Bushman Canine

The Bushman Canine is an unusual dental feature that was first observed among South African Bushmen (San, Khoisan people) by Oranje in 1934, and Galloway in 1937. The name was coined by David R. Morris in 1975. Its more formal name is "Canine Mesial Ridge", as it is known in the Arizona State University Dental Anthropology System (ASUDAS).

Since it is found at very high frequencies among the San people, and they are presented as the most diverse, divergent, and "oldest" living branch of the original African modern humans, this trait could probably shed some light on the origin and dispersion of our Homo sapiens lineage.

The Bushman Canine

Canine teeth are located on both upper and lower jaws, on each side of the incisors. They have a fang-like appearance, and are the teeth with longest roots. They are designed to seize, tear, and cut food.

Canines with the Bushman canine trait have a different shape, they resemble the premolars. Below is a depiction of a Bushman canine; the image comes from Irish & Morris, 1996:

Prevalence in different populations

This trait is found among the Khoisan, the allegedly "oldest" and most diverse human group, who live in Southern Africa. But it is also found in other populations around the world.

The following image gives the values for African populations, those closest, geographically, from the San Bushmen people. It shows the frequencies of incisor shoveling (see my post on dental shoveling), and Bushman Canine in African populations, from Irish, 1998, Table III.

As you can see, within Africa, there is a large variability in frequency, even in areas close to each other like Western Africa where it varies from 0 in Congo, to 35% in Togo Benin, with 6 or 7% in Gambia and Ghana.

Its frequency, according to Bailey, 2006 and (Sakuma, 1991), in other populations is the following: 0% in early Africans, modern Europeans and modern Asians, Indians, Polynesians, and Native Americans. But it reached 20% in Homo erectus, 33.3% in Early Neanderthals, 57.1% in late Neanderthals, and 16.6% in Upper Paleolithic Europeans.

This makes me wonder if its presence in ancient Europeans stems from Neanderthal or Denisovan introgression, or possibly a trait that is ancestral, and inherited from erectus.

In other populations, Irish, 1998 found the following frequencies for Bushman Canine: Sub-Sahan Africa: 18.1% ; North Africa: 6.1%; Europe: 4.8%; Sundadonts: 2%; Sinodonts: 1.2%; Australia: 5%; Melanesia: 3.2%.

Irish, 2026 reports it at 0% frequencies in Pima (Native Americans), Lower Ob Khanty (West Siberia), Aleutians, Chuckchi (Siberia), Vietnam, Philippines, Australia (North), Nepal, Italy, and South Finland; and at low frequencies elsewhere: Kazakhistan 2.4%, Monglolia 3.7%, Thailand 9.3%, Japan 4.7%, Malaysia 4.3%, Borneo 11.1%, New Britain (Melanesia) 1.9% Greece 8.7% Estonia 2.7% Lapps (Finland) 4.2%. (See Table S2 in that paper's supplementary material).

Hotspots with high frequencies Outside of Africa

The "Non-African" values mentioned above are all lower than 10% (except for Borneo). Interestingly, it has been reported at rates of 14% among inhabitants in Kerala and Odisha, in India (Nair, 2020). Very high frequencies were reported among Central American natives by Scherer, 2004, who studied Maya dental traits from the Classic Period (AD 250 to AD 900) and found a 7.1% frequency, which is high, some sub-populations like those at Barton Ramie, Aguateca and Seibal presented prevalences of 25% to 36.4%. These values are also mentioned by Domínguez Vázquez, 2020 in his thesis (see Annex 4, p.162). He reports it at 57.1% frequency among Yucatan pre-Hispanics, 100% among Yucatan pre-ceramic people (by pre-ceramic he means 4,500 to 12,000 year-old specimens) yet absent among Mexican and North American pre-ceramic populations, and 100% in Zhoukoudian, China humans, 35,000 years old.

The Sub Saharan San (Khoisan) Bushmen

But, why is it found among San people at frequencies of 43.1%? (Sakuma, 1991) There has been no Neanderthal introgression into Africa. Did erectus migrate into Africa, carrying it? Or, is it an ancestral trait that has not been lost over the course of hundreds of thousands of years?

These people have some odd ancestral traits, including the epicanthic fold, shared with Asians, pale skin pigmentation, shared with Europeans, they also display moderate dental shoveling, a factor which Nair, 2020, found as strongly associated with Bushman Canine among Indian populations, which hints at some kind of genetic link between both dental traits.

Back in 2018, I posted about research describing teeth recovered from a human who lived in Magubike, Tanzania, 45,000 years ago. The remarks were that they had Neanderthal and H. erectus traits. I revisited the post today, and reread the article by Willoughby et al., 2018 and noticed that its Table 11 reported that this person had Bushman Canine trait, and added that it was not present in other Mid to Late Pleistocene African fossils yet present in 20% of the Early Holocene Kenyan ones.

The paper also reported that the teeth had “Khoisanoid affinities” (due to the presence of the Medial Canine Ridge found in the San people, among other traits), yet these teeth were big, and had some traits that are not found in archaic Homo sapiens or H. heidelbergensis or other hominins from that period.

