It depends on where you draw the line between HSS and surving lineages of HSS, but the short answer is Afro-Asians, probably from Mesopotamia or the Levant.
There are several MtDNA studies which show that the first members of the species homo sapiens sapient to inhabit North Africa and to produce lineages which carry on in the region (and/or in the adjacent Canary Islands) until the present entered some 30-65,000 years ago and contained within their population the original portion of the haplogroup M which would later lead to the M1 subclades.
M is one of only two haplogroups which made it out of Africa in the original Homo Sapien Sapient expansion some 65-75,000 years ago and is generally associated with Asian peoples. The other of the two haplogroups is N which is generally associated with Eurasian peoples.
This expansion coincides with the region's Aterian culture, which began over 85,000 ybp and ended some 22,000 to 10,000 ybp, but which may have continued in some remote areas of northwest Africa and it's adjacent islands until historic times. Some of the earliest sites of the Aterian can be attributed to homo erectus, who inhabited the coasts from 200,000 ybp or earlier. There are differences in the eastern and western branches of the industry, and up until about 45,000 years ago some of the Aterian lithics can be attributed to Neanderthal man, whose fossils have been found in the far eastern portion of the region. Man is thought to have entered North Africa 85,000 ybp or later, and haplogroup M came with it at that time or up to 55,000 years later. however, only those peoples carrying the strand of haplogroup M, which would later mutate into M1 while in still in Africa, have continued living in the area until today. Ancestral "African M" also spread into (or was originally from) east Africa at about the same time as North Africa.
Haplogroup M was accompanied into North Africa, either at the original time of habitation or shortly thereafter, by peoples carrying haplogroup U6, a descendent of the Eurasian N lineage. We can't at present discern what male, or Y chromosomes, went into Africa with "hereditary" M1 and U6.
Common suspects for the Y culprit include E1b1b, which is still dominant among Ethiopians, Somalis, Eritreans and North African Berbers and Arabs, and E1a , which has been detected among samples obtained from Moroccan Berbers, Sahrawis, Burkina Faso, northern Cameroon, Senegal, Sudan, Egypt, and Calabria (including both Italian and Albanian inhabitants of the region), although E1a has only a small presence in North Africa.
Haplogroup E, the ancestor of both E1b1b and E1a, is thought to have originated in East Africa, but there are many who argue that it's dispersal points to a back migration into Africa from Asia. It's sister clade, D, is only found in Asia, though it's mother clade ED belongs to an "African" lineage.
E1a is found almost exclusively in Africa, while most non-african peoples who belong to haplogroup E belong to the E1b1b subclade.
The question partly comes down to whether the Guanches of the Canary Islands, generally agreed as the closests descendents of the Paleolithic North Africans, have or have ever had the y-Chromosome E1b1b in their population, and if so, whether they got it before or after the hostile takeover of their culture by subsequent waves of later peoples.
While E1b1b has been reported in the Guanches, the results have been questioned, and it may be attributed to Pheonician, Berber, Arab, and even Spanish influence during historical times. E1a has not been reported from indigenous Canarians, but that can likewise be attributed to hostile takeover and erradication of all original male gene lineages.
There are studies which show that at some time in the distant past the early original "M1 people" were subject to a violent takeover by peoples containing haplogroup R1*, which is a very rare subclade of R1.
With R1b1 and R1b1*(which might actually be R1*) this haplogroup is one of the only two or three subclades of R1 that seem to never have made it into Europe, at least until the common era.
R1* has two sister clades, R1a and R1b, which are the so-called "Cro-Magnon" haplogroups, and these or their common ancestor accompanied the first homo sapien sapient men into Europe somewhere between 45,000 to 25,000 years ago. The two groups make up 70% of the European male lineages, with specific countries, areas, and peoples often reaching over 90%.
At present, we can't tell if the takeover performed by R1* in this early North African population came before or after U6 and M1 spread into North Africa, but we do know that R1* was, in turn, nearly erraticated in the population at a later date....either by those very E1s who still exist in the descendent populations of Berber, Arab, and possibly Guanches today or by members of the later Capsian and Arab cultures that followed, and who carried their signature Haplogroup J along with their own E1 haplogroups.
We also don't know if R1* was already with U6 when he killed off M1's men and took the females for his own, or if, instead, M1 and U6 were already together in the original population and R1* brought no females when he came to call. But R1* is a good guess for the earliest match for U6, being that sister clades U5 and U8 are thought to have been the first into Europe, just like the brother clades of R1*.
In any case, R1* took over the clans that contained U6 and M1 either before or after they spread into North Africa 30-35,000 years ago. If it was after, it can only have been in the accompaniment of the Ibero-Maurusian culture some 18-22,000 ybp, because there is no real change in the Aterian culture from 35,000 ybp until that time.
