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Friday, July 1, 2016

Toward a More Logically Acceptable Model of Archaic Introgression

Because the old models of human evolution during the Paleolithic Revolution have yet to be reconciled with new fossils and new genetic data, I have had to reconstruct the most probable scenario from the evidence at hand. This is only so that my Prehistoric Fiction might stand the test of time, as new evidence is found over the coming years. I have tried to make many different models fit into our current understandings of prehistory, but only a few seem to work without discounting much of the data at hand. Perhaps the following is the best and simplest of my reconstructions of hominid evolution, and, in any case, an understanding of it is necessary to explain my theories on the origin of the Hobbits, Artificial Cranial Deformation, and introgression from Archaic Hominids such as Neanderthal, Denisovan, and others.

About 3.5 million years ago, the ancestor of all members of the genus homo was born into a population or subspecies of australopithecine, a chimp-like bipedal ape known only from Africa's fossil record. Most likely, this species of ape possessed fused chromosomes, a condition which had sexually isolated the population from other species of australopithecine. In this individual, a copying error occurred to produced a duplicate of the gene SRGAP2 known as SRGAP2B, which has been implicated in brain development.

This new gene became dominant in the population, and over the next half million years was positively selected. By 2.9 million years ago, one of the descendent populations, the burgeoning species which we will call Early Homo, had become abundant enough to leave fossils for scientists to find. These include the LD 350-1 hominid (Ethiopia) and possibly Homo Naledi (South Africa). Around 2.6 million years ago, Early Homo (and perhaps a sibling but non-homo species called A. Garhi) produced the first Mode 1 pebble tools (Oldowan) in East Africa (Ethiopia).

But by 2.5 million years ago, another Mode 1 industry (Madrasian) had sprung up in India, and by 2.4 another in Southwest Asia(Oldowan), bearing testament to the presence of Early Homo outside of Africa at least 700,000 (and probably a million) years before the date generally presumed by most popular media.

Another duplication error occurred in the brain of a member of Early Homo about 2.4 million years ago, producing SRGAP2C, which is thought to have greatly increased the benefits of the first mutation(or short-circuited it). Again this new gene began its sweep over the genome of Early Homo. By 2.3 million years ago, one subspecies of Early Homo had specialized into a niche in South East Africa and left enough bones behind to be given the taxon "Homo Rudolphensis." Rudolphensis may or may not have possessed the second "brain booster," SRGAP2C, but the species is not thought to have been a direct human ancestor.

By 1.8 million years ago (and probably earlier), Early Homo had reached South East Asia and Europe. Homo Georgicus with his Mode 1 technology and pebble core tools from Java attest to this. Also by this time, new subspecies of Early Homo had arisen in Africa, becoming prolific enough even to leave fossils behind, such as Homo Gautengensis, Homo Habilis, and Homo Ergaster).

Between 1.66 and 1.63 million years ago, pebble tools and Early Homo fossils are being deposited in both China and South East Asia, namely Lantian Man and Homo Erectus Modjokertensis.

Later radiations and subspecies of this first wave of Early Homo out of Africa likely include Early Homo Antecessor (1.0 - 1.2 million years old in Spain), Meganthropus paleojavanicus (1.2 - 1.4 million years old in Indonesia), Early Homo Erectus Pekinensis (780,000 years old in China), Homo Floresiensis (<50,000 to 780,000,000 years old in Indonesia), and the "Mystery Hominid" known only from studying the human OAS1 gene and the Denisova genome, which we'll discuss later. Except for the last, all of these subspecies can be classed as Early Homo by similarities in morphology to Georgicus, Habilis, or the Australopiths that are not found in later members of homo erectus, by the use of Mode 1 or pebble-core tools (though sometimes by their advanced use), and by brain volumes of less than 900cc. Also, possibly, by the lack of certain anatomical features that are found in later and supposedly more "sapien-like" examples of Homo Erectus.

Despite the success and diversification of Early Homo and his Mode 1 tools, a new lithic industry was on the rise by 1.75 million years ago; Mode 2 (Acheulean). In recent years scientists have become reluctant to ascribe any one species to this second lithic tradition, but the aforementioned Homo Ergaster derives its name from it due to their close association.