This seems to imply the presence of other unknown hominins in Africa admixing with the Khoisan people, super archaics? These could be related to the Homo erectus lineage, or relatives of Neanderthals / Denisovans. This could explain the high frequencies in India and Borneo, possible spots of Denisovan-human admixing. And, regarding America, I have already posted about human-Denisovan admixure in America.

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

Shoveled incisors an ancient hominin trait

Teeth shapes are a useful tool to identify the fossil remains of hominins. Neanderthals, Homo erectus, and modern humans have distinct teeth shapes (mass, multiple roots, shoveling, cusps, protrusions). Even people living in different continents have specific teeth shapes.

Dental Shoveling

One type of tooth shape is known as dental shoveling, where the front teet, known as incisors, instead of being flat on their inner side, are shoveled. This type of shoveled teeth has been observed in monkeys such as macaques, baboons, and gibbons. It is also found in contemporary and ancient humans, as well as hominins.

Hominins with shoveled incisors

Our hominin ancestors also displayed shoveled incisors: it has been found in ancient remains of Australopithecus, Homo erectus, and the fossils from Dmanisi, Georgia and Atapuerca in Spain (Homo georgicus and Homo antecessor). More recent, yet ancient specimens include Neanderthals ad of course, remains of Homo sapiens (Denton, 2011; Margvelashvili, 2008).

Shoveled incisors in contemporary populations

Among our lineage, Homo sapiens, modern humans, shoveling is found at vry higher frequencies in East Asians (Japanese, Chinese, Mongolians, Northeastern Siberians) and populations that originated in Northeast Asia like the Eskimos and North American Native Americans, and the South American natives who replaced the older, ancestral Paleo Indians in that part of the Americas. Surprisingly, the Khoisan in Southern Africa have shoveled teeth.

Other ethnic groups have very low frequencies or absence of shoveled teeth: the paleo-Indians in South America, South and South Eastern Asians, Melanesians, Europeans, Arabian and Middle East populations, North Africans, central Africans, Eastern Africans.

Below is an image showing a normal, and a shoveled tooth (from Benton), it cleary shows the difference in incissor shape due to shoveling.

A table with frequency of shoveling, and source

• 100% (Source) Mongolia
• 100% (Source) China
• 64.7% (Source) Ainu, Sakhalin
• 93.4 % (Source) Japan, Tokyo
• 23.8 % (Source) Tajik, Afganistan
• 6 % (Source) Philippines
• 87.3 % (Source) Polynesian, Easter Island
• 60-90 % (Source) Australian Aboriginal
• 73% (Source) Melanesian
• 98.3% (Source) Pima Native Americans
• 100 % (Source) Yanomana Native Americans, Amazonia Brazil, Venezuela
• 100% (Source) Northwest Argentina (Amerind, mestizo)
• 90.1 % (Source) Precolumbian Peru
• 49.5 % (Source) Lapps in Finland
• 9.4% (Source) England
• 20.1% (Source) Denmark
• 40.2 % (Source) Germany
• 10.3 % (Source) Uganda
• 24 % (Source) Egypt
• 20 (Source) Sudan
• 31.2 % (Source) Punjab, Pakistan
• 22 % (Source) Odisha, East Central India
• 15 % (Source) Kerala, Southern India
• 95.5 % (Source) Mysuru, Southern India
• 8 % (Source) Tamil population in Southern India
• 9 % (Source) Pakistan
• 5 % (Source) Syria and Jordania
• 20 (Source) Yemen
• 9 % (Source) Saudi Arabia
• 10.1 % (Source) Turkey Black Sea region

The variability of frequencies is notable, even within the same region (see India for instance). Amerindians and NE Asians have extremely high prevalence of shoveling. There is a cline from East to West across Eurasia, but even in SE Asia and Australia, frequencies are high.

Not all shoveling is the same

Shoveling in Neanderthals, erectus and modern humans differ in shape, and is probably driven by different genes. In this sense Yuan, 2019 notes that "The type of shoveling teeth found in Neanderthals and Pleistocene Homo from Atapuerca-Sima de los Huesos may either be a different type from that of Asians and Africans or come from early dispersal of Homo from Asia to Europe."

But which genes are involved, or if there is an interplay of several, is a mystery.

Kimura, 2009 reported that the Ectodysplasin receptor Gene (EDAR) in its "EDAR 1540C alleles... was strongly correlated with the tooth-shoveling grade... The effect of the allele was additive and explained 18.9% of the total variance in the shoveling grade." But it seems that other alleles are involved in shoveling. Interestingly EDAR 1540C is absent in Africans, Denisovans, and Europeans (where shoveling is low) and high in East Asians and modern Amerindians.

Who has shoveled teeth? Frequencies

Bailey, 2006 provides some frequencies on incisor shoveling in different populations: 100% in Homo erectus, early Neanderthals, and late Neanderthals. At 33.3% in Early Africans and Modern Asians, 45.5% in Upper Paleolithic Humans in Europe, and 37.5% in modern Europeans.

Denisovans, in the only incisor found to date, from the Xujiayao specimen of North China, 220,000 to 160,000 years old shows "marked shovelling." (Xing, 2015 in Masojc, 2026).

