D-statistic revised: Mutation rates and maybe we did not admix with Neanderthals or other archaics...

A preprint published on Aug. 25, 2026 in bioRxiv (Unexpected D-tour Ahead: Why the D-Statistic, applied to Humans, Measures Mutation Rate Variation and Not Neanderthal Introgression by William Amos and Eran Elhaik, bioRxiv 2024.12.31.630954; doi: https://doi.org/10.1101/2024.12.31.630954) suggests that the tool used to evaluate admixtures, introgression and sharing of alleles between sister species may be flawed.

The abstract states that D statistic "relies on two untested assumptions: mutation rate constancy between groups and a negligible contribution from recurrent mutations" in the paper, the authors argue that both assumptions are invalid!

They also report that "heterozygosity modulates mutation rate" and that it is this, the difference in heterozygosity that is the critical factor.

So, first of all, we will try to explain the D-statistic.

D-statistic, what is it?

The D-statistic or ABBA-BABA test was created by mathematician Nicholas Patterson (b. 1947) to evaluate if there had been gene flow between Neanderthals and humans (Patterson, 2012). It has been used since then to study admixtures and introgressions in different species.

This test looks into four populations, P1 and P2 that are closely related, P3 also on the same tree, but on a different branch, and a distant outgroup, O. The computer program, because that is what it is, compares the alleles of each population, which can come in two variants: A, the "ancestral" ones, found in the outgroup, and B, the "derived" ones (mutated, or more recent variants). There can be two patterns. One is the "ABBA" pattern, where P1 and the outgroup share the A (ancestral) variant while P2 and P3 have the mutated or derived B variant. The other is the "BABA" pattern, where P1 and P3 have the derived "B" variant while P2 and the Outgroup share the ancestral "A" allele. See the image below as a reference.

The outgroup, helps discriminate between ancestral (A) and derived (B) alleles.

ABBA and BABA patterns should appear with equal probability if there is no interbreeding, which results (see the formula below) in a value of zero for the D-statistic. (the numerator of the equation would be zero).

The reason for this is something called Incomplete lineage sorting or "ILS", which assumes that the ancestral alleles are distributed in a random manner among the descent of a species as it evolves and branches into new ones, so it is possible that some descendants inherit some alleles but not others. Interbreeding could add alleles to a branch that hadn't inherited them.

Therefore, if there has been gene-flow (introgressions), there will be a higher number of ABBA patterns than BABA ones, signfiying a positive value for D, and therefore, hybridization, with gene flow between P2 and P3.

In the equation, C stands for the counts of ABBA or BABA being observed (value = 1) or not (value = 0) at a given site in the genome (i), the symbol Σ means "sum", so you add up all the counts and perform the subtraction in the numerator, and the addition in the denominator.

However, there are some limitations. For instance, there may be an unknown extinct, and therefore unsampled species that introgressed into P3, which has not been included in the tree. Or "population stucture" could lead to discordant readings of the ABBA-BABA sequences. By "structure" it should be understood that a population is not homogeneous, it is composed of different subpopulations with significant differences between them regarding their alleles.

This example may help clarify the subject. The method works by anaylzing the strands of DNA. They are composed of bases Thymine (T), Cytosine (C), Adenine (A), and Guanine (G) in a given sequence. The DNA of different groups are sampled and aligned. At specific sites they are compared.

Then the similarities and differences at each site are calculated. P1 is Sub-Saharan African San, P2 is an Englishman, P3 is a Neanderthal, and O, the outgroup, is a chimpanzee. So, we will look at different sites and identify which base is found in the DNA, in the example they are either Guanine (G) or Adenine (A).

The formula is applied, and D is calculated. In this case it seems that Neanderthals introgressed into non-Africans.

But, apparently this method is flawed.

The incorrect assumptions

Amos and Elhaik argue that the two basic assumptions of the D statistic are wrong. Let's follow their logic:

• Mutation Rate is not constant. In genetic studies, it is taken for granted that mutation rates are invariable. They don't change. However, research has shown that this is not the case. Genetic sequences from family lineages have shown that mutation rates vary. Furthermore, the authors argue that mutation rates vary between primate species, human populations, and along chromosomes.
• Recurrent Mutations are common. Genetics assume that mutations in the same site are extremely rare. But this paper propses that they are far more common than we believe. A given single nucleotide polymorphism, or SNP, may flip from one to another base, with "at least two mutations since the most recent common ancestral base."

This is important because any change in alleles at a given site are assumed to have come from an introgression. But, say that the original, ancient variant is Adenine (A), found in chimps, and it mutated only once at a given site into Guanine (G), which is found in the Neanderthals. When we study the site in a human, who carries an Adenine (A), we would assume that there has been no introgression, but perhaps there was an introgression (G) that later mutated again, at the same site, to (A), this second mutation is what we see, but the first one was erased and therefore not counted.

Varying mutation rates have a similar effect, with one group accumulating mutations and the other one remaining stable, this violates the assumption that ABBA-BABA patterns appear at random, with the same probability across populations and lineages.

Mutation rates are different between Africans and non-Africans

The authors acknowledge that there are different mutation rates between modern Africans and people living elsewhere. They attribute it to enzyms that regulate the replication of DNA or, that heterozygosity affects it (see my post "Mutation rate is Faster in Africa"). So, the Out of Africa event, with its bottleneck reduced the heterozygosity of non-Africans and caused a drop in their mutation rate:

"Moreover, we find that differences in heterozygosity strongly predict relative mutation rate, with higher heterozygosity in Africa predicting a higher mutation rate in Africans"

The authors also point out that introgression is almost always found in genes that are subjected to "strong selection", and they reason that "Strong selection appears to impact local mutation rates... and we suggest a simple mechanism whereby selection impacts heterozygosity which in turn drives changes in mutation rate."

I must admit that their case is strong. Mutation rates are variable, and tools like the D statistic may be strongly influenced by facts like recurrent mutations and variable rates. We should not forget that these "statistics" are just models, tools, they are not facts, they are instruments to help us understand how the real world works. All models are fallible and can be improved.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

PRDM9 and our Ancestry

In my previous post about John Hawks take on the ancestral groups that merged (80:20 ratio theory) to form modern humans, I mentioned that he cited several research papers in his analysis. Today I will look into one of them, by Loya, H., Gupta Hinch, A., Palamara, P. F., Speidel, L., & Myers, S. R. (2026). Genome-wide genealogies reveal deep admixtures forming modern humans. bioRxiv. https://doi.org/10.64898/2026.04.17.719197

Loya et al., use a new algorithm called Ghostbuster to study ancient admixtures, and after validating it with recent admixture events, use it to evaluate PRDM9 alleles found in modern and ancient humans.

The PRDM9 gene is linked to genetic recombination. It codes for a protein called PR domain zinc finger protein 9, that plays an important role during meiosis (cell division that produces sex-cells, like sperm and egg cells, which carry half the normal number of chromosomes), it latches on to certain spots of the DNA strand and defines how the DNA recombines (how it is shuffled, and where snippets are added or removed).

Like all alleles, PRDM9 comes in different variants, that arise from mutations. And, these alleles are what Loya and team studied.

PRDM9 Alleles

Before continuing with Loya et al.'s paper, I would like to mention the frequencies at which different PRDM9 alleles are found in modern human populations. For an enlightening paper on the role of PRDM9, see Paigen and Petkov, 2019; it informs in an over-simplified manner that the alleles found globally "generally agree... with their “out of Africa” origins." It says so because Africans have a greater variety of alleles: 50% of allele A, 13% allele C and the other 37% a mix of different variants. The non-Africans, after the bottleneck caused by the OOA migration are almost exclusively A: "with ~90% alleles A and B (mostly A), which only differ in a serine/threonine substitution that does not affect binding specificity" Regarding the Denisovans and Neanderthals, this paper states that their "PRDM9 alleles closely related to rare contemporary alleles that are limited to Africans." I found this rather vague, so I looked for more information on alleles and their frequencies in different populations.

