populations studied, as well as their worldwide distributions. This type of descriptive analysis allows us to recognize the haplotypes either as being autochthonous or as having originated elsewhere (in regions such as sub-Saharan Africa, Europe, or the eastern Mediterranean).
 

Specific founder effect for some NW African haplotypes: an Upper Paleolithic differentiation? Although group III haplotypes H35, H36, and H38 are found in eastern and southern Africa, southern Europe, and the Middle East, their overall frequencies in NW Africa are, by far, the highest reported to date (Semino et al. 2000; Underhill et al. 2000). This is particularly true for H38, which clearly constitutes the male population core of NW Africa. By contrast, haplotype H35 is found mainly in Ethiopia (22.7%) and Sudan (17.5%), and H36 is most frequent among Khoisans (10.3%) and Ethiopians (6.5%) (Underhill et al. 2000). Given that H36 is directly ancestral to H35 and H38 and is found at moderate frequencies in Ethiopia and in southern Africa, this branch of the haplotype phylogeny may have been introduced into NW Africa from eastern Africa. On the other hand, the dramatic discontinuity in frequencies of group III haplotypes (especially H38) that is seen in northern Africa suggests that such differences originated under strong genetic drift in small, isolated populations. Such demographic conditions were probably found only before the population surge brought by the Neolithic, which may have prevented further significant differentiation by drift (Cavalli-Sforza et al. 1994), as shown by computer simulations (Rendine et al. 1986; Calafell and Bertranpetit 1993).

Use of classical genetic markers has suggested (Bosch et al. 1997) that the NW African populations may have a sizeable Upper Paleolithic component. This hypothesized Upper Paleolithic expansion may be represented today by the descendants of the haplotypes that share mutation M35 and that are further characterized by M78 (H35) and M81 (H38). It remains to be resolved whether the latter two haplotypes arose independently from H36 or share a common ancestor, yet to be discovered, that distinguishes them from the remaining haplotypes derived from H36.

Assuming a constant population size, an infinite-sites model, and population subdivision between NW Africa and Iberia, we used Genetree (Griffiths and Tavaré 1994) to estimate the age of M35 (giving H36) to be 53,000±21,000 years ago (ya), that of M78 (giving H35) to be 16,000±10,000 ya, and that of M81 (giving H38) to be 32,000±11,000 ya. Under the more likely condition of population growth (Thomson et al. 2000), the respective estimated ages were 30,000±6,000 ya, 7,600±6,000 ya, and 19,000±4,000 ya. Hence, the expansion that brought the ancestors of H35 and H38 (or even those haplotypes themselves) into NW Africa could have happened at any time after 30,000 ya, and, more specifically, it could have happened during the Upper Paleolithic. However, confidence intervals for those dates are large, even without the uncertainty in the effective population size or in generation time. Thus, any interpretation derived from these dates should be regarded with caution. The lower limit for the differentiation event that brought H35 and H38 to such high frequencies in NW Africa is set by the demographic conditions that are compatible with this magnitude, as discussed above, as well as by the genetic evidence, from classical genetic markers (Bosch et al. 1997), that suggests a strong Paleolithic background in NW Africa.

