Layers Of Language and Culture In West Eurasia (An Overview)

A useful way to think about the history and pre-history of Europe and West Asia since the Last Glacial Maximum is to frame it as a matter of techno-linguistic-cultural waves and layers.

The earliest layer is that of European, Levantine, and Caucasian hunter-gatherers, who are quite distinct and segregated populations from each other and not merely gradual clines of different degrees of genetic admixture. Prior to the Neolithic Revolution, Caucasian hunter-gathers and Levantine hunter-gathers were as distinctively different from each other (or more so) than an Irishman is from a man in China today, at a distance of a few hundred miles from each other or less, and while there was admixture during and after the Fertile Crescent Neolithic Revolution, the populations remained very genetically distinct until the Bronze Age, and remain distinguishable, genetically, even today.

Hunter-gather cultures in Anatolia and the Levant are just on the verge of transitioning to becoming Neolithic farmer cultures in sites like Göbekli Tepe and signs of proto-farming in the Levant, when the Younger Dryas climate event hits and postpones that breakthrough for a few thousand years after these false starts, until you get the Fertile Crescent Neolithic revolution.

The Fertile Crescent Neolithic Revolution collects crops and animals, which are native to different parts of the Fertile Crescent, to domesticate and then combined them into a Fertile Crescent Neolithic package over a few centuries (that was tweaked somewhat with a few more domesticated plants and animals in each direction it expanded , like the donkey added in Egypt and a few more crops in Greece and the Balkans), but still not fully merging ethnically (and probably not linguistically either) and instead remaining ethnically distinct first farmer cultures and populations.

The group of first farmers derived from an ethnicity known in ancient DNA circles as Caucasian hunter-gatherers heads east to bring herding and farming into West Asia. 

The group of first farmers in the Levant heads south to bring herding and farming into Egypt, Ethiopia, and the rest of North Africa. This branch can be associated with the Afro-Asiatic languages, although whether the Afro-Asiatic languages originated in the Neolithic Levant, or were a back reaction from some Northeast African hunter-gatherer language that dominated culturally over the original Levantine farmers and then expanded back to North Africa and the Levant from perhaps Ethiopia or upper Egypt, is hard to know.

The group of first farmers in Western Anatolia expands in two waves that split off from the same source West into Europe: the Linear Pottery Neolithic (LBK) farmers more or less to the north along central and eastern Europe's great rivers, and the Cardial Pottery (CP) farmers more or less along the northern coast of the Mediterranean. The archaeological record also suggest that the LBK farmers in turn had a fairly basal split into two distinct branches with different material culture traditions. Probably each of the three major components of the Western Anatolian first farmer traditions spoke different, related languages within a Western Anatolian first farmer language family.

As these first farmers expanded into the more agriculturally suitable areas that they encountered, they did so primarily as families that largely replaced the much more thinly populated existing hunter-gather populations (in many cases, the population density of farmers was as much as a hundred times greater than the hunter-gathers who had preceded them in the area and the farmers were also much more prone to stand their ground to protect their crops than to migrate away although early herders were more flexible and hunter-gathers transitioned to herding more easily than they transitioned to farming), although a small number of locals were integrated into each expanding first farmer population, mostly women taken as brides by first farmer men.

In most of Europe, societies in cultural continuity with these first farmers start to collapse. Maybe its due to unsustainable agricultural practices, maybe its due to natural climate changes. In the earlier stages of this decline, there is some introgression of people who were part of remaining European hunter-gather cultures (sometimes partially adopting herding), who had continued to persist in places unsuitable for Neolithic farming, mostly women, and some first farmer communities revert to herding and hunting and gathering almost entirely, largely abandoning farming.

A few places, like Sumerian, Egyptian, and highlands West Asian societies, that were some of the earliest adopters of the Fertile Crescent Neolithic Revolution start to innovate and adopt early metallurgy, proto-writing or a true written language, and larger scale political and social organizations with kings and high priests who are more than just chiefs and shamans of small tribes or bands. They ran kingdoms and established full fledged bureaucratically organized religious institutions.