Alleva et al., 2021 investigated this matter and found that the A allele is the most frequent one, ranging from 90% in Finns (highest value) to 48% among Yoruba people in Ibadan, Nigeria (YRI) and 49% in Luhya in Webuye, Kenya (LWK), in between are the Chinese Han with 75% (CHB). The B allele, on the other hand, is higher among the Chinese CHB, with 13% and YRI = 7%, and much lower in other populations (0 to 3%). Another variat named L14 allele is found in Africa, ranging between 11% in LWK, to 3% in YRI, and not present in other non-Africans. The most relevant one is the C allele, it is found in all populations: YRI = 10%, LWK = 8%, CHB = 6%, and also in South American Peruivians from Lima (PEL) = 8%, lower in Italians from Toscana (TSI) and Pakistanis from Lahore (PJL), both with 4%, lowest among Finns with 2%. So, it isn't an exclusive African variant, it is rare in Europe, but present there.

Then come many other variants at lower frequencies in all populations. They add up to a 16% of the global PRDM9 alleles, but they vary between different populations. Most of them at levels of 30% and 31% are found in Africans (LWK and YRI respectively), followed by Tuscans (TSI) with 14% —probably due to recent African admixture in historic times. The rest of the world has levels of 7% to 9% (PEL, CHB, FIN, PJL).

Some rare alleles are exclusive to certain populations, like M1, found at "enriched" levels in Chinese HAN yet absent in all other populations. The D allele was only found in Finns.

Below is an image depicting a Table from the Supplementary Material of Berg et al., 2011, which also provides some frequencies for Europeans, Indians from India, and Africans. Notice the different variants and how the rare ones are exclusive to some regions. I highlighted those higher than 4%. It does not include variant M1 (from the Chinese).

Figure S1 in Berg et al. 2011

Having said this, let's get back to Loya et al. who report that the most common one, PRDM9-A is found in all modern human beings at frequencies of 85% to 90%, and the PRDM9-C allele is less frequent globally, but high in Africa at 10% to 15% together with other rare variants (at 35% in Africa).

Their Ghostbuster algorithm provides evidence that the best model to explain this distribution is the following: there was an original basal lineage carrying the A allele as ancestral variant. Then it split into two stems, one carried the allele A (population HA), the other the allele C (population HC). These populations split and remained separated around one million years ago, then they met 300,000 years ago, merging, this was the time when modern humans, Homo sapiens appeared. Below is Figure 5c, from their paper, showing this scenario:

Original caption (c) Proposed model of deep human history showing the divergence and persistence of different PRDM9 lineages across Denisovans, Neanderthals, and ancestral humans over the past >1 million years. HA and HC denote population groups carrying PRDM9-A-like and PRDM9-C-like PRDM9 alleles, respectively. Fig 5c in Loya et al. 2026

The Neanderthals and Denisovans form part of population HC. Through introgression, the C allele entered modern humans, and the Neanderthals, through admixture with us, got the A allele and a 10% contribution from humans to their genome. Regarding the Denisovans, "Denisovan PRDM9 alleles are distinct from present-day human genotypes, and at least one allele is more similar to PRDM9-C than to PRDM9-A."

The authors suggest that HA population contributed with 20% of the human ancestry. It was a larger population.

Who were these HA people?

The HA population lived for a long time, starting over one million years ago, and surviving until at least 300,000 years ago. It was widespread as it had to mix with the other HC population (in Africa, in accordance with the African origin of modern humans and demanded by the Out of Africa theory), and also in Eurasia, where it met the Neanderthals and admixed with them too. However it seems not to have been present in Eastern Asia because it didn't mix with Denisovans (though, later, modern humans and Denisovans mated there).

Considering the higher prevalence of the A allele outside of Africa, and the lower frequencies inside of Africa, the HA population was surely an established Eurasian lineage that partly migrated into Africa blending with HC there.

Due to the controversy around Mid Pleistocene fossils (the "muddle in the middle" conundrum), it is not an easy task to assign fossils to the HA population. Loya et al. timidly propose that "fossils attributed to H. heidelbergensis are a candidate for HA, as they possess the geographic and temporal properties suggested above." Perhaps they were a group living in the Europe, and the Levant, originating and descending from Eurasian Homo erectus, like the Homo antecessor whose remains have been found in Spain

Pending questions

If so, we can imagine that the superarchaic group from which HA and HC split is also Eurasian. This split took place long ago. Could the basal lineage be an ancestor of Homo erectus, a group that was present in Eurasia, like the Dmanisi hominins from the Caucasus?

The figure above shows that the subpopulation within HC that mixed with HA to form modern humans within Africa split from the remaining HC branch before this stem, in turn, split into Denisovans and Neanderthals. So, according to this figure, there is an "out of Africa" around 600,000 years ago, of the branch leading to Denisovans and Neanderthals, that totally removes this HC group from Africa (note that Africans do not carry Denisovan or Neanderthal alleles).

As mentioned further up, C allele is present at similar levels in Africa 10-8% as in Asian and American populations: Chinese and Peruvians 8-6%, but is much lower in Europe. Why? wouldn't admixing with Neanderthals (carrying C allele) enhance the frequency in Europe and Western Eurasia? Perhaps the predominance of a large HA population in Europe contributed to a higher frequency of A allele there, diluting the C allele frequencies.

Perhaps the most interesting part is the 35% of rare variants found in Africa, these probably derive from mutations in other archaic branches like H. naledi or Homo rhodesiensis they did not come from the HC or HA populations. They were archaics that blended into the modern Africans, and added their ancient, rare PRDM9 alleles. Another piece of evidence suggesting that diversity within Africa is due to the introgression of archaic hominins.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

On archaic introgressions — John Hawks and the 80:20 theory

There is a gradual shift among mainstream scientists who are now accepting that modern humans are the result of different contributions from ancestral populations that took place within Africa (or possibly in the Levant), archaic people different from both Neanderthals and Denisovans.

An example of the above can be found in a recent article posted two days ago by renown anthropologist and professor at the University of Wisconsin–Madison, John Hawks, titled Modern humans evolved from an 80-20 mixture of two deep streams — Genome models are converging on a scenario in which two stem lineages separated 1.5 million years ago, and merged together 300,000 years ago.

In his post, Hawks looks into the different papers published over the past few years, that suggest ancient hominins admixed and the outcome was the human lineage. It is nicely written and mentions some of the research articles that I have mentioned in previous posts (like Padilla Iglesias, 2025, Mallick, 2016, Ragsdale, 2023, Rogers, 2026).

Hawks explores the suggestion that we evolved from the admixture of two archaic lineages, an event that took place 300,000 years ago. One of these groups provided 80% of our genes and the other one, 20%. He also suggests that the Homo naledi (Hawks played an important role in the discovery and analysis of this hominin) could be a possible ancestor of the older archaics because it split from our lineage around 1.5 million years ago (Mya). Other options are the Bodo and Kabwe hominins, but he points out that there is a gap in the fossils corresponding to the period spanning 1 to 1.5 Mya the date the two groups or "stems" split apart. Did they evolve "... from earlier, H. erectus-like ancestors"? or, as he suggests, (and this is very interesting) the basal ancestors were not living in Africa!

An Eurasian Origin

I will quote Hawks directly (highlighting is mine):

"A third scenario places the stem populations on different continents. Suppose that one of the stems, which gave rise to Neanderthals and Denisovans as well as the majority of the modern human gene pool, inhabited southwest Asia. The minority stem might then have inhabited part of East Africa or southern Africa. Putting the stem populations on different continents separated by the Sahara Desert would provide a biogeographic reason for the million-year separation of the stem populations. With low levels of gene flow like the Ragsdale model, it would be a multicontinental multiregional evolution scenario.
This scenario might fit the idea proposed by José Bermúdez de Castro and María Martinón-Torres in a 2013 paper, which proposed two “hotspots” of evolution for Pleistocene Homo, one in Southwest Asia and the other in East Africa. In their concept, the Levantine Corridor was an area with near-continuous habitation that gave rise to successive range expansions into Europe and other parts of Eurasia. Homo antecessor as represented in Spain was an early dispersal, possibly the first, and Neanderthals and Denisovans were later dispersals. The Levantine Corridor could well be the home of a Cousins-like Population A."

The suggestion of a population in the Middle East, spanning modern Israel, Lebanon, Syria, Jordania and probably Irak, and Saudi Arabia, is the origin of modern humans is very reasonable. That area was probably a vast refugium (or multiple refugia) safe from the rigors of harsh ice ages in northern Europe, and the desertification (for a recent paper on the climate in Eurasia see Zan, 2024) and arid conditions that prevailed for most of the Pleistocene. From there, pulses of hominins moved out, fanning into Eurasia (H. erectus, Dmanisi, the Flores Island hobbits, Homo antecessor, Neanderthals and Denisovans).