Haplotypes H35, H36, and H38 were found at a low overall frequency (5%) in the Iberian populations. Eight-locus STR haplotypes for the five Iberian group III chromosomes showed that four of them were identical to group III chromosomes in NW Africans and that the fifth was one STR mutation step away. This is clearly depicted in the reduced median networks in figure 3a and b. Given the fast mutation rate of STRs, the extreme similarity between the STR haplotypes in the two regions can be explained only if Iberian and NW African group III chromosomes have a common origin. The time necessary to accumulate this small number of differences was estimated at 700±600 years. Thus, recent gene flow, rather than common ancestry in the distant past, may have brought those chromosomes from NW Africa into Iberia.
Neolithic Y-chromosome traces in NW Africa and Iberia. H58 and H71(fig. 2e and f) are part of group VI, which is defined by the presence of M89 and by the absence of M9 and subsequently derived mutations. These haplotypes constitute 10% of the Iberian and 13% of the NW African Y chromosomes and are likely to have spread, with the Neolithic expansions, from the Middle East (Semino et al. 2000). Both haplotypes include chromosomes with the derived 12f2-TaqI*8kb allele (Casanova et al. 1985), as confirmed by Bosch et al. (1999) in a subset of the samples in the present study. Y chromosomes bearing that allele have been found all around the Mediterranean basin, with higher frequencies in the Middle East, and have been interpreted to have spread with the Neolithic wave of advance (Semino et al. 1996; Rosser et al. 2000). A steep cline in the frequency of both H58 and H71, with maxima in the Middle East and frequencies declining with geographical distance from the Middle East, is evidence for a diffusion from the Middle East westward through Europe. The presence of H58 and H71 in both regions could be due to two, not necessarily mutually exclusive, historical processes: (1) the parallel, independent advance of the Neolithic expansion from the Middle East, along the northern and southern shores of the Mediterranean, and (2) an early arrival of the Neolithic in either NW Africa or Iberia and the subsequent crossing of the Strait of Gibraltar. Two independent regional analyses of large sets of classical polymorphisms (Bosch et al. 1997; Simoni et al. 1999) found parallel gradients of genetic differentiation, along the northern and southern shores of the Mediterranean, which make the first scenario more likely. In the present study, STR haplotypes for H58 and H71 chromosomes seemed to be associated with the history of their lineages rather than with population history. In a reduced median network (fig. 4), STR haplotypes clustered by lineage, and a main subdivision was linked to an additional biallelic polymorphism, 12f2 (Casanova et al. 1985; data from Bosch et al. 1999 and additional typings reported here), the 8-kb allele of which was found only in some H71 chromosomes and in all H58 chromosomes. Thus, 12f2*8kb seems to have appeared in the phylogeny after M89 but before M172 (fig. 1). Given that the 12f2*8kb allele was found more often in NW Africa than in Iberia, a comparison, between NW Africa and Iberia, of Y-chromosome STR haplotypes would be, in fact, a comparison of different lineages (as seen in fig. 4, where population origin is not random in the main sections of the network) and would confound attempts to differentiate the two scenarios. A confirmation by the Y chromosome would need to establish an independent correlation with distance to the Middle East. Unfortunately, samples from the appropriate populations are not yet available, particularly from countries along the southern shore, such as Libya and Tunisia.
The European Paleolithic background in Iberia. Group IX haplotypes (fig. 2g–i) are found in the Middle East and are most prevalent in Europe (Underhill et al. 2000). Group IX also contains three local Iberian haplotypes: H101, H102, and H103. The latter, which is defined by derived mutation M167 (also known as “SRY-2627”), is equivalent to Y-chromosome haplogroup 22 as described by Hurles et al. (1999). These authors examined haplogroup 22 worldwide and showed that it has a geographical distribution almost restricted to northern Iberia. Moreover, on the basis of the dating of microsatellite and minisatellite diversity within haplogroup 22, they suggested that it arose in Iberia a few thousand years ago.

Group IX is found at a low frequency (3%) in NW Africa. In Iberia, 56% of the Y chromosomes carry H104, which is found across Europe, with increasing frequencies toward the west; its defining mutation, M173, may have been introduced by the first Upper Paleolithic colonizations of Europe (Semino et al. 2000). It may not have been the only lineage introduced into Iberia during the Upper Paleolithic, but it seems to have been the only one that has persisted in the extant Iberian gene pool. Of five H104 NW African chromosomes, one had an STR haplotype identical to that in an H104 Iberian chromosome, one was one mutation step away from Iberian H104 chromosomes, and the remaining three were two mutation steps away. Moreover, the mean repeat-size difference within 53 H104 Iberian STR haplotypes was 2.8 (range 0–11). The phylogenetic relations among H104 STR haplotypes is shown by a reduced median network (fig. 3c), in which the NW African chromosomes appear to be clearly embedded within the Iberian diversity. The time necessary to accumulate the STR-allele differences between NW African and Iberian H104 chromosomes was estimated at 2,100±450 years. This close STR-haplotype similarity seems to indicate that H104 chromosomes found in NW Africa are a subset of the European gene pool and that they may have been introduced during historic times.
Sub-Saharan gene flow into NW Africa. H22 (defined by mutation M2, also referred to, by Seielstad et al. [1994], as “sY81”; see fig. 2a) and H28, which belong to group III, show a sub-Saharan distribution pattern (Seielstad et al. 1994; Hammer et al. 1997; Underhill et al. 2000). The highest frequency of H22 was found in Mali (30%), and the highest frequencies of H28 were found in southern (51%) and central Africa (57%). Both haplotypes together constitute 8% of the NW African Y chromosomes, and, given their geographical distribution, their presence in NW Africa can be interpreted as resulting from sub-Saharan gene flow. The NW African contact with the southern peoples was especially important during the Almoravid Berber expansion (a.d. 1056–1147), which reached as far south as present-day Senegal and Mali (Kasule 1998), and it has been maintained, until recently, by the trans-Saharan commercial routes.

