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 © 

Kennewick Man, Ainus and Sunda

Two men were walking along the bank of the Columbia River in Washington, U.S. on July 28, 1996 when they came across a human skull, they quickly notified the autorities who inspected the area and, after a thorough search, managed to recover an almost complete skeleton, belonging to a man. This is how the "Kennewick Man" remains were discovered.

An initial inspection by an archaeologist [3] concluded that the man was not a Native American because the skull had a "caucasoid" appearance. A bone sample was sent to a laboratory and dated. We now know that these remains are between 8,000 and 9,500 years old. Which, by American standards is very ancient.

An act of Congress, The Native American Graves Protection and Repatriation Act (NAGPRA) was passed in 1990 with the goal of redressing the evils of the past, when Native American graves were profanated and their bones stolen and placed in Museums. The good intentions behind this legislation (which sought to return these remains to their original resting places in their tribal communities), has been distorted by the very extreme view held by some Native groups which are against any studies conducted on Paleo-Indian remains.

Actually most "modern" Native communities are relativley recent arrivals to their traditional tribal territories none go back more than one or two thousand years at most. Yet, they will go to court and in most cases succeed to get remains that date back thousands of years returned to "their homeland". In many cases the only link between the Paleoindians and the modern natives is the land where the remains were found.

Kennewick man was no exception, in 1996 some native groups took a hard stand and went to court to stop any further studies on his remains. Native American groups and the Federal Government of the U.S. combined their resources against the scientists who wanted to learn as much as possible from these extremely valuable ancient remains.

The legal battle took 8 years. In 1998 Kennewick man's bones were sent to the Burke Museum (Washington state) to protect them until the lawsuit was settled. They are still safeguarded at the Museum. [1]

The case was heard by a U.S. District Court which found that the remains could not be classified as "Native American". His ruling was appealed and the 9^th Circuit Court of Appeals upheld that decision in 2004. So the remains, which were found on Federal land, are now in custody of the U.S. Army Corps of Engineers and will remain so indefintely unless the Natives can somehwo prove in the future that Kennewick was a Native American. [1]

What have we learnt from the remains?

The remains comprise an almost complete skeleton (only some bones from hands and feet and the sternum are missing) which belonged to a tall man (about 1.73 m - 5.67 ft.) between 40 and 55 years old.

Although his DNA was intact, the tests were done on two samples but were inconclusive! (more on this below).

He had the broken off remains of a flint leaf-shaped serrated projectile stuck in his right ilium (hip bone). The wound was partly healed and the stone spear tip was quite large: 20 x 54 mm (0.78 x 2.12 in), [2] however, "The extensive amount of bone that has grown around the stone point suggests that the point was in place for a considerable amount of time and was not the cause of death." [3]

The surprising part is the skull, the cranial index marks it as dolichocephalic (long and narrow skull) instead of bracycephalic (short and wide skull) as found among the modern Natives. His face was narrow instead of broad and had a pronounced chin. This and other features gave him a "caucasoid" appearance. Yet, surprisingly his teeth were Sundadont like those found among South Asians. [2]

A clay facial reconstruction gives him a very "Caucasian" look indeed, [6] below is another view of this Kennewick man's face: [7]

Facial reconstruction of Kennewick Man. From [7]

Sundadonty is not a Caucasian trait, it is part of the "Mongoloid dental complex" [4], which evolved locally in Sunda Land (insular Indonesia); very similar teeth are found among Aboriginal Australians which "are also generally like those of Jomonese and some Ainus, suggesting that members of the late Pleistocene Sundaland population could have initially colonized Sahulland as well as the continental shelf of East Asia northward to Hokkaido" [4].

Map showing the Asian Range of Sundadonty (blue) and Sinodonty (yellow).. Copyright © 2014 by Austin Whittall

East Asians to the north of Sunda (i.e. Chinese, Japanese, Koreans, Mongolians) all have Sinodonty (dental shoveling). In general people belonging to Mongoloid groups (North Eastern Asians and, also American Indians) have the highest frequency of shoveled incisors while the rest of the world has chiseled ones. Shoveling is caused by the Ectodysplasin receptor gene (EDAR), also associated with hair thickness and the size and quantity of sweat and mammary glands. It is frequent in Asian populations and absent in Europeans, Africans, Denisovans and the Mal'ta remains from Siberia who carry the ancestral allele. The mutation or introgression (admixed through direct contact with H. erectus) is believed to have appeared in central China >30 Kya, a late date in my opinion since it was obtained by simulations restrained by a 15 Kya date for peopling America, which is far to recent. [5]

His teeth

The sundadonty of Kennewick man is an interesting find because it links him to Sunda (Insular Southeast Asia) and the archaic population of Japan, the Ainus.

The map above shows a discontinuity in the Sundadont range (blue). It is the intromission of Sinodonty (yellow) which cuts off Sunda in the south from Taiwan, Hokkaido, the Kuril Islands, Sakhalin and the tip of Kamchatka in the north.

This gap is currently occupied by sinodont populations (Chinese, Koreans and Japanese).

A simpe explanation for this discontinual distribution is that the sundadonts were the original people inhabiting the coastal areas of Oriental Asia and were later overlain or displaced to their current insular ranges by sinodonts in the central region (China, Korea, Japan).

Another option is that the Sundadonts moved from Sunda northwards, along the coastal areas in boats, or walked along the now submerged continental shelf alll the way to Hokkaido -the current Ainu territory (yet they seem to have failed occupying the main southern Japanese islands, or were later displaced from there by sinodonts).