In the meantime, primitive archaics populated Africa south of the Sahara (H. naledi and H. rhodesiensis) which later admixed with modern humans, providing diversity to the African genome. Below is a map depicting this scenario.

The origin of humans in the Levant . A. Whittall © 20206>

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

The history of humans in Africa (a new preprint)

The research article posted on July 28, 2026 in non-peer-reviewed bioRxiv, titled Ancient tree-topologies and gene-flow processes among human lineages in Africa, by Gwenna Breton et al. (doi: https://doi.org/10.1101/2024.07.15.603519) looked into the evolution of human beings inside Africa. The authors sampled the genes of different African populations and using AI ("extensive simulations and machine-learning Approximate Bayesian Computation (ABC)"), they compared different models to explain how our species evolved in that continent over the past couple of million years. The paper reports that the best-fitting model is one where "tree-like population histories with long periods of drift separated by short pulses of unidirectional gene-flow better explain the data than continuous gene-flow."

It should be noted that the authors state that their model did so "Without invoking archaic admixture"

The paper notes that instead of repeated migrations of populations, that lead to intermingling, the best models are those with "instantaneous gene-flow processes." This implies that isolated populations co-existed in Africa and time to time they quickly exchanged alleles: " ...our results unambiguously favor an evolutionary history of African lineages ancestral to a variety of Central and Southern African populations where Homo sapiens populations experienced long periods of isolation and drift, followed by short periods of possibly asymmetric gene-flow, which may in turn have induced some reticulations among lineages"

The first modern humans, according to this paper, were the Khoe-San (KS) people who are now found in South Africa and Botswana, they split from a basal lineage between 265,000 and 398,000 yearws ago. Then the rainforest hunter gatherers of Western Africa split from those of Central Africa (called RHG and RHGn, respectively, in the paper), between 127,000 and 190,000 years ago.

These findings differ from those of Ragsdale, A. P. et al., (2023) and the authors note that their dates are olde than the 110,000-135,000 year timeline for the KS split given in that study, and attribute the differenct to Ragsdale et al's use of "very ancient genetic structures, long before Homo sapiens emergence, a feature that is unspecified in our scenarios which considered simply a single ancestral population in which all extant lineages ultimately coalesce."

They agree that there are limitations to their study and that a "large effective population ancestral to all extant populations here investigated" could have existed, and future studeis should look into "very ancient substructures and reticulations within our ancestral population, prior to the original divergence between Southern and Central African populations." The authors also note that "where the ancestors of extant Khoe-San populations lived at that time remains unknown and is nevertheless needed to further elaborate possible scenarios for the causes of the genetic divergence here inferred... similarly as above for the Northern and Southern Khoe-San populations divergence, where the ancestors of extant Eastern and Western Rainforest Hunter-Gatherers lived remains unknown..."

We tend to consider modern Africans as being the same as the ancient ones, those living 100,000 or 200,000 years ago. But, they were not the same. There is a vast space of time of thousands of generations of people during which the genes accumulated mutations, admixing with archaics, and evolving under the pressure of natural selection. The paper recognizes these factors: "... Central African Rainforest Hunter-Gatherer and Southern African Khoe-San populations have had, respectively, extensive time for selection processes, including adaptive introgression processes, to have influenced independently both groups of populations as well as populations within each group separately."

Archaic admixture

The authors state that there is no need to invoke archaic introgression to account for the diversity in the genetic makeup of the Central and South African people. In fact, they "... did not explore possible contributions from unsampled lineages, whether from non-Homo sapiens or from ancient “ghost” human populations, and therefore cannot formally evaluate such possibilities."

But they know that archaics contributed even though they don't say so: they acknowledge that their method can be used to evaluate "scenarios comprising possible contributions from ancient or ghost unsampled populations; where the burden of proof lies on showing that scenarios with archaic admixture fit specific parts of the data significantly better than scenarios without such archaic admixture."

Interestingly, regarding the archaics and their contribution to modern Africans, the authors note that there is no ancient DNA available from ancient African fossils and that this lack of data is a handicap: "In any case, the complexification of scenario-specifications to account for possible past “archaic” or “ancient” introgressions will not fundamentally solve the issue of the current lack of reliable ancient genomic data older than a few hundreds or thousands of years from Sub-Saharan Africa 3,110,111. Indeed, analogously to archaic admixture signals that were identified outside Africa only when ancient DNA data were made available for Neanderthals and Denisovans, we imperatively need to overcome this lack of empirical ancient DNA data in Africa to formally test whether, or not, ancient human or non-human now extinct lineages have significantly contributed to shaping extant African diversity." I am convinced that when hard data is obtained from African fossils, there will be many interesting surprises.

Another finding is that the mixing after long periods of isolation took place around the time of the purported Out of Africa migration of modern humans: "Interestingly, we found strong indications for almost synchronic events of introgressions having occurred during the Last Interglacial Maximum in Africa 73, between ∼85,000 and ∼135,000 years ago... They involved gene-flow between lineages ancestral to Khoe-San populations and ancestors of Rainforest Hunter-Gatherer neighbors on the one hand and, on the other hand, between lineages ancestral to Khoe-San populations and the lineage ancestral to all Rainforest Hunter-Gatherers... we estimated that the instantaneous gene-flow event between the ancestral Rainforest Hunter-Gatherers lineage and that of their extant neighbors seemingly occurred synchronically to the genetic Out-of-Africa. This would imply that possible climatic and ecological shifts at that time may not have only induced population divergences and displacement, but may also have triggered population gene-flow."

This also implies that there was a blending of isolated populations, that contributed diversity to those remaining inside of Africa while another population that migrated out of Africa carried fewer alleles, and less diversity with them (see my post On the Diversity of Africans). It also implies that the Khoisan people were split from the rest, and isolated in Southern Africa with their genetics "enriched" by intermingling with other formerly isolated groups. Did they also interact with archaics like the Homo naledi?

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

Denisovans were very tall people!

An article published in Science on August 12, 2026, by Brown (Denisovans towered over their contemporaries and munched a bunch of meat) caught my eye. In it, Taylor Brown mentions two papers published in bioRxiv, one of which describes leg bones dredged up from the seabed between the Island of Taiwan and mainland China. The protein in the bones showed they were Denisovans, but they were also big. The article quotes an archaeologist (University of Washington), called Ben Marwick who said that "They were very tall! Previously we knew that Denisovans had big brains from the skull and teeth finds, and now we know they had big bodies also." But, how big?

Original caption reads: Fig. 2. Penghu 2 and Penghu 3 hominin leg bones. Anterior (a), medial (b), and distal (c) views of the Penghu 2 right femoral shaft. Anterior (d), lateral (e) and distal (f) views of the Penghu 3 right tibia. Symbols: m=medial, p=posterior. Scale bar=10cm. . Yousuke Kaifu et al., 2026

The paper in question is the following: Denisovan leg bones from Taiwan reveal large body size, Yousuke Kaifu, Chun-Hsiang Chang, Yumeko Tarusawa, Rikai Sawafuji, Shiori Yonemoto, Shigeru Shimamura, Masanaru Takai, Reiko T. Kono, Cheng-Han Sun, Cheng-Hsiu Tsai, Minoru Yoneda, Takumi Tsutaya, bioRxiv 2026.08.07.743438; doi: https://doi.org/10.64898/2026.08.07.743438.

The specimens are known as Penghu 2 and Penghu 3, different people, about 43,000 to 45,000 years old. The paper notes that they are similar to the Neanderthals, and different to Africans and also to the other hominins found in East Asia: "Among the premodern Homo groups, an anteroposteriorly thickened (non-platymeric) proximal shaft characterizes Neanderthals and in particular Middle Pleistocene European Homo, as compared to African Early-Middle Pleistocene Homo (Fig. 4b). Penghu 2 is similar to the Middle Pleistocene European Homo in this respect. Such a trend is not evident in the available small sample of H. erectus and Middle Pleistocene Homo from eastern Asia... Penghu 3 exhibits a strong tibial posterior pilaster, a shared feature of European Middle Pleistocene Homo and Neanderthals, as well as some Upper Palaeolithic modern humans"

This seems reasonable, Denisovans are a sister clade to modern humans and Neanderthals. We appeared later than Homo erectus, and should therefore share similar features.