Though it is unknown whether the sweep of the second brain gene, SRGAP2C, included rudolphensis, habilis, gautengensis, georgicus, or meganthropus, it is highly probable that Homo Ergaster possessed it. Ergaster's brain was nearly 300 cc larger than any hominid that had gone before it. Its face and morphology more closely resembles the morphology of homo sapien sapiens than any of its contemporaries. It is therefore more likely to be ancestral to us, and all homo sapiens have SRGAP2C. It might even be said that the brain gene mutation led to the innovation of the second mode of tool manufacture, because Mode 2 shows up in East Africa shortly after the first fossils of Homo Ergaster and the birth of the gene.

By 1.6 million years ago, Mode 2 had spread to South Asia. Though Homo Ergaster may have well been the species that brought the industry there, another population of Early Homo was already occupying India at this time. Later Mode 2 cultures in South Asia show local influence from advanced Mode 1 industries that had developed there before Mode 2's arrival. It is likely, then, that homo ergaster mixed with this Indian population, culturally and sexually, rather than driving it into extinction.

There are people living in India today who retain genes from a hominid that split from our lineage 3 million years ago (we'll address this later). This implies the area's late occupation by a population that split from and survived in isolation from the lineage that led to homo sapien through most of the Paleolithic, before recombination in the Late Paleolithic. Fossils in or near South Asia inevitably favor "Classic Erectus" morphology over "Homo Ergaster" for the whole of the timeline, and seem to radiate from the area at several times throughout prehistory. Unfortunately, despite ample Mode 1 and 2 Tools going back nearly as far as the record in Africa, India gives us no fossils until 600,000 years ago. Narmada Man has alternately been called an advanced Homo Erectus or an early Homo Heidelbergensis, but it is of the “classic erectus” shape like Java Man with a much larger brain volume and a suite of advanced features shared by some of the later “Advanced” or “Archaic” members of Homo.

It is now necessary to define the term "Classic Erectus," because its use has fallen somewhat out of favor and some modern researchers would like to put all of the aforementioned members of Homo under the broader classification of Homo Erectus, including most later hominids (with the exception of Homo Sapiens and Homo Neanderthalensis, perhaps). I am not in agreement with them, simply because names of genetic populations are useful to explain the hybrid nature of modern man. We can call them "breeds" for all I care, and that would make Modern Man a mutt who survived the extinction of the various breeds of “Homo.” We became a separate species from Homo Erectus only by assimilating and assembling what was left of the Homo Erectus genome after several devastations.

For the purposes of this article "Homo Erectus" will include only those members of the genus Homo whose cranial shape more resemble classic Java Man than Homo Ergaster or the earlier subspecies of Early Homo, and who possess an adult brain volume roughly equal to or greater than Homo Ergaster.

"The fossils found in Java are considered the type specimen for H. erectus...Java Man was about 5 feet 8 inches (170 cm) tall and his thighbones show that he walked erect like modern humans. The femur is thicker than that of a modern human. The skull was characterized by thick bones and a retreating forehead and no chin, as well as protruding brow ridges and a massive jaw. With 900 ccm, his cranial capacity was smaller than that of later H. erectus specimens. He had human teeth with large canines." -wikipedia

These fossils were found in Java, Indonesia and date from about 1.49 million years ago (plus or minus 130,000 years) and exhibit a more dolichocephalic (long) and sloped cranium than any Early Homo, including ergaster. I am proposing that the origin of this species is South Asia rather than South East Asia because "Classic Homo Erectus" traits seem to radiate from and are found in India until at least 75,000 years ago (Hathnora skullcap and others), and possibly until as late as 6000 years ago (Balangoda Man). They begin to show up in Java 1.49 million years ago, but are contemporary with Early Homo there, such as Meganthropus and more Modjokertensis-like fossils, which would be in line with an exodus from India.
These classic erectus traits begin appearing in ergaster specimens starting around 1.5 million years ago, but between 1.4 to 1 million years ago Africa yields no fossils at all. The fossils that show up after 1 million ybp in Africa are never called ergaster, and are always designated to Homo Erectus (until the arrival of the Bodo skull, which is sometimes considered an early heidelberg or archaic).

Classic Erectus traits show up in 600,000 year old fossils from Palestine with Galilee Man and at the same time in Europe with Ceprano Man, Samu Man, and Arago 21.