Interestingly, Yuan, 2019 notes that "native Africans such as Khoisans are well known to have certain East Asian features such as shoveling teeth, epicanthic fold, and lighter skins." The Khoi are said to be the most diverse and divergent, and therefore the most ancestral lineage of modern humans on the planet. But no Neanderthal or Denisovans have been found in Africa, and admxiture there between the ancestors of the San and the Neanderthals and Denisovans is not believed to have happened.

Comments

There are several genes that regulate teeth shape (Qing, 2025), making it a "polygenic" trait.

Shoveling has been linked to one variant of the EDAR allele (I have posted about it and its presence in Asia and the Americas).

Research by Kaya and Merriwether, 2024 has shown that Shovel Shaped Incisors (SSI) and Carabelli's Cusp (CC) are both linked to other genes: "Fifteen SNPs/s belonging to SOSTDC1, FGF3, FGF4 and WNT10A were significantly associated with SSI and/CC expression." So there are several genetic drivers for shoveled teeth.

Regarding the EDAR variant in Amerindians and Asians; did it arise spontaneously as a mutation in modern humans in Asia and then spread in Northern and Eastern Asia, and the Americas?, or was it introgressed from other hominins living in that region, like Homo erectus or the Denisovans?

Also, why do the San people of South Africa have shoveled teeth, even though there is no known introgression of Neanderthals, Denisovans or erectus in that region? Is it an ancestral trait found in archaic hominins which admixed from ghost African populations? The lower frequency in Europe and Western Asia, as well as North Africa suggests that there were hominin populations which lacked the shovel teeth alleles. This suggests, in my opinion, that it is possibly a modern human trait, and that shovel teeth introgressed from other hominins in Central or Eastern Asia (Neanderthals, Denisovans, or erectus), and back migrated from there into Europe, Western Eurasia and Africa.

A very interesting and thorough article published in 1985 by Yuji Mizoguchi "Shovelling: A statistical analysis of its morphology" suggests that shoveled teeth are stronger than regular non-shoveled ones and that the hunter-gatherer lifestyle selected in favor of strong faces able to "resist powerful biting forces, and one of the causes of the need for such a powerful biting mechanism was possibly hunting activity directly or indirectly associated with heavy meat-eating."

Misoguchi distinguishes three kinds of shoveling:

(1) weak or moderate, with "a slightly swollen basal part of the lingual surface of the tooth crown and lingual marginal ridges converging towards the cervix" Found in the ancient australopithecus and in almost all contemporary humans. This one continued to exist in Africa since the Australopithecines, and is found in modern Africans. Found in modern Asians and Europeans too.

(2) strong, with highly develped parallel ridges on the teeth's margins "a very deep lingual fossa and a large lingual tubercle." Found among East Asian Neanderthals and Homo erectus. Western Eurasian Neanderthals had a blend of types (1) and (2).

(3) strongly marked shoveling but the marginal ridges converge on the cervix. Lacking a "developed lingual tubercle" The tubercle is different from that of erectus and Neanderthals. Found among Innuit, Native Americans.

With these three varieties in mind, the evolution and dispersal of shoveling is the following, according to Mizoguchi>

"In Africa, the first type of incisor has continued since the emergence of the hominid, In Europe, the first type changed to the second type before or at the stage of Neanderthals, and, later, back to the first type. In Asia, as in Europe, the second type once changed into the first type after the Neanderthal stage, but some Asian anterior teeth developed in a different way from before, presumably as a result of direct or indirect adaptation to a severely cold environment. Such incisors are the third type of incisor observable in the modern inhabitants of the Arctic and Americas."

Perhaps the sequence is (1) ancestral African, evolved into a derived one, (2), in Homo erectus in Eurasia and later retained in Neanderthal. Those who would later become modern humans retained the ancestral version, globally, and the (3) variant appeared in East Asia, possibly from admixture with Denisovans.

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

Another post on dental enamel protein AMBN 273V or M in Denisovans, Neanderthals, and Humans

In three different posts I mentioned dental enamel protein as a tool to investigate ancient ancestry, admixtures, and archaic introgressions.

The first post looked into a paper by Quiamei Fu, et al., 2026, Enamel proteins from six Homo erectus specimens across China, published in Nature on May 13, 2026 (vol. 655, pp 141–147 https://doi.org/10.1038/s41586-026-10478-8), it suggests archaic introgression from Homo erectus into Denisovans, and from them, into modern humans, which shared a protein found in teeth enamel, named AMBN(M273V) from erectus via Denisovans to us, Homo sapiens.

The second post mentioned another protein found in teeth, called COL17A1 that is found in its ancestral form in Paranthropus robustus and Homo naledi, and in derived form in--Denisovans, Neanderthals, and modern humans. Suggesting that naledi may not be a homo (see Madupe P, Taurozzi A, Koenig C., et al., Proteomic analysis of dental enamel from 20 Homo naledi individuals shows no male markers. Cell, 2026; 189, 4415-4424.e11).