The two specimens, who may have been male, were tall and big: Penghu 2 was 1.8 m tall and weighed 83 kg (5.9 ft. - 183 lbs ) and Penghu 3 was 1.9 m tall and 91 kg (6.23 ft - 201 lbs), similar to the largest known European or African hominins from that period, but much taller and massive than the Asian erectus.

The authors suggest that "large-bodied Denisovan individuals inhabited both northern and southern regions of Asian mainland, and large brain sizes seen in some Denisovan individuals (Harbin 1, and possibly other specimens such as Xujiayao 6 and Xuchang 1) was at least partly a consequence of their large body sizes, as has been suggested for Neanderthals."

Comments

This study is very interesting. First of all, we have a random sample of two specimens, out of a Denisovan population, and both are very, very tall, even by modern standards. Could they have inspired tales of giants among shorter hominins that lived in the same region? Did they move on, into America and settle there, giving birth to myths among the later arrivals (modern humans) like the Si-Te-Cah, a race of red-haired giants mentioned by the Northern Paiute people in Nevada, U.S? (see my posts about them)

Large body sizes are efficient when living in a cold environment (the paper mentions this), so they were well equipped to deal with the icy conditions that they would have come across on their journey north, into Eastern Siberia and Beringia.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

The Island of Giants, Samson, and the Falklands-Malvinas

Recently, it was Falkland Islands' Day in the UK, commemorating the discovery of the islands by John Davis on August 14, 1592. As expected, with these islands claimed by Argentina, as the Islas Malvinas (claim that, unfortunately, led the last military dictatorship to invade them in 1982, in a stupid and sensless war that caused an irreparable breach with the islanders as well as hundreds of young men, dead and maimed, on both sides, Britain, and Argentina), their discovery is also contested.

The Argentine side argues that they were discovered by a cartographer named Andrés de San Martín who, on board of the ship San Antonio, made the first map of the islands. This ship was part of Magellan's voyage to discover a soutern pass from the Atlantic into the Pacific Ocean, to reach the spice islands of the Far East. In this post we will look into this alleged discovery.

The Story of the San Antonio

When Magellan left Spain in 1519, he had five ships, Trinidad, San Antonio, Concepción, Santiago, and Victoria (the only one to make it back to Spain after sailing around the globe). Of these, the Santiago capsized on the coast of Patagonia at the mouth of the Santa Cruz River in 1520 with no loss of life. y la Santiago. The four remaining ships set sail after wintering in San Julian, in what is now Argentina's Santa Cruz province and entered the Strait of Magellan in October 1520.

Magellan sent the Concepción and the San Antonio into the Strait to explore it. They reached a spot called Cape Forward, that jutted out into it, and marked a place where there were two separate channels to follow. They returned, reported their findings to Magellan who advanced with his fleet to the Cape.

Magellan decided to send the San Antonio and Concepción to explore the two channels, but he first called his pilots and captains to a meeting. Esteban Gómez, the pilot of the San Antonio (which was commanded by Alvaro de Mesquita, Magellans nephew) expressed his point of view: the fleet lacked supplies and was not equipped to face the stormy seas that lay ahead. He proposed that they turn back, and return to Spain. Magellan would have none of this. He ordered the two ships to proceed ahead into the channels. The Concepción returned, but the San Antonio vanished. Magellan waited for it, and finally decided to head west into the Pacific. The loss of the San Antonio was a blow, with a displacement of 120 tons, it was the largest ship in the fleet, and it was packed with supplies and food. This led to famine for the remaining ships as they sailed across the Pacific.

What Magellan didn't know was that the San Antonio had turned around, and in the dark of night, sailed past the ships in the Strait, and headed by the route of Guinea (Africa) to Spain. The crew, led by Gómez seized Mesquita and took control of the ship on November 8, 1520. They reached Seville and complained about the behavior of Mesquita and Magellan, accusing them of treason against the King of Spain (both were Portuguese). Mesquita was sent to prision, and only released when the Victoria returned to Spain in 1522 and its crew, led by Sebastián Elcano told the truth.

During the return trip, from the Strait, to Africa, is where the mythical story of the discovery of the Falklands arose.

There is No written record or map or entry in the ship's log mentioning any islands. But the Argentine version is that they discovered some islands during a stormy spell, and named them San Antón Islands, or Sanson (Samson) Islands.

An article published in the Boletín de la Socieded Geográfica de Madrid, 1899 Volume 41, Annex VIII, p 212 in Pigafetta's First Voyage Around the World quotes a contemporary letter written at the time of the arrival of the San Antonio to Seville:"prendieron al dicho Mezquita en ocho de Noviembre del dicho año veinte é vinieron derechamente á este puerto... Sevilla 12 de Mayo 1521" ("They arrested the said Mezquita on the eighth of November of the said year '20 and came directly to this port... Seville, May 12, 1521") no reports of islands or anything along their voyage back to Spain. They didn't even head north, to San Julián to rescue two men (Cartagena and Calmette) who had been cast asho, in exile, by Magellan to punish them (Cartagena was the King's overseer, and Calmette a priest) for mutiny.

The islands, were not mentioned during the following expeditions to the Strait, conducted by Spanish navigators. This is added proof that they had not been discovered.

Thevet

The map often attributed to the San Antonio is, in fact, the work of André Thevet (1502?–1590). Thevet was a Frenchman, a Franciscan friar, and cosmographer to several kings of France. He traveled extensively and visited South America in 1555. Upon his return to France, he published his experiences in the books "Les singularitez de la France Antarctique" (1557) and "Cosmographie Universelle" (1575). I have discussed Thevet in my book, because he mentioned two strange Patagonian creatures, the Succarath and the Pyrassouppi.

Thevet also wrote a book titled "Le grand Insulaire et pilotage d'André THEVET, Angoumoisin, cosmographe du Roy, dans lequel sont contenus plusieurs plants d'isles habitées et deshabitées et description d'icelles" (The Great Insular Book and Pilotage by André Thevet, native of Angoulême and Cosmographer to the King, containing various plans of inhabited and uninhabited islands and descriptions thereof), a description of the most important islands in the World. It is a manuscript, unpublished, and the original handwritten text can be seen online here. It mentions on page 269, the Isles des Geants (Islands of Giants) he states that they were discovered by the Portuguese sailors that accompanied Magellan on his voyage and that a captain from the Portuguese city of Lisbon told him about them. See the text below:

He included the map, naming them Samson Islands - Giants Islands, shown below (see plate on p. 268)

In the Islario general de todos las islas del mundo (General list of all the islands in the world) by Alonso de Santa Cruz (1505-1567), first published in 1541, the author mentions the islands of Patagonia as reported by Magellan and (see it here online), on page 552, says "...they discovered some islands that were to the east of San Julian, by eighteen leagues and they named them Islands of Samson and the Ducks because on them they found many and very fat that could hardly walk, and half-plucked of which they took many for their voyage and are at 51 degrees of latitude..."

The location is far too close to the coast of Argentina to be the Falkland-Malvinas; 18 leagues is 87 km or 54 miles. The islands are 571 km - 354 mi from San Julian.

The ducks were probably penguins. Looking at the map published by de Santa Cruz, reproduced below, you can see Cabo Blanco, located at 47° S on the upper left side of the map — north is left, south is right (see it in Google maps), a Rio de los Trabajos river (perhaps modern Rio Deseado), and nearby, the Sanson Islands.

Interestingly, just south of Rio Deseado is a Provincial Natural Park, called Isla Pinguino (Penguin Island) with some other smaller islands, that protects a penguin colony there (see it in Google maps). Could these bre the Samson-Ducks islands?

The name Samson, and Giants Islands surely came from the large-sized Patagonian natives, the Patagons reported by Magellan's expedition. Samson, is mentioned in the Bible (Book of Judges ch. 13-16) as an extremely strong man, so he is comparable to the large, muscular Patagonian natives.

Summing it up, I doubt that the San Antonio, or Magellan's ships set eyes on the Malvinas-Falklands Islands. Thevet probably took note of de Santa Cruz's Islands book and included it in his own version.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

Misterious Middle Eastern (non-African) homo, not Neanderthals or modern humans (Denisovans?)