Map showing a hypothetical route from Sunda to America for sundadont dental morphology. Copyright © 2014 by Austin Whittall

Did a branch of these migratory sundadonts reach America before the sindonts whose dental pattern now prevails among Amerindians? or, since sinodonty is a Homo erectus trait, did the admixture leading to Amerindian sinodonty take place in America, between an early arrived sundadont H. sapiens population and the ancient H. erectus settlers in America?

Turner [8] proposes a theory where "Sundadonty or more likely proto-Sundadonty, [is] the ancestral pattern for all modern humans". He bases this on the "generalized" appearance of Southeast Asian population: "they possess various external physical features of many geographic races, although usually in relatively low frequency". Interestingly "when South Siberian teeth are compared with those of Sundadonts, they show remarkable similarities. Because South Siberian [...] hybrid condition retrodicts the probable dental pattern of the common ancestors of Europeans and Asians before these derived groups drifted to their distinctive patterns by late Pleistocene times..." [8].

So Sunda would be the source of this dental morphology, coinciding with the place from which Y chromosome Haplogroup C radiated into New Guinea, Australia, India, China and Northeast Asia. see my post on haplogroup C. South Siberians also have hg. C at low frequencies.

The Ainu

Regarding the Ainu people of northern Japan, their similarity to Sunda populations such as the Bataks of Sumatra and the Dayaks of Borneo (shee photos below) was reported back in 1872 [9]. More recently, Genetic studies link the Ainu to Amerindians, and one should reread those studies within the hypothesis mentioned above. Perhaps both groups (Amerindians and Ainu) share a common sundadont ancestor which is the main reason for the similarities detected in these genetic studies.

Ainu people (Notice their curly hair in the top image and the clearly non-Mongolian look of the bearded man in the bottom one)
The impression I get is that they are two different kinds of Ainu.

Batak from Sumatra and Dayak from Borneo

His DNA

The DNA samples taken from the Kennewick man did not give any conclusive results. The tests were botched (since I don't believe in conspiracy theories, I can only guess that they were done in an inadequate manner or that the technology used was rudimentary, perhaps the samples were unwittingly contaminated, etc.). Below is the conclusions by the National Parks Service [3]:

"Thus, two separate amplifications from two different extractions suggested that Kennewick Man does not belong to haplogroup D (because the fragment was at least partially digested at np 5176 by Alu I restriction enzyme) while a single amplification from one of the two extractions suggests he might belong to haplogroup D. Given that at this point it was still unclear whether or not either of the two extractions were clean (i.e., uncontaminated), amplifications from the extracts had given conflicting information and neither extract had been successfully tested for the diagnostic markers for haplogroups B, C or X, it was impossible to determine to which, if any, of the common modern American Indian mtDNA haplogroups the Kennewick remains belong. At very most, our results provided, at this point, no evidence that the Kennewick remains belonged to haplogroups A, B or C [3]

In other words he does not belong to mtDNA haplogroups A, B or C. And the two tests gave differing results for hg. D (one was positive, the other negative). So he may or may not belong to haplogroup D. He may belong to X or M or any of the other haplogroups for which he was not tested.

I have not found any records regarding his Y chromosome analysis.

Closing Comments

Despite the theories put forth by some blogs and forums regarding a European origin for Kennewick man and the links they try to build between him and the Solutreans and Cro-Magnons of Europe, I am inclined towards an Asian origin for the Kennewick man

His sundadont teeth clearly set him apart from any modern Europeans. He is closer to the more ancient modern human migrants that reached Sunda, Sahul and what is now insular Eastern Asia in the Out of Africa initial migration some 70 kya. He probably represents this basic and most archaic line of modern humans who may have reached America not long after their departure from Africa. They do not, in my opinion, represent the more recent "white Europeans" as some propose (many of these forums have some white supremacist viewpoints, which I abhor).

A clear typing of his mtDNA and Y chromosomes will settle the issue of this man and lead us to ask interesting questions regarding why are there no contemporary Caucasoid-looking Native Americans? and maybe clarify what is the exact link between these Sundadont populations, perhaps some other gene similar to the EDAR of the sinodonts?

I will dig deeper into the Ainu, they intrigue me and I want to learn more about them.

Sources

[1] Burke Museum, Kennewick Man - The ancient one
[2] James C. Chatters, (2004). Kennewick Man. Smithsonian Institution.
[3] U.S. National Parks Service Article 1, and Article 2
[4] Turner C. G., et al., (1990). Major features of Sundadonty and Sinodonty, including suggestions about East Asian microevolution, population history, and late Pleistocene relationships with Australian aboriginals. Am J Phys Anthropol. 1990 Jul;82(3):295-317
[5] My post on an Early Peopling of America
[6] Sciencephoto.com, Clay reconstruction of the Kennewick Man
[7] Chelbea Fair, The Desert Town Right Around the Riverbend
[8] G. Richard Scott, Christy G. Turner, (2000). The Anthropology of Modern Human Teeth: Dental Morphology and Its Variation in Recent Human Populations . Cambridge University Press, pp. 303.
[9] Vivien de Saint-Martin, (1872). L'ethnologie du Grand Archipel d'Asie... races humaines. L'Anée Géogr. 9:90-97

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Patagonian Monsters - Cryptozoology, Myths & legends in Patagonia Copyright 2009-2014 by Austin Whittall ©