The third post mentioned another recent paper that questions the archaic introgression hypothesis proposed by Fu et al. and suggesting other alternatives for the presence of AMBN 237V in Denisovans and humans, that don't require an introgression with Homo erectus (see No signal of a super-archaic origin of the Denisovan AMBN gene, a comment on the protein affinity of Homo erectus and Denisovan enamel proteins, Ioannis Patramanis, Frido Welker, Laurits Skov. bioRxiv 2026.08.11.744165; doi: https://doi.org/10.64898/2026.08.11.744165).

Today I will discuss a fourth paper on the subject of dental enamel proteins, published last week, on August 26, 2026 by Paula Kotli; Enamel Palaeoproteomics Successfully Distinguishes between Denisovans and European Neandertals (bioRxiv 2026.08.22.746388; doi: https://doi.org/10.64898/2026.08.22.746388).

Kotli compared DNA from the Penghu jawbone, dredged up from the sea in Taiwan, and a Neanderthal from Portugal and found that the AMBN V273 variant (with a Valine aminoacid substitution there —hence the "V") was exclusively Denisovan while the Neanderthal carried a Methionone aminoacid there (M273 variant). These two proteins can serve as markers to identify the origin of ancient teeth when DNA is not correctly preserved; tooth peptides last longer than DNA.

Most modern humans carry the M273 variant, shared with Neanderthals, while not only Penghu, but other Denisvoans (like the Denisova 3 specimen) carry the V273 variant. Unsurprisingly, the region with the highest levels of Denisovan admixture, the Philippines, also has the highest level among all modern humans of the V273 variant: It reaches "a notably elevated frequency of ∼21% in modern Philippine populations (compared to <1% globally)."

Africans and archaic admixture

While researching about ancient teeth proteins and their prevalence among modern humans, Neanderthals, and Denisovans, I came across a good paper on the subject: Zanolli C, Hourset M, Esclassan R, Mollereau C. Neanderthal and Denisova tooth protein variants in present-day humans. PLoS One. 2017 Sep 13;12(9):e0183802. doi: 10.1371/journal.pone.0183802. PMID: 28902892; PMCID: PMC5597096. In it, they mention the presence of rare variants found in Denisovans and Neanderthals, inside Africa. This is quite unexpected because both of those ancestral lineages are not known to have existed within Africa and the current view is that they admixed with the humans that left Africa in the Out of Africa migration. So, how did these rare variants find their way into the genes of Africans?

Zanolli et al. write (highlight is mine): "The finding in living Africans of rare derived polymorphisms that were present in Neanderthals (Fig 2: ENAM G389S, 5%) or Denisova (Fig 3: CEMP1 R80H, 0.5%) is difficult to reconcile with the fact that interbreeding of AMH with Neanderthals occurred out of Africa. However, as mentioned above, this could be explained by the inside Africa admixture between the ancestors of AMH and archaic hominins, or by the possible back migration of Neanderthal gene flux. Further genome analysis of the Neanderthal and Denisova polymorphisms in AMBN, ENAM, and CEMP1 certainly deserves attention as they could trace back an African ancestor common to Neanderthals, Denisova and early anatomically modern humans" The final comment is expected: these rare alleles could be the faint trace of the last common ancestor shared by Humans, Neanderthals, and Denisovans. I choose to believe that they are the result of admixing between archaics and recent Africans within Africa.

The ENAM G389S variant is, according to the supplementary material found at rates of zero in Europe, South Asia, and East Asia, at 5% in Africans, and at 0.3% in America. So how did it exist in Neanderthals but leave no trace elsewhere, but in Africa and America (Maybe slave trade brought it to America?). The other variant, the Denisovan CEMP1 R80H is found at zero levels everywhere except Africa where it reaches 0.5%. Denisovans in Africa?

K 55 E variant

Another ancient (ape variant) is found at surprisingly high levels in Africa, with 26%, but only at 2% in Americas, 0.2% in Europe, zero in East Asia, and 0.1% in South Asia. This variant is known as K55E (rs13331643) in CEMP1, and is commonly shared in this ancestral form with gorillas and chimpanzees.

So, basically, all modern humans in the rest of the world carry the derived allele of K55. Surprisingly, the ancestral variant, found in Africa at 26% frequency, is also found in Denisovans and Neanderthals. Again, a contradiction to the belief that Africans never mingled with those hominins.

The paper tries to explain this incongruity as follows (note that AMH means "anatomically modern humans"): "This suggests that the ancestral variant may have not been favorable to AMH outside Africa or, conversely, that the derived allele was specific to AMH and evolved out of Africa due to a founder effect. The ancestral archaic variant could also explain in part the tendency of people with African origins to have larger crowns and root volume and surface. Indeed, this variant is also found in Neanderthals teeth, characterized by high crown dimensions and frequent taurodontism. The presence of this variant in Denisova is also compatible with such traits."

Why would the non-Africans, who mated with Neanderthals and Denisovans not uptake their ancestral K55 variant in higher frequencies, and instead retain a mutation (the derived form), while Africans, who supposedly never met the Neanderthals & Denisovans, carry the ancestral variant?

I don't see how this fits in with the Out of Africa theory. The admixing of Denisovans in Asia can explain its presence there, and also in America, and Europe where Neanderthal admixture is higher, but why is it so prevalent in Africa? Maybe, the cause is an introgression of super-archics into the recent African populations.