Two 2021 papers published in Science, one by Israel Hershkovitz et al. (A Middle Pleistocene Homo from Nesher Ramla, Israel, Science 372, 1424-1428. DOI:10.1126/science.abh3169 🔒), the other by Yossi Zaidner et al. (Middle Pleistocene Homo behavior and culture at 140,000 to 120,000 years ago and interactions with Homo sapiens.Science 372, 1429-1433. DOI:10.1126/science.abh3020 🔒) describe hominins living in what is now Israel using stone tool technologies associated to both modern humans and Neanderthals yet different two both of those lineages. These fossils have been dated to around 120,000 ti 140,000 years ago (120 - 140 kya), in the Middle Pleistocene (or MP).

Not Modern Humans or Neanderthals

The second paper states that "The evidence from Nesher Ramla demonstrates that late MP Homo fully mastered advanced Levallois technology that until only recently was linked to either H. sapiens or Neanderthals." It also mentions that this stone knapping technology was due to contact between different human groups ("cultural diffusion"), and this implies that they admixed: "These results are consistent with a growing body of genetic studies suggesting that a gene flow existed between divergent archaic Homo populations and H. sapiens during the late Middle and early Late Pleistocene. Our findings provide archaeological support for close cultural interactions between different human lineages during the Middle Paleolithic period and suggest that contacts between MP Homo and H. sapiens had already occurred prior to 120 ka."

The first paper proposes that these Middle Pleistocene hominins, which they call NR, after Nesher Ramla, the site in Israe where they were found, are distinct from both modern humans and Neanderthals. They present the following unrooted phylogenetic tree (Fig 2b) which they describe as having "NR-1 close to the origin of the branch leading to African MP Homo, close to the split from the H. erectus branch and to European MP Homo and Neanderthals (including Atapuerca SH), and far from early and recent H. sapiens".

Phylogenetic tree for the Nesher Ramla homo. Fig 2b in Hershkovitz et al., 2021

Notice how it is midway between the Eurasian Homo erectus, and the African MP hominins (like the ergaster, and H. rhodesiensis).

The authors point out that their analysis of the fossils show that they are a mossaic, with "a unique combination of archaic and Neanderthal features, supporting the existence of a local, Levantine population at the final MP... showing that an affiliation of the NR fossils with early and recent H. sapiens is highly unlikely, but that it is impossible to establish whether NR fossils are more likely to be classified as MP Homo, Neanderthal, or H. erectus (the latter for the parietal only)" So, it is different from the Homo sapiens lineage yet similar to erectus, Neanderthals or MP homo. They probabilities assigned by the study are: 41% Homo erectus, 34% MP Homo, and 25% Neanderthal.

Due to their recent age of 120 to 140 ky, the researchers suggest that they are the last survivors of an ancient Southwestern Asian homo group that could include other specimens found in the Middle East, like the Quesem and Zuttiyeh Caves individuals, and also Tabun Cavre. If this is the case they lived here between 420 and 120 ky ago. They could have admixed with the Homo sapiens of Misliya Cave, and also the Neanderthals of Skhul and Qafzeh.

They propose that the Nesher Ramla Homo is the source of the populations found in Western Europe such as the Arago and Atapuerca Sima de los Huesos (SH) specimens: "... contribution of Levantine Homo groups carrying Neanderthal-like traits to European Homo lineages. The NR Homo, carrying Neanderthal-like traits, could thus represent the “source” population... according to which Western Europe was repopulated through a series of successive migrations." The paper's bibliography cites de Castro et al., 2019, who proposed an explanation for the differences between them: "...Whereas the SH teeth present an almost morphological identity with European Neanderthals, the Arago teeth exhibit a combination of plesiomorphic as well as some Neanderthal-derived features. In addition, the Arago crown dimensions are remarkably larger than those from SH, the differences being statistically significant for most variables. We hypothesize that during the Middle Pleistocene the European continent was settled at different points in time by hominin groups coming from Southwest Asia, probably from a common mother population evolving in this latter region."

What I find interesting, is that these are Eurasian hominins, all of them, none are African. It somehow hints at a Middle Eastern origin of the population that led to Neanderthals, with a link to Homo erectus, and which is also associated with Mid Pleistocene Western European hominins like the Atapuerca Homo antecessor. The MP African hominins are not even close to these Levantine people.

The paper does not state it, but it follows logically, that modern humans could have originated in this region too.

Denisovans?

A very smart suggestion posted by John Hawks is that these Nesher Ramla people were Denisovans!

Hawks wrote: "One possibility may fit well, and I've not seen anybody mention it. We already know that there was a third population out there: the Denisovans. The DNA data from central Asia show us that Neandertal and Denisovan populations were sometimes in contact with each other, across more than 100,000 years of time, and that they mixed where they met. We do not know what the geographic range of Denisovans may have been across the southern tier of Asia. Many researchers have emphasized the Denisovans as an “eastern” population, due to the high representation of Denisovan ancestry in island southeast Asia and Oceania. But we have very little fossil evidence about the ancient populations of India and Pakistan, and the known Neandertals from Teshik-Tash, Uzbekistan and Shanidar, Iraq, are more recent than 100,000 years. So why not Denisovans in the Levant before 120,000 years ago?"

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

The odd thing about Homo rhodesiensis

A research paper published in Nature in April 2020, titled Dating the skull from Broken Hill, Zambia, and its position in human evolution, that I didn't spot at that time just when the pandemic of Covid 19 locked us all up in Argentina, dated the original skull of Homo rhodesiensis, known as BH1 (Broken Hill 1) and it is much younger than expected for such a primitive hominin.

The paper by Grün, R., Pike, A., McDermott, F. et al. (Nature 580, 372–375 (2020). https://doi.org/10.1038/s41586-020-2165-4) reports how they studied the Pleistocene skull discovered in 1921 in a mine at Broken Hill (now Kabwe), in what then was known as Rhodesia (now Zambia), from a site known as Broken Hill. The specimen had been described by Woodward that same year and named the Rhodesian man (Homo rhodesiensis), a new species. Currently it has been included into the same line as the European Homo heidelbergensis and therefore, the ancestor of Neanderthals and maybe, also modern humans.

Those who studied it, dated it to around 500,000 years ago but did so assuming it was similar to the European Homo heidelbergensis which is that age, and also using estimations based on correlations with other sites in Africa and fauna found at the site.

That age makes it old enough to be an ancestor of both Neanderhtals and Homo sapiens, and inside Africa it provides support to the African origin of both species. However, and this is a crucial point, the original site at Broken Hill was razed as it was mined away, and in the 1920s, dating of ancient specimens was more guess work and good will than science. So, Grün et al., decided to date it with modern technology, analyzing the skull itself.

A very recent primitive hominin

They found that its "best age estimate of 299 ± 25 thousand years" which is coincidental with the date given for the first modern humans, in North Africa, and the very primitive Homo naledi in South Africa! The authors write: "The result suggests that later Middle Pleistocene Africa contained multiple contemporaneous hominin lineages (that is, Homo sapiens, H. heidelbergensis/H. rhodesiensis and Homo naledi)" which is quite surprising. They try to play down this odd situation by saying that in Asia the situation was similar: "where Homo neanderthalensis, the Denisovans, Homo floresiensis, Homo luzonensis and perhaps also Homo heidelbergensis and Homo erectus were found contemporaneously". Indeed, but it should be noted that the minute Flores and Luzon people were isolated on insular habitats. Erectus was on the brink of extinction, and regarding Denisovans and Neanderthals it is widely accepted that they admixed and trysted with modern humans, and with each other (and, regarding Denisovans, very likely with erectus). But when it comes to Africa we are asked to believe that modern humans appeared and never mixed with any of these archaics that populated the continents.

But the researchers can't escape the fact that a very recent yet primitive hominin co-existed inside of Africa with modern humans, the write: "The age estimate also raises further questions about the mode of evolution of H. sapiens in Africa and whether H. heidelbergensi/H. rhodesiensis was a direct ancestor of our species."

Correct! If modern humans had already evolved 300 ky ago, then, how could the Homo rhodesiensis with that same age, be our ancestor? Furthermore, it has prmitivie traits: solid skull (relatively large at 1280 cm³, low forehead, prominent brow ridges.