The Ibero-Maurusians, also known as Oranians and Maghreb or Afoulu man, were a seemingly violent peoples, who knocked out two to four teeth of every male and female in the tribe as a rite of adolescence around the age of 16. They are described as "hyper-robust" and often equated with "Cro-Magnon" man in the old texts (when the term was still in wider use). The main differences between the "Afoulu" or "Oranian" group and the so called "Cro-Magnon" group were said to be that the Ibero-Maurusians were slightly shorter (but still nearly 6 feet tall, being that the two skelotons found in Dordogne at the time had suggested an average height of 6'5 for Cro-Magnon) and had a slightly more primitive skull shape, with a nasal cavity more similar to non-Cro-Magnon specimens from the Paleolithic found in Africa and eastern Europe.
But the Ibero-Maurusian clans don't seem to have been influencial enough on North Africa to have slaughtered every man in the entire region, for at that time the Aterian spread over all of North Africa and even into central and east Africa as well . In fact the Ibero-Maurusians don't seem to have traversed North Africa's entire width, barely extending the culture from the Atlas Mountains and Morrocco to Tunisia. the culture was influenced and eventually succeeded by the Capsian culture, which entered the region from the east around 12,000 ybp.
Another problem with R1* being equated with the Ibero-Maurusians is that the clade is also found all the way over in Turkey and India associated with a very ancient date, and it's "twin" sister clade R1b1* is found among the Bantu and the Cameroon. It is thought to have spread into Africa from the east shortly after it's seperation from all the other R1 clades that went into Europe, some 25,000 to 45,000 years ago. This date and locale squarely places R1*, U6, and M1 together with the Aterian culture somewhere around 35,000 ybp or more. This means that R1* must have performed it's hostile takeover of the "M1" clan before or immediately after it's spread into North Africa, and a very long time before the Ibero-Maurusian.
The best assumption is that from at least 30,000 ybp until at least 22,000 ybp, all of North Africa was near completely dominated by the Aterian culture along with haplogroups M1,U6, and R1*.
It is possible that the Ibero-Maurusians contained most of the E1b1b now found in Berbers, Arabs, and possibly Guanches, entering North Africa from the east and/or south of North Africa 8,000 years before the Capsian. But it is also possible that the Capsian and Arab cultures brought these lineages later, along with haplogroup J.
The Cro-Magnon robustness of the Ibero-Maurusians, as well as supposed lithic similarities, has been attributed by some scientists to an origin in the Iberian Penninsula, from which the culture gets it's name, and from where Cro-Magnon could have conceivably arrived at such a remote date. If this is so, then the R1b they undoubtedly carried has been wrongly attributed (in Guanches and Berbers) to colonization in historic times by Europeans, or else the Ibero-Maurusian R1b lineage was wiped out by the invading Capsians and the later Berbers themselves.
We know from a recent study on Guanches that the Basques in Iberia obtained at least one matriarchal lineage from the Aterians (or Aterian/Ibero-Maurusians) sometime around 20,000 ybp. Since the Canary Islands are thought to have been uninhabited at the time, they can only have gotten the haplogroup from North Africa. This implies one of four possibilities; that there was either a human trade or matrimonial connection between the Ibero-Maurusians and Basques at the time, that the Ibero-Maurusians represent an extinct African branch of the ancestral Basque populations which maintained trade and communication across the Straight of Gibralter with it's mother population, that the Basques raided North Africa for women independent of the Ibero-Maurusians at roughly the same time as the latter's appearance in the region, or that Aterian/Ibero-Maurusian tribes entered Iberia and were there absorbed by the Basque population alone.
As there is no evidence of a distinctly Ibero-Maurusian or Aterian presence in Iberia itself at this time, the invasion from North Africa into Iberia seems unlikely.
The fact that the "robustness"comparisons of Cro-Magnon to Ibero-Maurusians were made before modern dating techniques and assumed that the North African samples were of the same age as the early "Cro-Magnon," which was comparably robust but is actually over 10,000 years older, is problematic at best for the Iberian "Cro-Magnon" origin model; by 20,000 years ago most Europeans had scaled down their hyper-robustness, even the ones that descended from the original Rb1.
If the Ubero-Maurusians were part of an ancestral Iberian population, then that Population could only have descended from the anomolous cro-magnoid "hybrids" found in the caves of northern Portugal and dated to 24,500 ybp, who were the only known Iberians robust enough at the time to fit the Ibero-Maurusian bill.
But the date of 20,000 for the Basque/Guanches shared lineage marker coinciding with the beginning of the Ibero-Maurusian in North Africa makes an independent voyage to the region by Basques coincidental, if possible.
Of course, not all of these possibilities explain who the Ibero-Maurusians actually were. A hyper-robust people arriving mysteriously out of some other part of Africa is not impossible, and could explain the E1b1b and E1a in Berbers quite nicely. And since other cultures in Africa practice tooth-knocking it would provide a cultural context.