mtDNA control-region sequence analysis (Rando et al. 1998) detected female-mediated gene flow from sub-Saharan Africa to NW Africa. In particular, 21.5% of the mtDNA sequences in a set of different NW African populations were found to belong to haplogroups L1, L2, and L3a, which constitute most of the sub-Saharan mtDNA sequences.

So far, our analyses have allowed a clear dissection of almost all NW African and Iberian paternal lineages into several components with distinct historical origins. In this way, the historical origins of the NW African Y-chromosome pool may be summarized as follows: 75% NW African Upper Paleolithic (H35, H36, and H38), 13% Neolithic (H58 and H71), 4% historic European gene flow (group IX, H50, H52), and 8% recent sub-Saharan African (H22 and H28). In contrast, the origins of the Iberian Y-chromosome pool may be summarized as follows: 5% recent NW African, 78% Upper Paleolithic and later local derivatives (group IX), and 10% Neolithic (H58, H71). No haplotype assumed to have originated in subSaharan Africa was found in our Iberian sample. It should be noted that H58 and H71 are not the only haplotypes present in the Middle East and that the Neolithic wave of advance could have brought other lineages to Iberia and NW Africa. However, the homogeneity of STR haplotypes within the most ancient biallelic haplotypes in each region indicates a single origin during the past, with possible minor reintroductions, with the Neolithic expansion, from the Middle East. Thus, Neolithic contributions may be slightly underestimated.
Detection of Gene Flow across the Gibraltar Strait
The detection of gene flow between both geographical regions may provide a measure of the reciprocal contribution of Y chromosomes that has occurred during the past. In particular, we have shown that Iberian chromosomes carrying H35, H36, and H38 originated in NW Africa and were brought recently to the peninsula. Their frequency in Iberia will allow us to estimate the maximum NW African male contribution to the Iberian Y-chromosome pool. Since not all NW African Y chromosomes carry those haplotypes, gene flow from NW Africa must have brought other chromosomes. Thus, to estimate the NW African contribution, the proportion of H35, H36, and H38 chromosomes in NW Africa must be taken into account. Therefore, we estimated the overall NW African contribution to the Iberian Y-chromosome pool as being 5% (the frequency of H35, H36, and H38 in Iberia) divided by 75% (the frequency of those haplotypes in NW Africa)—that is, 7%, with the highest level of contribution (14%) being found in Andalusians from southern Iberia. Conversely, since group IX chromosomes in NW Africa may have an Iberian origin, the Iberian (or European) contribution to NW Africa can be estimated, as above, as being 4%.
A small NW African genetic contribution in Iberia is also detected with mtDNA, the female counterpart of the Y chromosome. Rando et al. (1998) suggested a NW African–specific origin for mtDNA haplogroup U6, which is found at frequencies of ~10%–20% in NW Africans and is absent or nearly absent in Europeans and other Africans. The presence of this NW African mtDNA haplogroup in Iberia can be used as an indicator of NW African–female contribution. Such a contribution seems to be small, since haplogroup U6 is found at very low frequencies: it has been found in 3 of 54 Portuguese and in 2 of 96 Galicians and is absent in Andalusians and in 162 other Iberians (Bertranpetit et al. 1995; Côrte-Real et al. 1996; Pinto et al. 1996; Salas et al. 1998).

We have detected male-mediated gene flow from NW Africa to the Iberian Peninsula; gene flow in the opposite direction, as shown by the Nm and admixture estimates and by ages obtained from STR haplotypes, occurred at lower levels and is more ancient. However, date estimates integrate all the gene flow between the two regions and should be regarded as giving an average rather than as pinpointing a single event. In that respect, the more ancient age estimate for the north-to-south gene flow could have been caused by the fact that it occurred on a haplotype background, H104, that is slightly more diverse than its south-to-north counterpart, H38 (compare figs. 3c and 3b, respectively), thus carrying a more diverse set of Y chromosomes from Iberia into NW Africa.