The ergative language speaking linguistic groups of the highlands of Anatolia, the Zargos mountains, and the Caucasus mountains, and of Mesopotamia constitute one group of early metal age societies which emerged in specific communities that used their technological advantages of conquer societies that were stuck in the Neolithic farming era and didn't adopt as fast. These specific communities probably emerged from an ergative language first farmer society. 

The Ancient Egyptians were another early adopter of metal age technologies whose military, economic, and cultural influence spread its non-ergative language Afro-Asiatic substrate across an ergative language first farmer substrate that extended, at least, to parts of North Africa, either via the Southern coast of the Mediterranean, or via Iberia.

The Harappan culture in the Indus River Valley and Northeast India is a third early adopter of technological advances of Copper Age technology that manages, somehow, to become dominant with much division or warfare.

Meanwhile, Pontic Caspian steppe hunter-gathers who had adopted herding and domesticated the horse, and made advances in making practical use of simple animal pulled wheeled vehicles enjoyed a fruitful synthesis with nearby West Anatolian derived first farmers from the LBK branch of their expansion into Europe, probably in Sredny Stog, that gave rise to the first proto-Indo-Europeans. Maybe the farmers had early metallurgy already, maybe the herders borrowed it from neighboring societies like the Caucasians, or maybe it arose independently not long after the synthesis that created proto-Indo-European society. But, the Indo-Europeans were already a society of Bronze Age horsemen and charioteers before their main expansion. One of the earliest expansions from this PIE core is east into the Tarim Basin where they become the Tocharians.

When a large area arid climate event strikes West and South Eurasia from Ireland to the Middle East and North Africa to Harappan territory in South Asia, in a time frame that peaks around 2500 BCE to 1500 BCE with climate leading its impacts on the affected societies, societies that were still Neolithic farming societies or were less advanced entirely collapse, while societies that had advanced to the copper age or very early Bronze Age are weakened and vulnerable to conquest, but don't actually collapse.

Indo-Europeans expand in all directions into the vacuum as the climate event strains existing cultures. In most places that had not reached the metal age when this happened and had almost fully collapsed, the Indo-Europeans almost fully wiped out existing hunter-gather and first farmer men, while assimilating some of the women from the existing culture, in patriarchal clans. This was the story for the Corded Ware Culture, more or less in central and eastern Europe, and the story of the Bell Beaker culture invaders of Great Britain and Ireland.

In places where early metal age cultures were still standing and just weakened when they arrived, like the Harappan culture of South Asia that gave rise to Sanskrit speaking Indo-Aryan society (that briefly extended as far as the Mittani empire at the Hittite Empire's border, before it was replaced by the Indo-Iranian Zoroastrian culture), the Hattic culture of Anatolia which came to be ruled by the Hittites, and the Aegean (where they conquered the Greeks and Hattic-related Minoans but adopted considerable parts of local language and culture), and Basque Country (it isn't entirely clear if this is a first farmer society remnant or if it was a distant and first farmer substrate influenced far outpost of early metal aged culture, probably the latter), there was more give and take, with a greater substrate influence on the conqueror's language through substrate influence, on the conqueror's religious beliefs and identity, and other deep rooted aspects of local culture (like Harappan curry). Isolating geography in Sardinia, and in the highlands of the Italian Peninsula's Etruscans and the Alpine kin, and in the highlands of Basque country (possibly assisted by its cluster of RH negative blood types) resisted longer and were changed less by the Indo-Europeans, although most ultimately fell to them, aided by somewhat less collapsed societies with somewhat more advanced early metal aged cultures (and by adopting technologies from the Indo-Europeans who sought to conquer them).