While the 1.6 million year old Lantian Man fossils and the 780,000 year old Early Peking Man fossils in China favor Early Homo and have a smaller brain volume (780 for Lantian, latter unknown), the 500,000 Late Peking Man fossils are larger brained (1000+ average) and have the Classic Erectus look in spades. In both Europe and in China, Classic Erectus shows up between 780,000 and 600,000 years ago in areas that are already occupied by Early Homo subspecies- Early H. Antecessor and Early H. Pekinensis (respectively).

It could be that Homo Ergaster and Homo Erectus were the same species, and it is tempting to even say that there is no difference between them, since only the earliest ergaster fossils lack Classic Erectus traits and predate Java Man himself, which is the earliest of the type known. However, certain traits found in Ergaster but not in classic erectus continue in Homo Antecessor, Homo Heidelbergensis (African and European), and maybe even in Modern Man, suggesting at least some continuation in the West for 1.8 million years. The aforementioned 3 million year old gene also suggests separate African and Indian populations that were at least somewhat sexually isolated from each other. But the shared Mode 2 technology, increased brain size, and similarity of certain late Homo traits between the two lineages might suggest that both populations enjoyed the benefits of SRGAP2C and at least occasionally interbred.

The Saharan Pump Theory does well to explain the dynamic, and can be applied to the Thar Desert as well. Both Africa and India alternated between severe dry periods and lush, fertile periods during the Ice Ages. Some researchers believe they were all but abandoned during the dry periods. The period between 1.4 million and 1 million years ago was one of these. Though no ergaster or erectus fossils show up in India or Africa during this time, and few tools, the closely Ergaster-related Homo Antecessor and the Classic Erectus "Java Man" both appear first in areas adjacent to Africa and India at the beginning of the dry spell. 
It is likely that members of these the African and Indian populations were also driven into the area between the two desert pumps- South West Asia and the Middle East. The repeated hybridization of the two populations during dry periods eventually led to the rise of Archaic Homo Sapien.
It should be said that though Homo Ergaster favors Modern Man more than does Homo Erectus in the areas of body ratios and facial features, several studies have shown that we owe more of our dental structure to Classic Asian Erectus. Though African Archaics and Modern Humans have a head shape more like that of Ergaster, most European Heidelberg and all Neanderthal crania are elongated like those of Classic Erectus, and all of these show similarities not seen in any of the earlier hominids.

Around 1 million years ago, a third brain gene duplication occurred, SRGAP2D, and it is likely that it happened in a member of this hybrid population, somewhere in Southwest or Central Asia. SRGAP2, of course, is probably not the only gene that led to the eventual world-dominance of Archaic Homo Sapiens. Several other genes of this age have been identified which made total or near-total sweeps over our genome. A hybrid population would have had access to many new and beneficial genes developed during periods of separation. However, SRGAP2 is a great genetic marker to use in attempting to unravel prehistoric interchange.

It is interesting that 500,000 years after SRGAP2B, the first member of the genus Homo appears along with Mode 1 tools. 500,000 years after SRGAP2C, the first Advanced Homo (Ergaster) appears along with Mode 2 tools. 500,000 years after SRGAP2D, the first Archaic Homo specimens appear along with Mode 3 tools. Perhaps it takes 500,000 years for a brain gene to make enough of a sweep over a significant population to create a new stage in species-wide brain development?

Returning to Early Homo, as opposed to Ergaster and Erectus, it is useful to look at each case individually.
In China, smaller brains and Mode 1 tools predominate until at least 500,000 years ago (Early Peking Man and Lantian), whereupon brain capacity increases markedly and the tools become more advanced (Late Peking Man). By 200,000 years ago some fossils there are showing definite Archaic (sapien-like) features, such as Dali and Jinnuishan, as well as yielding evidence of Upper Paleolithic technology.

Europe shows a similar pattern- Homo Antecessor of 1.2 million years ago used Mode 1 tools, had many traits reminiscent of Georgicus, Earliest Ergaster, and even Australopithecines, and likely had a small brain case. Antecessor of 500,000 years ago exhibits a Late Homo sized brain and anatomical features in common with Neanderthal and Heidelberg. Mode 2 tech also shows up in Europe around 500,000 years ago, along with the very “Classic Erectus-like” Ceprano Man. By 350,000 years ago, true Archaics are showing up in the form of Heidelbergensis and proto-Neanderthal, together with Mode 3 tool technology.