A paper by Balzeau et al., 2017 studied the shape of the BH1 skull, and its dimensions and thickness, comparing it to other archaic hominins. It reported that its thick parietal bones resemble the Homo erectus and is unlike the "morphology observed in Neandertals and H. sapiens." Together with the Petralona skull (equally primitive, but found in Greece and also recent —286 ky) the Rhodesian man shares "a thick frontal torus or a thickened postero-lateral parietal bone... The main peculiarity of BH1 is related to its large, thick frontal torus and to high thickness values for the vault." The paper concludes that it is distinct from Neanderthals and modern humans and similar to the Homo erectus sensu lato (in a broad manner): ".. this fossil does not exhibit the apomorphic conditions observed in either Neandertals or among H. sapiens."

See the skull below, and judge for yourself:

Side and front views of the Kabwe skull, Broken Hill, Rhodesian man. Source

I have posted often, that Africa was a melting pot of primitive hominins, blending into the stock of African Homo sapiens, and adding diversity to its genetic pool.

H. Rhodesiensis

Some scholars have proposed a new name for the specimen, dropping "Rhodesian man" and renaming it Bodo man, or Homo bodoensis (yes, wokeness seeks to decolonize anthropology and remove the tainted, imperialist reference to Cecil Rhodes who gave his name to Rhodesia). However, this will not solve the issue of its recent age and perhaps, its location on a dead end branch of hominins. In a recent post we mentioned a paper that analyzed the shape of the faces of humans and ancestral hominins. It reported that "The evidence instead suggests that the large and non-H. sapiens-like faces of Bodo and Broken Hill 1 ... represent taxonomic diversity in the African MP record, which could exclude the large fossils assigned to H. heidelbergensis and H. rhodesiensis as representative of an ancestral morph for H. sapiens." Meaning that they are not our ancestors.

Finally, the similarity of the Rhodesian man and Petralona was noted by Falgueres et al., 2025 and they place them together, it in a parallel and independent lineage to that of Neanderthals: " our view, the similarities in midfacial and cranial contours between Kabwe and Petralona and the absence of true Neanderthal apomorphies in their midfaces suggest that European and African samples often assigned to H. heidelbergensis (Stringer, 2012; Rightmire, 2017) remained closely related during most of the Middle Pleistocene, alongside and independent of the Neanderthal lineage. Our results from dating the matrix attached to the Petralona cranium suggest that like the Kabwe cranium, the Petralona cranium may date to about 300 ka, consistent with their persistence into the later Middle Pleistocene" They also note that their work supports the notion put forward by Quam et al., 2023 and Ashton et al., 2016, that Middle Pleistocene hominins can be grouped into two separate lineages:

One lacking Neanderthal-like features, that includes the following speciments: "Mauer, Mala Balanica, Montmaurin, Visogliano, and one of the Arago mandibles" as well as the more recent Petralona skull. The second group has Neanderthal-like features, and comprises "the Atapuerca, SH fossils, Arago 2, and specimens from Payre, Ehringsdorf, and Bau de l'Aubesier."

Taken together the data seems to suggest that Homo heidelbergensis and the Rhodesian homo from Africa may not be linked to Neanderthals or modern humans.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

Blood Groups support an "Into Africa - Out of Eurasia" migration of modern humans

Rereading a research paper published in January 2025 (Mazières S, Condemi S, El Nemer W, Chiaroni J. Rapid change in red cell blood group systems after the main Out of Africa of Homo sapiens. Sci Rep. 2025 Jan 23;15(1):1597. doi: 10.1038/s41598-024-83023-0. PMID: 39848961; PMCID: PMC11757997), I noticed that the authors attempted to adapt the facts to the Out of Africa theory even though they point in an opposite direction.

The abstract states that "we noted that the blood group allele diversity in the Neandertals remained unchanged since 120,000 yBP, while H. sapiens conquered Eurasia with blood group alleles presently exclusive to non-African populations, suggesting they may have differentiated right after the Out of Africa, between 70,000 and 45,000 yBP. Notably, Ust’Ishim possessed unknown alleles that may illustrate the lost genetic heritage of the early Eurasians. Lastly, Neandertals shared a unique Rh haplotype from which we updated the current RHD phylogeny."

The paper notes that the oldest modern humans in Eurasia carry red blood cell variants that are absent in Africans: "the oldest genomes available (Ust’Ishim, Bacho Kiro, Zlaty Kun, Tianyan, Oase1, Kostenki14, 36-45 ka) show that H. sapiens entered in Eurasia with a set of functional alleles absent in Neandertals and Denisova, and already derived from their ancestral state... Presently, some of these blood group alleles are only found in non-African populations."

How could this be possible? If humans originated in Africa, the original pool should be found there, and the Eurasians, after a bottleneck due to the Out of Africa migration, should have fewer original variants. But instead, they find none of these variants in Africa.

The paper invents a standstill in Iran, the "Persian Standstill" where the new Eurasian humans evolved their own, distinct variants! Below is the paper's text:

"Given the wide distribution of the derived alleles amongst the earliest H.sapiens of Eurasia coupled with their total absence in Africa, a first scenario could assume that these alleles have emerged in African H. sapiens before the Out of Africa, spread in Eurasia, then disappeared in Africa only. It would be tempting to point to natural selection, but the joint disappearance of independent alleles coding for phenotypes of a wide variety of susceptibility to natural selection is unlikely. Moreover, given that the highest gene diversity is found in Africa and that the expansion of H.sapiens is characterized by an initial founder effect, these alleles should be found in African populations. An alternative could be an outbreak of red cell blood group alleles outside Africa right before the conquest of Eurasia. Genetic, lithic and paleoecological evidence from the Initial Upper Paleolithic indicated that the late phase of H. sapiens expansion outside Africa circa 70 − 60,000 years ago was followed by a standstill of at least 15,000 yBP in the Persian Plateau. Analogously to the Beringian occupation ca. 20,000 yBP during which the Beringian populations paused several millennia and differentiated before entering the Americas with genetic polymorphisms shared by Native Americans only, H. sapiens could have resided long enough in the Persian Plateau to differentiate. This view makes Western Asia a good candidate for the place of incubation of the functional ABO*O.01.02, FY*01, RHCE*03, KEL*02.07, FUT1*01.02 c.35 C > T, and GYPB*04 before expansion towards East (Tianyaun), North (Zlatý kůň, Ust’Ishim) and Western Eurasia (Oase1, Bacho Kiro) more than 45ky ago."

Why propose a standstill in Persia? An unexplored alternative is that humans originated in Eurasia, and lost part of their diversity as they migrated out of Asia and into Africa. There they settled in Eastern Africa and gradually spread into the continent, losing diversity as they advanced, and taking up archaic versions as they admixed with older homo lineages in Africa.

Neanderthals, Denisovans, and Humans

Surprisingly, the paper stated that "H. sapiens entered in Eurasia with a set of functional alleles absent in Neandertals and Denisova, and already derived from their ancestral state" so, they were present in modern humans in Asia and absent in our Neanderthal and Denisovan ancestors but, and this is relevant, "Neandertal and Denisova, two archaic hominid populations inhabiting Eurasia, expressed blood groups currently found in sub-Saharans... For two blood group systems, RH and KEL, the Neandertal alleles are almost exclusively found in modern populations from Sub-Saharan Africa" This suggests that the ancestral populations of Neanderthals and Denisovans, that originated in Asia, migrated into Africa, surviving through admixture with hominins inside AFrica. Their variants are now only found in the red blood cells of populations South of the Sahara. Outside of Africa, the larger population that evolved into modern humans mutated into a derived form, absent in both Denisovans and Neanderthals.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

East Asian hominins in the Pleistocene

A paper published by Vialet, 2018 attempts to understand how modern humans settled East Asia. It reports that different "lineages" must have interacted to explain the outcome. People from the islands in South East Asia could have contributed to the final admixture, depending on variable sea levels. Vialet analyzes a scenario with "demographic “sources” and “sinks” and where repeated colonization, extinctions and recombination may help explain the morphological variability of the population." Clearly, Denisovans, possible extant archaics like Homo erectus interacted with modern humans in this region.