Indo-European expansion wasn't unchecked, however. The Afro-Asiatic cultures of the Levant and Egypt managed to resist them as did Mesopotamia, which Afro-Asiatic people had conquered from the Sumerians before this collapse. Geography prevented them from expanding past the Tarim Basin into East Asia and Central Asia. In Europe's far Northeast, as far as what is now Finland, northern Sweden, the Baltic states, and much of what is now Russia, as well as (much later) Hungary, Uralic people arrived as a post-Indo-European layer, or arrived to replace hunter-gather-fishing populations that the Indo-Europeans either never reached or didn't manage to hold onto for very long.

Then, another climate event triggered Bronze Age collapse around 1200 BCE. This only slightly unsettles the division of the world between Indo-European societies and Afro-Asiatic ones, for example, with the migration of Mycenaean Greek sea people into the Southern Levant after being fended off by more robust Egypt, to become the Philistines.

During Bronze Age collapse and its immediate aftermath, the waves of migration and war in Europe and West Asia become intramural fights between Indo-European people beyond the standoff between the Bell Beaker Indo-Europeans in Western Europe (known for their archery) and their successors, and the Corded Ware people to their east in Europe and their successors (know for their battle axes), whose respective territories shifted only a little (most notably midway through the Bronze Age in the vicinity of Denmark) from about 2000 BCE to 1200 BCE.

As Bronze Age collapse struck, Anatolian Indo-Europeans and Greek Indo-Europeans fought the Trojan War until it all collapsed into the Greek Dark Ages. Around that time that the Greek dark ages ended and classical Greek civilization started to emerge, Celtic peoples expanded from somewhere around Czechia as far west as Ireland and as far southeast as Anatolia. As Europe and West Asia emerged from the dark ages, the Iron Age technologies that the Hittites has managed to prevent its neighbors from acquiring prior to Bronze Age collapse, became ubiquitous. Iron wasn't actually a superior metal for weapons to bronze, but it was easier to mass produce and almost as good (and in isolated communities in what is now Iran, high quality steel far superior to bronze or iron was invented, around 1000 BCE, only to be lost centuries later before it could be adopted by the Roman Empire or the Islamic Empire).

The Iron Age was followed by Greco-Roman classical civilization eventually leading to Greek conquests of West Asia up to India (only to ultimately lose most of this territory one way or another in several big installments), and to the Roman Empire at its peak that reached Hadrian's Wall in Britain, Romania, the Levant, Egypt, North Africa, Southern Europe, and Anatolia. This lasted until another climate event led to the Western Roman Empire's collapse. A century after Rome collapsed, the Islamic empire expanded into vast swaths of territory in North Africa, Southwest and West Asia, Iberia, Malta, and the Balkans and beyond into South Asia and Southeast Asia and the Sahel, while the Christian Byzantine empire held on for several more centuries as it gradually lost territory, until it fell entirely to what had morphed in several rounds of Islamic regime changes into an Ottoman Empire that absorbed and coopted into Islam, Turkic invaders with origins in Southeast Asia, with the little interruption of the Mongol Empire along the way.

While the Islamic empire expanded, non-Muslim Europe collapsed with Germanic, Slavic, and Uralic tribes roaming about Europe in the migration period (giving rise to Hungary whose Magyar rulers swiftly converted from being pagans to Christians), Charlemagne and feudal lords running Europe in a fig leaf of a Holy Roman Empire under the Pope and launching several largely futile Crusades into the Levant, Viking raiders plundered Europe and briefly settled in North American only to collapse there and retreat (one branch of Vikings became the founding dynasty of and rulers of Russia's slavs), the Mongol Empire spanning from Persia to Korea before collapsing in the 13th century, multiple waves of the Black Plague ravaging Europe from its source in the east, missionaries re-Christianizing Europe from Ireland to the east until the last pagans of Europe are in the Urals and almost everyone else has nominally become Catholic by the late 11th century CE, and monastic orders doing what they could to preserve classical civilization for the thousand years of the Middle Ages that ended with the Columbian Exchange as Iberians and other Europeans colonized the Americas, with the Renaissance that finally restored classical Roman levels of technology and social order, with the Protestant Reformation, and with the end of the Reconquest of Iberia from the Moors. I'll end this survey with the start of this early modern period around 1500 CE.