Besides being members of Early Homo, both early and late Peking Man and Antecessor share several other traits in common, including a type of gracile jaw that is unique to these two species alone.

The simplest explanation for these parallel developments is that the regional variants of Early Homo (Early Antecessor and Early Peking Man) were assimilated first by variants of Late Homo (Erectus) beginning about 800,000 years ago, and then later assimilated by variants of our SRGAP2D "hybrid," Archaic Homo, beginning sometime after 600,000 years ago.

Late Antecessor is the result of the mating between Early Antecessor and insurgent Homo Erectus Mauritanicus, who brought Mode 2 tech and SRGAP2C. European Heidelberg is the result of mating between Late Antecessor and the insurgent Ceprano Man, who brought SRGAP2D, and the cultural exchange led to Mode 3 (Mousterian).

Late Peking Man is a result of the mating between Lantian Man and an insurgent Classic Erectus, bringing Mode 2 -like tech and SRGAP2C. Dali is the result of mating between Late Peking Man and the insurgent Jinnuishan, bringing Mode 3 and SRGAP2D.

African Erectus is the result of mating between Ergaster and insurgent Classic Erectus, inspiring the creation of Mode 2 and sharing SRGAP2C. Rhodesian Man (and ultimately Homo sapien) is the result of the mating between Late Ergaster/Erectus and the insurgent Bodo Man Archaic, bringing SRGAP2D and leading to Mode 3.

In Java, the situation is a little different. Modjokertensis, Meganthropus and the earliest Sangiran specimens are undoubtedly Early Homo, and Classic Erectus shows up between 1.49 million and 700,000 years ago. However, later specimens from Ngandong and the Solo River (as well as Homo Floresiensis and his 780,000 year old ancestor) have traits that tie them both to Early Homo (and even possibly to the Australopithecines in the case of the Hobbits).

Java was alternatively an island and part of the mainland many times during the Paleolithic, and between 800,000-900,000 years ago was devasted by a meteor shower. This meteor storm may have affected the entire planet, as geneticists have detected a bottleneck in several species including our own from this time, and there is a marked difference between hominids in strata before and after this date all over the world. This is also the time of a pole shift, which helps scientists to date fossils before or after by their magnetism.

Classic Erectus may have found himself trapped in or nearby Java to be assimilated by Early Homo in a strange reversal of fortune, with the resulting hybrids driving their “Hobbit ancestor” siblings across the Wallace Line to evolve in isolation.

It is important to note that a small brain does not necessarily mean lower intelligence. Soloensis and especially the Hobbit had smaller brains than contemporary hominids, but their technologies have been considered Upper Paleolithic despite showing no influence from Mode 2 or Mode 3 traditions. Their cultures, though unique to them, are more advanced than that of some cultures of Modern Human. It is likely that Microcephalin D was assimilated from one of these two species, and this brain gene has proven so beneficial to our own species as to make a 70% sweep of our entire genome in less than 40,000 years.
As we move on to the arrival of Neanderthals, Homo Sapiens, and Anatomically Modern Humans, let us put the above model of hominid evolution with the facts of Archaic Introgression.

"In 2012, researchers studied high-coverage whole-genome sequences of fifteen Sub-Saharan hunter-gatherer males from three groups—five Pygmies (three Baka, a Bedzan, and a Bakola) from Cameroon, five Hadza from Tanzania, and five Sandawe from Tanzania—finding signs that the ancestors of the hunter-gatherers interbred with one or more archaic human populations,[39] probably over 40,000 years ago.[41] They also found that the median time of the most recent common ancestor of the fifteen test subjects with the putative introgressive haplotypes was 1.2–1.3 mya." - Wikipedia, Archaic human admixture with modern humans.

In other words, a group of the aforementioned Advanced Erectus living somewhere in or between East Africa and Asia migrated into (Central) Africa around 1.3 million years ago, assimilated Ergaster, and evolved there in isolation until Homo Sapiens assimilated them in turn around 40,000 years ago. This group most likely possessed SRGAP2B and SRGAP2C, but not SRGAP2D, since the last is thought to have only evolved after their split from our common ancestor.