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

Denisovans may not have admixed with erectus after all

A preprint published on August 11th, 2026 proposes an alternate explanation to the teeth protein shared by Denisovans and Homo erectus, suggesting different scenarios that could result in the same outcome and not require admixture or introgression from erectus into Denisovans.

This article was published in the non-peer-reviewed bioRxiv: No signal of a super-archaic origin of the Denisovan AMBN gene, a comment on the protein affinity of Homo erectus and Denisovan enamel proteins, Ioannis Patramanis, Frido Welker, Laurits Skov. bioRxiv 2026.08.11.744165; doi: https://doi.org/10.64898/2026.08.11.744165 and disputes the claims put forward by Quiamei Fu, et al., 2026, Enamel proteins from six Homo erectus specimens across China, Nature, https://doi.org/10.1038/s41586-026-10478-8

Background

Last May, I posted about the original paper by Fu et al., that reported that a variant of a tooth enamel protein (AMBN) found in 400,000 year-old Homo erectus specimens, known as (M273V) was also found in Denisovans, suggesting a "super-archaic introgression, some of which later passed to modern humans, are likely to have originated from H. erectus. Late Middle Pleistocene H. erectus may have coexisted with Denisovans in parts of East Asia, where these interactions are presumed to have occurred." However, this new research suggests other alternatives.

The image below is from the original paper (Fig. 4) and it shows "a possible model of gene flow related to AMBN(M273V) among H. erectus associated with the populations of Zhoukoudian, Hexian and Sunjiadong, Denisovans and modern humans." The red hexagon represents AMBN(M273V), which passes (right to left) from erectus into Denisovans, and through them, into modern humans.

The image also shows a new allele found only in erectus (yellow pentagon), called AMBN(A253G) absent in all other hominins (Denisovans, Neanderthals, Dmanisi, Homo antecessor, and modern humans).

Now those findings are disputed

The new paper by Patramanis et al., says that the erectus admixutre explanation of Fu et al., is based on an assumption: that the AMBN (M273V) variant found in Denisovans has a super-archaic origin (via erectus). The authors suggest that it isn't super-archaic. Instead, they say, there are four different possible explanations for the presence of that variant in both Denisovans and modern humans.

Below is Fig. 1 from Patramanis, 2026, with the alternatives B, C, D, and E, that could explain the presence of the allele. Alternative (A) is the one shown further up, in Fu et al.

These are the four possible alternatives:

• (B). In this case, the variant appears, by chance in Denisovans, who in turn pass it on to H. erectus and modern humans. In this case, the allele is relatively recent, and not super archaic. The erectus also carry their specific variant, AMBN(A253G).
• (C). The allele is Denisovan. That have admixed with erectus and received AMBN(A253G). This group is mistaken by Fu et al. to be erectus. They are actually admixed Denisovans.
• (D). This model is one where Homo erectus did not admix at all with Denisovans. The Denisovans are a structured population, and some carry the AMBN(A253G), others carry the AMBN (M273V), some carry the two or none.
• (E). This alternative excludes admixtures. The variant was present in the common ancestors of Denisovans, Neanderthals, erectus), but through incomplete lineage sorting (ILS), by chance this variant was only inherited by erectus and Denisovans, but not by Neanderthals or Homo sapiens. Later admixing passed it from Denisovans into modern humans.

I favor the original theory. Denisovans lived in East Asia, and had plenty of time to admix with erectus there. Let's wait for Fu et al.'s reply to this paper, and see what future research finds.

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

Neanderthals treated a tooth cavity 59,000 years ago

Although this post has nothing to do with the main theme of my blog, I decided to include it because it gives us a glimpse of the intelligence and humanity of Neanderthals. They were not lumbering semi-human brutes. They were human beings, just like us, with a brain and a mind similar to ours. In this post I will comment about a very recent publication about how a Neanderthal from the Chagyrskaya site in the Altai region of Siberia 59,000 years ago, perforated a molar to relieve the pain caused by a caries. The oldest example of dentistry on record.

The paper is the following: Zubova AV, Zotkina LV, Olsen JW, Kulkov AM, Moiseyev VG, Malyutina AA, et al. (2026) Earliest evidence for invasive mitigation of dental caries by Neanderthals. PLoS One 21(5): e0347662. https://doi.org/10.1371/journal.pone.0347662

Over the years, there have been studies (the links lead to the research that backs these statements) that suggest that the Neanderthal buried their dead, and took care of their elders and injured companions, disabled (Down syndrome) and, mended those that were sick. This implies some kind of elevated mental function, and compassion.

In this paper the authors analyzed a molar (one located on the lower left side of the mouth) which had a large man-made hole bored into it (see image) this hole was made while the individual was alive, and the person survived after it was done. It was perforated by using a stone "perforator" that drilled or rotated against the enamel of the tooth to reach the inner pulp and relieve the pain caused by the caries with "severe pulp exposure." This procedure was done without any type of anesthesia, and would have taken (based on the experiments conducted by the researchers) between 10 minutes and two and a half hours, but they point out that "the individual would have had little choice: remove the dentin or face potentially fatal consequences."