Now, I'm writing this post from memory, which is partially why I've left out some things, particularly in West Asia and the details in far Northeast Europe and North Asia. I've omitted the stories of the Armenians, the Kurds, and the Druze. I skipped the process by which the Sumerian language was replaced by a succession of Semitic languages in Southwest Asia and Mesopotamia's trade ties to the Indus Valley Civilization and the eastern coast of Africa. I haven't adequately covered the history of the Caucuses. I've left out the fairly well understood history of the Jewish people after the fall of the Second Temple around 70 CE and haven't said enough about the Phoenicians and the Punic people. I haven't discussed the rise of Christianity during the Roman Empire or the demise of Zoroastrianism and polytheistic paganism. I haven't explored the connections between the Jewish, Christian and Muslim accounts of Southwest Asia's history through the legendary histories of their holy texts and the historical realities. Nor have I discussed why the Bronze Age is so full of what I call "legendary history" that is a mix of fiction and fact in purportedly fictional accounts that is so predominant across cultures in this era. I've left out the expansion around the time of the domestication of the camel that gave rise to the Berber people. I've barely touched on the repopulation of Europe by hunter-gathers from basically three refuges after the Last Glacial Maximum. I haven't mentioned the Silk Road or the history of Central Asia before and after the Mongols. I didn't cover how pottery made its way from the coasts of China and Japan about 16,000 years ago, over thousands of year to produce the transition from the pre-pottery Neolithic in the Levant to the pottery Neolithic of Europe and the Levant, or how the Black Plague got to Europe. I omitted the trans-Saharan slave trade, and Roman, Greek, Egyptian, Anatolian, Germanic and Viking slavery practices.

But the point isn't to cover all of  the details. That would take a book, not a blog post. It is to provide an overarching framework into which the details can fit and be better understood. And, of course, this post only minimally touches upon history and pre-history become its title's West Eurasian scope, even though there was lots going on in the rest of the world which is much less well known. Africa, for example, didn't start to fall decisively behind the rest of the world technologically and economically until perhaps five or six hundred years ago, and it took quite a while before the gaps became as stark as they are today (a process that only really started in earnest around the time of the trans-Atlantic slave trade that started after the time period covered by this post).

Actually, to be honest, I'd started trying to frame the issues presented by the origins, cultures, and linguistic history of Greece, West Asia and Southwest Asia up through the early Iron Age, and ended up going a little further afield with a forward outlining Western Civilization from its earliest roots which would be about ten pages in print. I did touch on that and frame it in a narrative (admittedly one that is sometimes hotly contested by legitimate academics even in the present). But, I can refocus on that in a more detailed and referenced manner in a future post. 

A Grammatical Gender And Ergativity Linguistics Refresher

Grammatical gender rules are not a feature that is shared by all Indo-European languages, or even a feature shared by all languages in the Germanic language family. 

Ergativity is is a grammatical feature with more uniformity, but is not uniform within the Indo-European or the Berber language family within the Afro-Asiatic language family.

Grammatical gender

Some of the Germanic languages (Icelandic, Norwegian, German, and Yiddish), the Slavic languages, and Greek have three grammatical genders (masculine, feminine, and neuter).

The subset of Germanic languages made up of Swedish, Danish, Dutch, and Flemish have a "common" and a neuter grammatical gender (the masculine grammatical gender and the feminine grammatical gender are merged relative to the three gender system).

The Celtic languages of the British Isles, the Romance languages, the Baltic languages (Lithuanian and Latvian), the Northern Kurdish languages, and the non-Indo-European Afro-Asiatic languages of Europe and the Mediterranean and the Middle East (Arabic including Maltese, Hebrew, Aramaic, the Berber languages, Coptic) have two grammatical genders (masculine and feminine). But, they don't have a neuter grammatical gender.