According to the nuclear DNA analysis, Denisovans split off from our ancestral line over a million years ago. 700,000 years ago, this group also split in two, hence the Denisovan-like nuclear DNA in the Heidelberg/Antecessor Atapuerca hominid. In other words, one group split off from the aforementioned Advanced Erectus 1 million + years ago and headed north into Central Asia (Basically the same time as the African split). Around 700,000 years ago, this group split into an Eastern group that assimilated Lantian Man to create Peking Man and a Western group that assimilated Antecessor to become European Heidelberg.

"...haplogroup D of microcephalin, a critical regulatory gene for brain volume, originated from an archaic human population.[18] The results show that haplogroup D introgressed 37,000 years ago (based on the coalescence age of derived D alleles) into modern humans from an archaic human population that separated 1.1 million years ago." -wikipedia, Archaic human admixture with modern humans."

In other words, a group of Advanced Erectus moved into South East Asia over 1.1 million years ago (Basically the same time as the African split). It assimilated Modjokerto and Meganthropus to later become Homo Erectus Soloensis. There it developed Microcephalin D, which introgressed into Homo Sapiens 37,000 years ago when Anatomically Modern Humans first arrived.

According to the mtDNA genome analysis, "The estimated average time of divergence between Denisovan and Neanderthal sequences is 640,000 years ago, and the time between both of these and the sequences of modern Africans is 804,000 years ago." Wikipedia, Denisovan

In other words, one group split off from Archaic Homo (Advanced Erectus with the addition of SRGAP2D) around 800,000 years ago and headed north into Central Asia. Around 640,000 years ago, this population split into two groups. One went West and assimilated European Heidelberg, eventually becoming Neanderthal, and the other went East and assimilated Peking Man, eventually becoming Jinnuishan and Dali Man.

True Archaic Homo

Sometime after 1 million years ago, the ancestor of Homo Sapiens and Homo Neanderthalensis was born in the area between or including East Africa and India. The resulting species (Archaic Homo) assimilated the various subspecies of Homo Erectus in much the same way that Erectus had assimilated Early Homo, throughout the Old World. Hence the Neanderthal-like component in the Denisovan genome and the presence of so-called "Neanderthal introgression" in all Modern Humans outside of sub-Saharan Africa. One branch of this Neanderthal/ Sapien ancestor assimilated the Homo Erectus in Africa, which led to the African Archaic Hominids (Idaltu, etc.) that were contemporary with Neanderthal Man. From the interbreeding of several populations of African Archaics in Africa, Anatomically Modern Humans evolved around 300,000 years ago. This species was largely contained in the area between and/or including Africa and India by Neanderthaloids until sometime after 70,000 years ago. After the eruption of Mount Toba had thinned populations of Neanderthaloids and Erectines considerably, Anatomically Modern Human was able to expand.
Around this time, an Anatomically Modern Human population with significant Neanderthal genetic contribution arose, probably arising in the Levant, Middle East, or North east Africa). He began assimilating all Erectines, Neanderthaloids, and other Anatomically Modern Humans. Therefore, all Modern Humans, with the exclusion of some isolated groups of Sub-Saharan African, have pieces of the Neanderthal genome. However, Sub-Saharan Africans have pieces of the African Homo Erectus Genome, and some West Africans have introgression from a 400,000 year old "heidelberg" y haplogroup. This likely means that the African Archaics were far better suited than the Neanderthal hybrids for the Sub-Saharan environment and took the Anatomically Modern Humans in as they fled from the invasion. But we will get to the dynamic between Homo Sapiens and Neanderethals in a later article; it is only useful to give a brief summary for the purposes of our current subject.

Suffice it to say that no pure Homo Sapiens exist any more, though Y Adam and Mito Eve show that they probably once did. They were assimilated by the hybrids, and  Modern Man is the hybrid. 

There are 200,000 years worth of Anatomically Modern Human Y DNA lineages between Y haplogroup A00 and A1 that we hybrids wiped out or drove to extinction with the help of introgressed genes. There are 150,000 years worth of Anatomically Modern Human mtDNA  between LM3 (Mungo Man) and mitoEve that we hybrids wiped out or drove to extinction with the help of introgressed genes.