Fig 2. Chagyrskaya 64 molar tooth and its macro-features. 1 General view of the tooth in five projections; a–c. Macro-photographs of the crown’s occlusal surface features: a. superior view of the concavity; b. stepped groove on the concavity’s wall; .
From Zubova et al., 2026

The authors point out that the procedure required "... two distinct types of manipulations requiring different tools, in addition to the drilling/rotating technique, necessitating complex finger movements, indicates that the Chagyrskaya Cave Neanderthals possessed the cognitive capacity to intuit the source of pain, comprehend the feasibility of its elimination, and deliberately select the most efficacious dental intervention. These patterns bring Neanderthal behavior closer to modern humans and differentiate that behavior from the instinctive actions of other primates."

Interestingly, the enamel thickness on Neanderthal teeth is thinner than in human teeth, meaning that it is easier to perforate it and reach the pulp than in modern humans. The edges and walls of the perforation are smooth and rounded suggesting that the individual was alive during the procedure and that lived after it, using the tooth. The authors wondered if the hole had been filled with wax, resin, bitumen, tree sap or antiseptic herbs to serve as painkillers but a Raman spectroscopical test did not reveal the presence of other materials.

Yes, it must have been a painful intervention, and the aftermath was also painful as it resulted in " complete pulp exposure. On the one hand, this could have caused pain for some time; on the other hand, pulp exposure would have accelerated nerve destruction, after which the pain should have disappeared." Was it done by chance? or did these people know how to treat a caries?

The paper concludes that this molar "offers new insights into Neanderthal self-regulation, revealing their capacity for deliberate, sustained actions that required enduring pain – a behavior not documented in other higher primates, which typically rely on instinctive responses. While genetic evidence confirms that Neanderthals exhibited heightened pain sensitivity compared to Homo sapiens, their ability to engage in goal-directed behaviors involving pain tolerance suggests advanced volitional control. It can be argued, therefore, that Neanderthals exhibited volitional strategic goal-setting behaviors that may have aligned more closely with those of Homo sapiens than with non-human primates... challenging traditional distinctions in cognitive archaeology between anatomically archaic and modern members of Homo sapiens and highlighting their shared legacy of biological and cultural adaptations."

Closing Comments

Neanderthal were smart, compassionate, and human. By the way, the earliest Homo sapiens dental intervention discovered to date was in an individual from 14,160 to 13,820 years ago, who lived in Ripari Villabruna in Italy. This person shows evidence of enamel scraping on a caries more of a toothpicking procedure rather than a drilling one.

One can hardly imagine the pain suffered by countless generations of human beings before the discovery of analgesics and anesthesia. We are fortunate to live in the present time! I recall the scene in the movie "Castaway" (2000) where Tom Hanks, stranded on a desert island, in terrible pain due to an untreated toothache that he had prior to his trip that led him to the island, does away with the tooth using a rock as a hammer and the blade of an ice skate as a chisel, passing out after the procedure.

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

An earlier Human - Neanderthal split? (400,000 years earlier!)

Aida Gómez-Robles published a paper in Science Advances three days ago (May 15, 2019 - Dental evolutionary rates and its implications for the Neanderthal–modern human divergence, Vol. 5, no. 5, eaaw1268 DOI: 10.1126/sciadv.aaw1268) in which she finds evidence that "support[s] a pre–800 ka last common ancestor for Neanderthals and modern humans unless hitherto unexplained mechanisms sped up dental evolution in early Neanderthals".

This is really remarkable, as it pushes the conventionally accepted date of divergence back in time from 400 to 800 thousand years ago.

This far older age was worked out by studying ancient Neanderthal teeth from the Sima de los Huesos site in Spain.

Gomez-Robles compared fossilised teeth from archaic hominins such as Australopithecus afarensis, Australopithecus africanus, Paranthropus robustus, Paranthropus boisei, and ancient ones such as Homo habilis, Asian specimens of H. erectus, a Sima de los Huesos H. neanderthalensis and of course, us: Homo sapiens, Other Neanderthal teeth from different sites in Europe were deliberatel left out of the comparative study.

The idea behind the comparison was to see how the shape of teeth in hominins changed over time. Gomez-Robles found that they evolve, and do so at a steady pace. This let estimate when each of these species split from each other.

The conclusion: "The simplest explanation of the results presented in this study is that Neanderthals and modern humans diverged before 0.8 Ma ago", our Last Common Ancestor (LCA) with Neanderthals dates back to 800,000 years ago.

Gomez-Robles goes on to spell out the implications of this more ancient split:

" If the phenotypic LCA of Neanderthals and modern humans was older than 800 ka, this would imply that all fossil hominins younger than this age are no longer valid candidates to occupy this ancestral position. Some fossils younger than this age, however, are frequently considered to be part of the last common ancestral species to Neanderthals and modern humans. These fossils, usually ascribed to Homo heidelbergensis, include European and African specimens, such as Mauer, Arago, Petralona, Bodo, Kabwe, etc., and maybe even some Asian specimens. If Neanderthals and modern humans diverged earlier than 800 ka ago, then all these fossils have to be related either to Neanderthals or to modern humans, or they can be part of a sister lineage to both of them. These fossils, however, cannot be ancestral to Neanderthals and modern humans because they would postdate their evolutionary divergence. An evolutionary relationship between these fossils and both Neanderthals and modern humans would be possible only if they were part of an older ancestral species that persisted in time as a relic species after the actual split of both lineages. Effectively, this scenario would mean that the H. heidelbergensis fossils are part of a sister group to Neanderthals and modern humans but that the evolutionary change from their putative ancestral populations did not involve speciation."