English (a Germanic language), the Central Kurdish languages, the non-Indo-European Uralic languages (Saami, Finnish, Estonian, and Hungarian), and the non-Indo-European Turkish languages do not have grammatical gender. Modern English, in common with Icelandic, Norwegian, and German does, however, have a masculine, feminine, and neuter third person singular pronoun (he, she, it), and Central Kurdish has a masculine and feminine but not neuter third person pronoun.

The Non-Indo-European Basque language has an animate noun class and an inanimate noun class that is called a grammatical gender, rather than an actually gender based grammatical gender system.

All of these languages are Indo-European language, except Basque, Turkish, the Uralic languages, the Afro-Asiatic languages (Arabic including Maltese, the Berber languages, and Hebrew).

Ergativity

Ergativity is another grammatical feature that doesn't strictly follow language family lines (probably due to substrate influences). Basque is ergative, as is Kurdish (which is spoken in an area where extinct ergative languages were once spoken), as are some Berber languages.

What is ergativity?

I'll quote the Wikipedia link above to make sure that I get it right:

In linguistic typology, ergative–absolutive alignment is a type of morphosyntactic alignment in which the subject of an intransitive verb behaves like the object of a transitive verb, and differently from the subject of a transitive verb. All known ergative languages show ergativity in their morphology, and a small portion also show ergativity in their syntax.

The ergative-absolutive alignment is in contrast to nominative–accusative alignment, which is observed in English, where the single argument of an intransitive verb behaves grammatically like the agent (subject) of a transitive verb but different from the object of a transitive verb. In ergative–absolutive languages with grammatical case, the case for the single argument of an intransitive verb and the object of a transitive verb is called the absolutive, and the case used for the agent of a transitive verb is called the ergative.

By one measure, 17% the world's languages use an ergative alignment in the marking of noun phrases. Examples of ergative-absolutive languages include Basque, Georgian, Mayan, Tibetan, Sumerian, and certain Indo-European languages such as Pashto, the Kurdish languages and many others.

The Population Genetics Of Egypt Have Been Stable For A Long Time

An ancient DNA sample from ca. 2500 BCE in Egypt reveals a great deal of continuity in the population genetics of Egypt then and the population genetics of Egypt today. 

I didn't have a lot of time to look carefully at this study, but prior studies have shown a modest increase in sub-Saharan African admixture since then, due to the trans-Saharan slave trade in more recent time periods.

Medieval Supernova

Astronomy is one of the oldest sciences. Advanced scientific astronomy calculation instruments were in existence in the Greek classical period and megalithic solar observatories came into existence independently shortly after agriculture was invented in multiple places. There is even evidence of astronomy being done in a stone temple in Anatolia built before agriculture was invented. By around Y1K, astronomy was taught in universities, and was done in a quite scientific manner in multiple civilizations around the globe. Prior to the Renaissance, however, it was all done with the naked eye. 

The remnant of the historical supernova SN 1181 is under discussion: While the previously suggested G130.7+3.1 (3C58) appears too old (3000-5000 yr), the unusual star IRAS 00500+6713 with a surrounding nebula (Pa-30) has an expansion age not inconsistent with a SN Iax explosion in AD 1181 under the assumption that neither acceleration nor deceleration occurred. 

Previously, only reports from China and Japan were known, pointing to an event near the northern circumpolar region. Any further reports from other cultures can therefore be highly relevant. 

We present here an Arabic poem in praise of Saladin by the contemporaneous author Ibn Sanā' al-Mulk (Cairo, Egypt). We re-date its composition to between Dec 1181 and May 1182. It contains a new bright star, which can be identified as SN 1181. The poem also provides new and independent information on the object type (called `najm' for `star'), location on sky (in or near the Arabic constellation al-Kaff al-Khabīb, lit. the henna-dyed hand (five bright stars in Cassiopeia), and brightness (brighter than alpha Cas, 2.25 mag). 