Papuans have high Neanderthal admixture as well as Denisovan and Early Homo genes, some Indians have both Early Homo and Neanderthal genes, and nearly all modern populations have introgression from the Microcephalin D hominid and LM3, a group of Anatomically Modern Humans that branched off at least 150,000 years before Mitochondrial Eve (as evidenced by the DNA of Mungo Man and the LM3 insertion found in Modern Humans).
In short, all Modern Humans are Neanderthal Hybrids except a few isolated groups in Sub-Saharan Africa, who are hybrids with a comparable Archaic Hominid that was likely better suited to Sub-Sahara. 70% of modern Neanderthal hybrids are also Erectus hybrids, as evidenced by Microcephalin D, and some populations in South East Asia, Oceana, and India are also hybrids with late surviving Early Homo, as evidenced by the introgression of OAS1.

Fernando L. Méndez et al., Global genetic variation at OAS1 provides evidence of archaic admixture in Melanesian populations. Molecular Biology and Evolution 2012;

"Recent analysis of DNA extracted from two Eurasian forms of archaic human show that more genetic variants are shared with humans currently living in Eurasia than with anatomically modern humans in sub-Saharan Africa. While these genome-wide average measures of genetic similarity are consistent with the hypothesis of archaic admixture in Eurasia, analyses of individual loci exhibiting the signal of archaic introgression are needed to test alternative hypotheses and investigate the admixture process. Here, we provide a detailed sequence analysis of the innate immune gene, OAS1, a locus with a divergent Melanesian haplotype that is very similar to the Denisova sequence from the Altai region of Siberia. We re-sequenced a 7 kb region encompassing the OAS1 gene in 88 individuals from 6 Old World populations (San, Biaka, Mandenka, French Basque, Han Chinese, and Papua New Guineans) and discovered previously unknown and ancient genetic variation. The 5′ region of this gene has unusual patterns of diversity, including 1) higher levels of nucleotide diversity in Papuans than in sub-Saharan Africans, 2) very deep ancestry with an estimated time to the most recent common ancestor of >3 million years, and 3) a basal branching pattern with Papuan individuals on either side of the rooted network. A global geographic survey of >1500 individuals showed that the divergent Papuan haplotype is nearly restricted to populations from eastern Indonesia and Melanesia. Polymorphic sites within this haplotype are shared with the draft Denisova genome over a span of ∼90 kb and are associated with an extended block of linkage disequilibrium, supporting the hypothesis that this haplotype introgressed from an archaic source that likely lived in Eurasia."

This was first brought to my attention by the "For What They Were, We Are" blog, who's author goes on to comment on other materials in the paper:

"I find particularly notable that the haplotype has been found at very low frequencies in South Asia and nowhere else West of Wallace Line. It can be backflow but may also be indicator about the possible location of the admixture event."

To me the answer to the riddle is right before us; Chinese "Denisova" assimilated a subspecies of Early Homo in China. The genes don't show up in modern Chinese people because Neanderthal and extinct lineages of Homo Sapien assimilated Denisovans in China before Modern Man ever got there. South East Asian "Denisovan" assimilated a separate but related subspecies of Early Homo in Sundaland, and was in turn assimilated by Modern Papuans. Extinct lineages of Homo sapiens or Western Neanderthals assimilated a separate but related subspecies of Early Homo in South Asia, and were in turn assimilated by certain groups of Modern Man in India.

I'm not saying that this is exactly how it all happened, but writing prehistoric fiction that might stand the test of time requires a model that reconciles the genetic data with the fossil and lithic record. Modern scientists seem afraid to propose a model at all, and the brief sketches they provide in encyclopedias and text books don't quite make sense unless you leave out a few facts or dismiss them as dating errors or anomalies. Additionally, there is a similar but alternative model I've considered that might work as well. It involves a different dynamic between the two Advanced Erectus subspecies that lived in or between East Africa and India during the Paleolithic. It may not have been just two different populations, but two different niches, that created the difference between the Ergaster and Erectus type and facilitated the alternating genetic exchange and isolation between them. I will discuss this in a future article I plan to entitle, "The Tall and Short of Human Evolution: Dispelling the Repeating Hominid Dwarfism Myth."

Until then, consider the possibilities and let me know how the theory can be improved in the comments, it's not a static model!

Joe Lyon Layden is a prehistoric fiction author and primitive musician. To receive a free copy of this entire novella "The Man from Parkho Khatune Bears Favor," as well as three free songs and monthly updates, freebies, and discounts on Joe's ongoing work, please sign up for the newsletter below.

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