Of course, there are confounding factors that can explain the structural differences in teeth without requiring such an older age, for instance: the teeth evolved quicker in a small population of Neanderthals, isolated from other human groups, making them seem older while they are actually only 400 Ky old.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2019 by Austin Whittall © 

Some Odd teeth in China

A paper (Late Middle Pleistocene hominin teeth from Tongzi, southern China, by Song Xinga, María Martinon-Torres and Jose MariaBermudez de Castro) reported that some teeth roughly 200,000-years-old found in the Yanhui cave in China, which were originally thought to belong to Homo erectus, may belong to some other hominin.

They have a mix of modern and archaic traits, which lack some characteristic features of H. erectus, but don't belong to Neanderthals either!

Since modern H. sapiens was still in Africa at that time, these teeth (because they are large) may have belonged to the mysterious Denisovan group.

Expect some surprising findings to come out of China in the coming years. Asia was teeming with "humans" long before the purported "out of Africa" migration took place. It may be the cradle of humankind.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2019 by Austin Whittall © 

Atapuerca Sima de los Huesos remains were closer to the Neanderthals

Four and a half years ago I posted about a site in Spain, known as Atapuerca (What Science is all about), in it, I mentioned two studies that came to different conclusions: one stated that the human remains there resembled Neanderthals, the other that they were closer to the Denisovans than to the Neanderthals.

Today I read a paper that was published on Nov. 30, 2018 and which states that the teeth of remains from Atapuerca's Sima de los Huesos site shows that they had a close relationship to the later Neanderthal groups found in Europe.

The paywall protected abstract says:

"Enamel and dentin patterns have awakened a considerable interest in phylogenetic studies. However, almost nothing is known about the dental tissue proportions of European Pleistocene hominins, apart from Neanderthal populations. This study aims to assess the three-dimensional dental tissue proportions of permanent canines belonging to the extensive sample of hominin teeth at Sierra de Atapuerca (Spain) through the use of microtomographic techniques. Our results show that early and middle Pleistocene populations from Atapuerca exhibit large coronal and root dentine dimensions, as well as a thinly enamelled pattern, which has been traditionally considered an autapomorphic Neanderthal trait. Therefore, these results might support an early enamel thickness decrease which is already observed 800 kyr ago in Homo antecessor and maintained in later groups such as Sima de los Huesos and Neanderthal populations during the middle Pleistocene."

This is important because it shows that the Sima de los Huesos remains had a close relationship with the Homo neanderthalensis.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2018 by Austin Whittall © 

45 KY old African teeth from Magubike with Neanderthal and H. erectus traits...

This paper Middle Stone Age human teeth from Magubike rockshelter, Iringa Region, Tanzania, by Pamela R. Willoughby, Tim Compton, Silvia M. Bello, Pastory M. Bushozi, Anne R. Skinner and Chris B. Stringer. Published: July 31, 2018 https://doi.org/10.1371/journal.pone.0200530, discusses human teeth found in Africa:

They are significant because they come from Tanzania (the Magubike rockshelter) and are apparently fom Middle Stone Age (MSA) deposits dated and a "conservative estimate of their minimum age is 45,000 years" according to the paper.

As we all know, finding human remains in this part of Africa, especially those from the Middle Stone Age (which took place between 320,000 and 30,000 years ago) is a very rare event.

The San ("oldest" humans) and this paper

The authors try to find similarities between these ancient teeth and the San people (the San are always mentioned because they are assumed to be a "relict" of the original Homo sapiens that left Africa, the "oldest" extant humans, the "most diversified" members of our species.

And they say so, albeit cautiously, in this paper: "... a number of characteristics in the Magubike teeth suggest possible San affinities.".

But as we go into the details, it seems that these teeth are indeed quite unique, and, in my opinion, not at all like those of the San people:

Quoting the authors: (highlight is mine)

"In summary, the Magubike teeth have the characteristics typical of African sub-Saharan Homo sapiens Holocene teeth, but aspects of the morphology of the upper canine and third premolar are atypical for the African mid/late Pleistocene sample. However, this may be at least partly due to its North African bias. Certain traits present ― e.g. the root shape of the lateral incisor and canine, and the presence of a cingulum pit on the lateral incisor ― are rare or absent in the comparative samples. Traits specific to Homo heidelbergensis / archaic Homo sapiens and Aterian teeth are not present. Despite their large size, there is some suggestion of San affinities, though this interpretation must be very tentative, being based on a single individual."

The interesting part is the similarity with Neanderthal and Homo erectus features: (again highlight is mine)

"... The pronounced buccal vertical curvature of the canine root found at Magubike is seen in early teeth, Homo erectus and Neanderthal, but not in samples of early modern Homo sapiens (Skhūl and Qafzeh) or European Upper Palaeolithic Homo sapiens.
In recent teeth, the buccal surface is usually straight or has a mild curvature. The few canine roots that could be observed in the Kenyan early Holocene sample, and the two in the Gwisho sample, are straight, and in the mid/late Pleistocene sample only the Broken Hill canine has similar curvature to that of the Magubike canine."