Vialet cites Zanolli (2013) who notes a reduction in the size of teeth, which differs from the classic Asian Homo erectus s.s. (sensu stricto) and, interestingly ntoes that "Such a degree of dental reduction, from both a morphological (i.e., molar inter-ratio, loss of main cusps) and a metric perspective has been only identified in the Middle Pleistocene sample from Sima de los Huesos, in Atapuerca" in Spain. This means that teeth became smaller and are similar to the Atapuerca hominins, similar to those of modern humans, but, they don't know why!

Atapuerca remains have provided nuclear DNA that links them to Neanderthals, however, their mitochondrial DNA placed them very close to te Denisovans.

Zanolli is at a loss when it comes to explaining this situation, and speculate that the smaller teeth may be due to "more complex scenarios envisaging frequent population interchanges (possibly from sources to sinks) and/or regional variation by isolation" and also "possible connections between Asia and Europe."

They tend to believe that this dental reduction "is more likely resulted either from replacement or introgression by admixture with continental immigrating groups" Interestingly the African Homo heidelbergensis" and the Chinsese Homo erectus as well as the early specimens from Sanigran in Indonesia share similar featues.

These articles show that Denisovans, Neanderthals, and their Spanish ancestor in Atapuerca are linked to East Asian people. Somehow, these people replaced the classic Eurasian erectus. They surely interacted with them, admixing and replacing them. The hominins living in East Asia when modern humans reached the area (or evolved there, becoming modern humans), carried the signs of Eurasian people. Where does the Out of Africa myth fit in this panorama?

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

Homo ergaster wasn't an erectus

When the skulls of Homo erectus were discovered in Java, Indonesia, and Beijing, China ~100 years ago, they cemented the notion that modern humans evolved from these Asian hominins. They had relatively big brains, made stone tools, and were upright creatures. There was no similar fossil in Africa until a skull from Kobi Fora was discovered in 1975 and assigned to a new species which was named Homo ergater ("working" man) it was erectus-like (but differed from it), and it was old. This boosted the position of the African origin of humans, so it fitted neatly into the Out Of Africa (OOA) theory. The erectus of Asia were assigned to a branch named Homo erectus sensu stricto, or strict-sense erectus, the classic fossils. And they, together with the African ergasters were grouped as sensu lato (broad sense) erectus.

However, there has been a controversy ever since, and it is likely that they are not related, and that ergaster vanished in Africa without leading to the ancestors of modern human beings.

Antón 2014 summarizes the situation as follows: "Proponents generally allocate early African Homo erectus fossils dating from c. 1.8–1.4 Ma to Homo ergaster, mainly on the basis of smaller, more gracile crania and a lack of robust cranial suprastructures. In this schema, the geographic distribution of Homo erectus sensu stricto is restricted to Asian localities with a few examples (e.g., OH 9) of back migration into Africa. Homo ergaster includes the early African and Georgian specimens attributed by others to H. erectus and is therefore posited as the ancestor to Homo erectus, Homo heidelbergensis, and Homo sapiens..." This quote mentions an "Into Africa" back migration of Eurasian erectus, assigning an OH 9 fossil to this event. Remarkable!

The OH 9 sample was discovered by Louis Leakey in Olduvai Gorge, Tanzania, in 1960. It is approximately 1.4 million years old and has a large brain. In fact, it is the oldest hominin discovered until now, with a braincase capacity above on litre (it is 1,067 cm³). Leakey himself recognized it as erectus, calling it "Chellean man" because in the same sedimentary layer he uneaerthed Acheulean-like stone tools that are the hallmark of Homo erectus. Yet, it is unlike the ergaster fossils, and more or less overlaps with the last of them.

So, according to Antón 2014, ergaster is more gracile (smaller, slender) and the ancestor of the Dmanisi specimens in the Caucasus and ancestor of the sensu stricto erectus, implying that ergaster migrated out of Africa and originated the Homo georgicus found in Dmanisi, and the later erectines. However, it may be possible that the situation is exactly the opposite: that Homo ergaster descends from a Eurasian lineage of hominins, that originated the Dmanisi and erectines, and also migrated into Africa, leading to the hominins there, like Homo habilis, Homo rudolfiensis, and later, Homo ergaster. Real erectus originated and evolved in Asia, and migrated into Africa much later.

The Eurasian origin of Homo erectus

Chris Stinger in an interview (see it online here) suggests that the oldest hominins may not even be homo, they were something else, below I transcribe the relevant part, starting at 35' 45" in the video:

"...I mean it depends again what we call homo erectus. So on our Yunxian paper we obviously analyzed some of these earlier skulls and for us there is a clear erectus group based on the original fossils from Java and the ones from China. They are the real erectus if you like and then there are earlier things in Africa and in Dmanisi in Georgia. Those things on our analyses are probably not actually Homo erectus. They are something different. I mean, you know, we couldn't solve everything in our paper and say what they were, but we think that they are maybe not what you could call real erectus. There's another name around called Homo ergaster, workman, work human. Um, which was given to some fossils from Africa. Um, so sometimes called African Homo erectus, but also Homo ergaster. So that name is also around and could be applied to some of these earlier things. They would still be homo and therefore human in my view. But yes, there might be an earlier species before erectus. Um and in fact there might be even more than one species. So the Dmanisi material from Georgia 1.8 million years old that's been called the first Homo erectus out of out of Africa. Um some people and I'm beginning to join them think that that isn't really Homo erectus. It's something more primitive. Uh and that is the sort of creature that might have given rise to, you know, Homo floresiensis eventually, over in Flores or Homo luzoinensis eventually, over in the Philippines. So we could be looking at something that actually is prehuman even though at the moment there's a homo name attached to it. Some of those things could turn out to be prehuman. So for me, Homo erectus as we know it from Asia is is the first definite human for me and it dates back to you know one and a half million years probably. And then there are some things before where it's tricky to know whether they really are over the line to be homo. Some of them are still small-brained. Some of them still had adaptations in their skeleton which shows that they were either spending time in the trees or they had those remains still in their skeleton showed the traces of tree living. Whereas erectus for me as we know it later on is the first committed ground living in completely you know given up life in the trees and fully committed to life on the ground walking on two legs. So that for me is a big step forward and humans have done that. Some of these earlier things at around 2 million maybe are still in that process of transition."

Stringer mentions the paper The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans co-authored by himself and Feng et al., 2025. He is quite categorical about Dmanisi, and ergaster as being prehuman, or, not homo. They are possibly the same creature that originated the Flores Island hobbit or the Luzon hominins.

Ergaster

The label "Homo ergaster" has been assigned to a collection of fragmentary skulls, and assimilated to the erectine group. But these African "erectus" are different from the genuine Asian ones. They are more dimorphic (Sexual dimorphism is when males and females of the same species have different features, like colors, markings, and importantly in the case of primates, body sizes). Large, aggressive males can control more females by fending off rivals if they are bigger bodied. Modern humans are far less dimorphic as we don't rely on aggressive behavior but on cooperation between males. Asian erectus also show a lower dimorphism in comparison to African hominins.

Villmoare 2019 explored the dimorphism of erectines analyzing footprints and found African ones to be futher from modern human sizes, and Asian ones to be closer:

"Here we present an alternative, using footprints from near Ileret, Kenya, to assess the sexual dimorphism of presumptive African Homo erectus at 1.5 Ma... Our results indicate that East African Homo erectus was more dimorphic than modern Homo sapiens, although less so than highly dimorphic apes... As a species, H. erectus shows considerable temporal and geographic variation and, as Plavcan notes, only the African sample (in which he includes Dmanisi) shows a sufficiently high degree of variation to falsify the hypothesis that the species had acquired the relatively monomorphic human-like condition. In his analysis, the high variation in the African sample was driven by the small size of KNM-ER 42700 and the large size of OH 9 (which are separated geographically by 800 kilometers, and temporally by more than 300,000 years), although the Dmanisi sample also increases the variation within H. erectus considerably. The Chinese and Javan samples show a level of variation in cranial measures very similar to that seen in modern humans. The different patterns in the African and Asian samples raise the possibility of a change in patterns of dimorphism, and potentially social behavior, in the H. erectus lineage over time and/or across different geographic regions."

So, about 1.5 million years ago, the "erectus" in Africa had a high level of dimorphism, large males, smaller females. Could we be identifying the remains of males and females as belonging to different species? In other words, could these African hominins assigned to two different species belong to a single species with high sexual dimorphism?