If the neanderthal hybrids became the Ibero-Maurusians, then their culture changed between 24,500 and 21,500 while crossing to Africa, which is not surprising when two cultures collide. Their robustness and primitive features can be attributed to the so-called Portuguese "hybrids" and full blooded neanderthals around the Rock of Gibralter which were evident in the area only a few thousand years earlier. It is not surprising that there is a gap between the Neanderthals and hybrids of 24,500 years ago and the Ibero-Maurusians of 21,500 years ago if they are indeed a continuation of the same race, as the shorelines between them have shrunk and only a small portion of artifacts and fossils deposited at such a remote date survive. Furthermore, the absense of the Gravettian lithic and cave painting in southern Iberia during this period lends strength to the idea of neanderthal/hybrid occupation throughout. And 3000 years is plenty of time for the Ibero-Maurusians to have developed the practice of "tooth-knocking," which is otherwise unknown in Europe.
Turning back to haplogroup U6, we can see that it's subcalde u6a is the most common, whereas U6b is only (or mostly) found in Basques(where it has the most diversity) and Guanches, whereas subclade U6c is only found in Berbers and Guanches. A variation of U6b called U6b1 is found in guanches, but not in Basques. This shows that U6b and U6c had to have been seperate populations when the Basques and Berbers aquired them, and sometime afterwards rejoined.
Despite the coincidence in dates, it seems most probable that the Ibero-Maurusian lineage was U6c, and the Aterian U6b, because the Ibero-Maurusians would have been more adjacent to the invading Capsians and Berbers (who obtained only the U6c lineage) whereas the last bastion of Aterian culture was by that time pushed south of the Atlas mountains, or elsewise already heading to the Canaries (although the latter is unlikely since first signs of habitation are dated to no more than 5000 ybp).
If this is the case, then either the Ibero-Maurusians delivered only females to the Basques from Aterian clans that they did not yet fully control, or the Basques sailed around the lands held by Ibero-Maurusians when they came to collect their ladies. This is not to absolutely rule out the possibility that they brought some men back too, but the small amount of E1b1b in Basques is generally agreed to be from the much later Moorish invasions of the Iberian Penninsula, and R1* and/or R1b1* is unknown. R1b makes up around 90% of the Basques lineages, and the remaining 10% is diverse.
The Aterian Culture
From Wikipedia:
The Aterian industry is a name given by archaeologists to a type of stone tool manufacturing dating to the Middle Stone Age (or Middle Palaeolithic) in the region around the Atlas Mountains and the northern Sahara.
The industry was probably created by modern humans (Homo sapiens), albeit of an early type, as shown by the few skeletal remains known so far from sites on the Moroccan Atlantic coast.
Bifacially-worked leaf shaped and tanged projectile points are a common artefact type and so are racloirs and Levallois flakes. Items of personal adornment (pierced and ochred Nassarius shell beads) are known from at least one Aterian site, with an age of 82,000 years.
The Aterian culture gets a bad rap because it is seen as being comtemporaneous with the Aurignacion Culture, which didn't begin until about 45,000 ybp and only happened in Europe proper. The Homo Sapiens in the Levant of 82,000 years ago had not yet gotten an upper hand on the neanderthals of the region, who had experienced both overlapping campsites and cultures there since 95,000 ybp, and who would continue doing so for the next 7,000 years or so. The Aterian tradition was the first to have used Bows and Arrows, pushing that date from a mere 15,000 ybp to 75,000 ybp or more. The neighboring Levant was slow to take on Aurignacion culture when it form in Europe 45,000 years ago, and always received the innovations from Europe last over the next 25,000 years. But the Aterian culture did not participate in the Aurignacion or succeeding cultures at all, and instead continued with the Mousterian traditions. The use of the Levalloise technique sometimes seperates Upper Paleolithic from Middle Paleolithic for researches, so North Africa is often not given an Upper paleolithic date until 15,000 ybp. This is unfortunate because here the Mousterian techniques are taken to their final fruition, resulting in thousands of beautiful arrowheads, spearheads, and other weapons and tools. The Aterians were the first to use desert glass, and have early cave paintings that rival similar dates in Europe. They are the guys on the Discovery Channel storing water in austritch eggs for later trips across the 800 miles of Sahara desert to and from Tunisia, though I doubt they looked anything like they were portrayed. One of the earliest, if not THE earliest, mines is attributed to the Aterian Culture, from where chert was transported to the Tunisian coasts 20,000 years ago. Through much of the Aterian era, the Sahara was greatly diminished, and North Africa was covered with forest and savannah along the coasts, with abundant game to boot, and even the first true fishhooks are claimed for the culture.
But. many archies don't assign Upper Paleolithic staus to North Africa until the Capsian period starting 10-12,000 ybp because of the blade techniques, which mistakenly implies that both the Aterian and Ibero-Maurusian were both "Old Stone Age" cultures.
http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6WJS-45F4X3...
http://mathildasanthropologyblog.wordpress.com/2008/07/16/berber-and-...
http://www.springerlink.com/content/5677p31651255155/fulltext.pdf
http://mathildasanthropologyblog.wordpress.com/2008/05/02/the-asian-o...
http://freepages.genealogy.rootsweb.ancestry.com/~donegalstrongs/u6b....
http://carnby.altervista.org/troe/02-08.htm
http://www.oldstoneage.com/smugglers/aterian.htm
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