In other words it is not found among early modern Homo sapiens, only in Neanderthals and H. erectus. The authors continue, finding some examples of curvature in recent African samples (7% of them - a low figure) which may have come from out of African migrants in more recent times...:

"However, four instances were found in the recent sub-Saharan East/South African sample (7%). Similarly, the incomplete root of the Magubike lateral incisor has pronounced buccal vertical curvature in contrast to the more evenly tapered form of recent teeth. A lateral incisor of this form occurs at the Klasies River Main site and in one of four in the Gwisho sample, but in just one out of seventy-five in the recent East/South African sample."

As we see in the preceden paragraph, there are yet more morphological traits that are not at all related to modern African samples.

And the authors admit the teeth's unusual uniqueness very clearly:

"The six Magubike teeth are differentiated from African archaic Homo sapiens / Homo heidelbergensis and Aterian teeth in terms of size and the lack of morphological traits that could be associated with these groups. They are also differentiated from the Kenyan early Holocene and Gwisho samples and more recent teeth in terms of the relative size of the incisors, the incisor crown heights, the relative crown heights of the two premolars, and the long premolar roots. However, the morphological evidence is contradictory, principally in relation to the canine having traits that are rare or absent in the African mid/late Pleistocene sample, though this may be partly due to the small size of this sample and its North African bias."

They attribute this to the small sample size and its "North African bias."

But why try to find them similar to San teeth? Well, orthodoxy requires it: oldest African remains of a Homo sapiens and "oldest" extant humans (SAN) "must" be related:

"Small bodied individuals recovered from East African LSA sites have often been described as having “Khoisanoid affinities”, implying an assumed connection with the small bodied people of South Africa, including the San. There are also still people in northern Tanzania today whose languages are often related to Khoisan (the Hadzabe and Sandawe), implying some possible past connections with southern African peoples."

But what if... the teeth resemble Homo erectus and Neanderthal teeth because these migrated INTO AFRICA? As we can see they don't resemble the African teeth of that time or those of early Modern Homo Sapiens...

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2018 by Austin Whittall © 

On the genes that regulate teeth, archaic hominins and Africans

A paragraph in this paper: Neanderthal and Denisova tooth protein variants in present-day humans) by Clément Zanolli , Mathilde Hourset, Rémi Esclassan, Catherine Mollereau (Published: September 13, 2017https://doi.org/10.1371/journal.pone.0183802) caught my eye. It pointed out an anomaly: Africans sharing a trait with Neanderthals and Denisovans!

This is what it says:

"The finding in living Africans of rare derived polymorphisms that were present in Neanderthals (Fig 2: ENAM G389S, 5%) or Denisova (Fig 3: CEMP1 R80H, 0.5%) is difficult to reconcile with the fact that interbreeding of AMH with Neanderthals occurred out of Africa [75, 76]. However, as mentioned above, this could be explained by the inside Africa admixture between the ancestors of AMH and archaic hominins, or by the possible back migration of Neanderthal gene flux. Further genome analysis of the Neanderthal and Denisova polymorphisms in AMBN, ENAM, and CEMP1 certainly deserves attention as they could trace back an African ancestor common to Neanderthals, Denisova and early anatomically modern humans."

Well, it is not "difficult to reconcile" if the fact is wrong and AMH interbred with Neanderthals inside Africa...

The ENAM G389S variant is found not only in Africa at 5% but also at very low frequencies in the Americas 0.3% (maybe imported by African slaves or perhaps a genuine Amerindian-Neanderthal admixture). Its global prevalence is only 1.3% and is virtually absent in Asia and Europe. Denisovans carried the ancestral "G" type, found in chimps and gorillas. Neanderthals and modern humans carry the derived "S" type.

They also found a some proteins in Denisovans (K55 E variant) that appear at 26% frequency among Africans and 7% globally (the highest outside of Africa being in America at 2% and far lower elsewhere). This is the ancestral type, found in Denisovans, chimpanzees and modern humans, but it was not found in Neanderthals.

The paper also hints at an admixture in Africa between H. Sapiens and some archaic homo 35,000 years ago:

"Most of the amino acid changes are found in the dentin and enamel proteins that are encoded by genes located in two close clusters on the chromosome 4 (Fig 4) and supposed to derive from an ancestral gene encoding a secretory calcium-binding protein [68]. These clusters fall apart from known haplotypes on chromosome 4 and do not correspond to candidate Neanderthal or Denisovan gene flow regions in non-African living humans [69, 70]. However, a signal of archaic introgression in the locus 4qMB179 on chromosome 4 has been evidenced in Central African populations, and is supposed to result from admixture with an archaic Homo some 35 kya [71]."

Admixture with archaic hominins in Africa would "enrich" the diversity of African DNA and therefore contradict the prevailing theory that Africans are more diverse and therefore they are the "original" source of humanity.

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2018 by Austin Whittall ©