The gracile bones assigned to Homo habilis and Homo rudolfiensis roughly 1.3 m tall (4.3 ft.) and weighing around 35 kg (70 lbs) could have been the females and the larger H. ergaster measuring 1.8 m (6 ft) and weighing 60 kg (130 lbs) could have been the males. Their brain sizes were very similar, smaller than that of genuine erectus:

Bolhuis, 2014 includes the following image; notice how the ergaster brain size clusters close to Homo habilis/Homo rudolfensis:

Brain sizes of different hominins. Bolhuis, 2014

Did this Homo ergaster - rudolfiensis - habilis survive in some parts of Africa to evolve into the Homo naledi? A branch that survived in South Africa for over one million years, living as recently as 300 ky ago.

The stone tools

Acheulean hand axes are the typical stone tool industry of Homo erectus, however the ergasters of Africa were not so clear cut. Semaw et al., 2020 wrote a research paper with the suggestive title "Co-occurrence of Acheulian and Oldowan artifacts with Homo erectus cranial fossils from Gona, Afar, Ethiopia" where African "erectus" (i.e. Homo ergaster") made more primitive Oldowan tools. Their comments are shown below. But notice how the mistake genuine erectus that back migrated into Africa, from Busidima, with the smaller, gracile African "ergasters" from Dana Aoule North. They mistakenly attribute the differences to dimorphism:

"We report that both Acheulian and Oldowan artifacts and Homo erectus crania were found in close association at 1.26 million years (Ma) ago at Busidima North (BSN12), and ca. 1.6 to 1.5 Ma ago at Dana Aoule North (DAN5) archaeological sites at Gona, Afar, Ethiopia. The BSN12 partial cranium is robust and large, while the DAN5 cranium is smaller and more gracile, suggesting that H. erectus was probably a sexually dimorphic species. The evidence from Gona shows behavioral diversity and flexibility with a lengthy and concurrent use of both stone technologies by H. erectus, confounding a simple “single species/single technology” view of early Homo"

The smaller ergasters used the more primitive Oldowan tool kit. They were not erectus.

The Asian erectus that migrated back into Africa, evolved there as reported by Asfaw et al., 2002 who studied a 1 million year old skull known as Daka: "A hominid calvaria and postcranial remains recently recovered from the Dakanihylo Member of the Bouri Formation, Middle Awash, Ethiopia... These ∼1.0-million-year (Myr)-old Pleistocene sediments contain abundant early Acheulean stone tools... Here we report that the ‘Daka’ calvaria's metric and morphological attributes centre it firmly within H. erectus. Daka's resemblance to Asian counterparts indicates that the early African and Eurasian fossil hominids represent demes of a widespread palaeospecies. Daka's anatomical intermediacy between earlier and later African fossils provides evidence of evolutionary change. Its temporal and geographic position indicates that African H. erectus was the ancestor of Homo sapiens." So the Daka person is more evolved than the older ergasters, and is the ancestor of modern humans.

Pearson, 2021 noted the differences between ergaster and the Asian erectus; they took a sample of specimens (a hypodigm), and usied Computed Tomography to analyze the brain case, such as the middle cranial fossa (MCF), which hints at the development of the temporal lobe, which is linked to memory, language and social skills and cognition:

"A total sample (n = 51) of cranial Computed Tomography scans (CT) from extant H. sapiens and fossil specimens attributed to African Homo ergaster and Javanese H. erectus were generated into three-dimensional (3D) virtual models with three linear metrics digitally measured on the MCF and two metrics to approximate cranial size...
relative MCF proportions showed extant H. sapiens had a larger MCF to relative to cranial size, while specimens attributed to Javanese H. erectus which had a relatively longer and wider MCF compared to African H. ergaster. These findings indicate that despite sampling limitations, the fossil sample did not show differences in allometric scaling compared to extant H. sapiens, but these specimens attributed to Javanese H. erectus and African H. ergaster share the same total cranial size but had different MCF proportions relative to cranial size. The paleoneurological implications are that these specimens attributed to Javanese H. erectus had longer and wider temporal lobes relative to total brain size compared to African H. ergaster suggesting different patterns of cerebrocranial organization within the Homo erectus hypodigm."

The Asian erectus were therefore smarter than the ergaster lineage.

Regarding the African fossils themselves, Tattersall, 2007 cautiously points out that they are many differences between the specimens, highlight is mine: "However, while this separation at the species level of the African and Indonesian hominids is certainly justified, the species H. ergaster as thus constituted still embraces a bewildering morphological variety. Indeed, although this grouping of African fossils seems to form a fairly coherent clade, it also appears quite diverse."

Conclusion

The ergaster line is more primitive than the Asian erectus, it has smaller molars and premolars, a less specialized skull. Add this to the difference in cognition, tool kits (Oldowan vs. Acheulean) and the timelines, and you can only conclude that they are different species and not a single pan-African-Eurasian group.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall © 

Homo Floresiensis (Flores Island Hobbit) was more primitive than we believed

The Flores Island hominin, known as the hobbit (Homo floresiensis) was originally believed to have mastered fire, but new research published in Science on July 3, 2026 (E. Grace Veatch et al., Taphonomic analysis at Liang Bua reveals the behavioral and technological capabilities of Homo floresiensis. Sci. Adv. 12, eaeb7219(2026). DOI:10.1126/sciadv.aeb7219) suggest that they were much more primitive.

The paper's abstract indicates that "Since its discovery, Homo floresiensis—an extinct, short-statured, and small-brained hominin species from Flores, Indonesia—has often been ascribed unexpectedly advanced behaviors, such as hunting large game and using fire. Here, we report... that Komodo dragons likely had primary access to these remains leaving behind only low-utility elements for H. floresiensis to scavenge. Moreover, no signs of intentional use of fire are present in the stratigraphic units associated with H. floresiensis. Together, these results suggest that H. floresiensis was not as behaviorally advanced as originally suggested and provides critical insights into the behavioral repertoire of H. floresiensis, raising important questions about its ancestry."

The authors point out that the traces of fire usage in the sites where hobbits were found are more recent, and ocurred after these hominins became extinct. Fire was made by modern humans:

"The controlled use of fire and coordinated hunting of large game are considered critical milestones in hominin evolution ... The fact that H. floresiensis was originally described as having these behavioral adaptations continues to be a source of intense debate. However, with no evidence of in situ burning in the Stegodon or murine assemblage sampled thus far from H. floresiensis–bearing sediments at Liang Bua, it is reasonable to conclude that past fire use at the site was the sole result of H. sapiens behavior that occurred from ~46 ka until present day, well after H. floresiensis and Stegodon disappeared from the area."

This isn't surprising, the paper notes that although fire has been used for about 1 million years (My), it was only used widely over the past 400,000 years. Since the H. floresiensis reached Flores earlier than 1 My ago, they probably never mastered the art of fire making.

The researchers report that they were not hunters either, only scavengers. Since they couldn't cook meat, and ate it raw, they did not benefit from a better use of animal protein derived from cooking. The authors attribute this limited "behavioral repertoire as ... possibly due to an ancestry in which large game hunting and controlled use of fire did not evolve." This suggests that they evolved from a primitive, or more archaic branch of hominins and not from Homo erectus.

Comments

As mentioned in a previous post (about Stringer's remarks about the Out of Africa origin of humans), the origin of the hobbits is a mystery. It could be possible that they originated from an early lineage of hominins like those that led to Dmanisi - Homo georgicus, predating the Homo erectus line. They could even be the original basal line that originated in Eurasia, and in Europe, led to the hobbits proposed by Ludovic Slimak as living in France over 400 ky ago (see my previous post about Slimak).

Ancient hominins like these would have developed primitive stone technologies (Stringer calls them "prehuman"), and they may have not learned to use fire. They were probably the first hominins (were they even members of the homo clade?) who left Africa (or probably evene originated in Western Eurasia). They could be the ancestors of the Dmanisi people in the Caucasus (H. georgicus), and the erectines, and migrated into Africa to originate the hominins there.

In my next post we will look into the hominins that are said to have preceded Homo erectus inside Africa, and discuss Homo habilis, H. ergaster, H. rudolfensis.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2026 by Austin Whittall ©