The Islamic (Arab and Berber) occupation of the Iberian Peninsula, which began in a.d. 711 and, in the south, lasted until a.d. 1492, left a rich cultural heritage, from science and philosophy to agriculture and architecture. Islamic rule lasted longest, until 1492, in southern Iberia. Our results suggest that the demographic contribution linked to that occupation (and to movements in the opposite direction) must have been small but not at all negligible.

This study has demonstrated the unprecedented power of the use of Y-chromosome biallelic polymorphisms for the dissection of paternal lineages, which has allowed us to cut through the historic layers in the Iberian and NW African gene pools in much the same way as archaeologists excavate prehistoric layers at a site.

The oldest mini skirts in history.

Now archaeologists say the true origins of the mini go back to the very dawn of civilisation.

They have unearthed evidence that Stone Age women were wearing mini-skirts – along with short tops and bracelets – more than 7,500 years ago.

A series of stone figurines wearing the prehistoric fashions were unearthed at one of Europe’s oldest known villages – a community that nestled between rivers, mountains and forests in what is now southern Siberia.

The finding pushes back the origins of fashion and art in Europe by hundreds of years to a time when our ancestors were first getting to grips with farming.

“According to the figurines we found, young women were beautifully dressed, like today’s girls in short tops and mini skirts, and wore bracelets around their arms,” said archaeologist Julka Kuzmanovic-Cvetkovic.

The unnamed tribe lived between 5400 and 4700 BC in the 120-hectare site at what is now Plocnik. Remains at the site reveal that they knew about trade, handcrafts, art and metallurgy while a thermal well nearby might be Europe’s oldest spa.

“They pursued beauty and produced 60 different forms of wonderful pottery and figurines, not only to represent deities, but also out of pure enjoyment,” said Dr Kuzmanovic.

Women, it seems, have always paid attention to their appearance

Little is known about the life of people – known as the Vinca – who made the figurines.

The Vinca culture flourished between 5500 and 4000 BC in Bosnia, Serbia, Romania and Macedonia. It got its name from the present-day village of Vinca on the Danube River near Belgrade where eight villages have been found.

The latest discoveries suggest these early farmers had developed a sophisticated division of labour and organisation.

Houses had stoves, there were special holes for trash, and the dead were buried in a tidy necropolis. People slept on woollen mats and fur, made clothes of wool, flax and leather and kept animals.

They were especially fond of children. Artefacts include toys such as animals and rattles of clay, and small, clumsily crafted pots apparently made by children at playtime.

One of the most exciting finds for archaeologists was the discovery of a sophisticated metal workshop with a furnace and tools including a copper chisel and a two-headed hammer and axe.

“This might prove that the Copper Age started in Europe at least 500 years earlier than we thought,” Dr Kuzmanovic said.

The Copper Age marks the first stage of humans’ use of metal, with copper tools used alongside older stone implements. It is thought to have started around the 4th millennium BC in south-east Europe, and earlier in the Middle East.

The discovery of Europe’s oldest mine at the nearby Mlava river suggested at the time that Vinca could be Europe’s first metal culture, a theory now backed up by the Plocnik site.

“These latest findings show that the Vinca culture was from the very beginning a metallurgical culture,” said archaeologist Dusan Sljivar of Serbia’s National Museum. “They knew how to find minerals, to transport them and melt them into tools.”

The metal workshop in Plocnik was a room 25 square yards, with walls built out of wood coated with clay. The furnace, built on the outside of the room, featured earthen pipe-like air vents with hundreds of tiny holes in them and a prototype chimney to ensure air goes into the furnace to feed the fire and smoke comes out safely.

In Bulgaria and Cyprus, where other workshops have been found, the early metal workers blew air on to the fire using straws rather than relying on chimneys.

They probably experimented with colourful minerals that caught their eye – blue azurite, bright green malachite and red cuprite, all containing copper.

The village was destroyed at some point, probably in the first part of the fifth millennium, by fire.

The Plocnik site was first discovered in 1927. Some findings were published at the time but war, lack of funds and objections from farmers meant it was investigated only sporadically until digging started in earnest in 1996.

The copper forges have been dated to about 5,500 BC, and a copper ax head the same age has now been found in Serbia too.