In addition, we present another Arabic text on SN 1006, also from Cairo, by the historian al-Maqrīzī, probably based on the contemporaneous al-Musabbihī

J.G. Fischer, H. Halm, R. Neuhäuser, D.L. Neuhäuser, "New Arabic records from Cairo on supernovae 1181 and 1006" arXiv:2509.04127 (published August 19, 2025 at 346 Astronomical Notes e70024).

The Punic People Were Mostly Greek, Not Levantine, In Ancestry

Ancient DNA from the Iron Age and classical Greco-Roman era reveals that the Punic people were much closer genetically to the Greeks and modern Sicilians than to the Phoenicians of the Levant who founded this maritime empire in the Western Mediterranean.

Punic people from this time period had been expected to be genetically similar to the Phoenicians were had often been assumed to be the ancestors of the Punic people, since archaeological and historical information indicated that the Phoenicians founded Carthage and other Punic cities. Linguistic information had also supported this expectation:

The Punic language, also called Phoenicio-Punic or Carthaginian, is an extinct variety of the Phoenician language, a Canaanite language of the Northwest Semitic branch of the Semitic languages. An offshoot of the Phoenician language of coastal West Asia (modern Lebanon and north western Syria), it was principally spoken on the Mediterranean coast of Northwest Africa, the Iberian Peninsula and several Mediterranean islands, such as Malta, Sicily, and Sardinia by the Punic people, or western Phoenicians, throughout classical antiquity, from the 8th century BC to the 6th century AD.

To the extent that the Punic people were genetically different from the Greeks, this was predominantly due to Iberian and Northwest African admixture, rather than due to Levantine admixture. 

Levantine admixture was completely absent from the Punic sample, except in three individuals (about 5% of the Punic sample analysed with Admixture) who were predominantly Levantine, and another four individuals who were predominantly North African in ancestry with very minor Levantine admixture (but with no Greek, Iberian, or other kinds of ancestry). 

This suggests a narrative in which a 95% Greek-like Punic people may have mostly replaced (without meaningful admixing with) a society in which some people with nearly purely Levantine Phoenicians, and some people were assimilated indigenous Northwest Africans with minor Phoenician ancestry.

Likewise, none of the contemporaneous ancient DNA from the Levant showed any Greek admixture at all, although three of eleven samples had small amounts of North African ancestry, and a fourth had small amounts of Iranian and Iberian ancestry (but no North African admixture).

The paper is Harald Ringbauer, et al., "Punic people were genetically diverse with almost no Levantine ancestors" Nature (April 2025).

As Bernard explains at his blog (via Google translate from French):

Phoenician culture emerged in Bronze Age city-states in the Levant. By the early first millennium BCE, the Phoenicians had established an extensive trade network along the Mediterranean coast as far south as the southwest shores of the Iberian Peninsula, spreading their culture, religion, and language. 

By the mid-sixth century BCE, Carthage, a Phoenician colony in present-day Tunisia, emerged as a major center of power in the central and western Mediterranean, as Levantine influence declined as their cities fell under the control of the Neo-Assyrian and Neo-Babylonian empires. Carthage subsequently came into conflict with Greek city-states in the fifth and fourth centuries BCE, and then with the Roman Empire in the third and second centuries BCE, before its final destruction in 146 BCE. 

In this article, the term Punic is given to all archaeological sites in the central and western Mediterranean associated with Phoenician culture, dated between the sixth and second centuries BCE, corresponding to the hegemony of Carthage in the region.

They analyzed the genomes of 210 ancient individuals from 14 Phoenician or Punic archaeological sites located in the Iberian Peninsula, Sardinia, Sicily, North Africa and the Levant dated between 600 and 150 BCE. There are no individuals older than 600 BCE, because before this date cremation was the most common burial method in these communities.

We don't know if the Phoenician founders of Carthage were later replaced by Greeks, if the original Bronze Age Phoenician colonists were recruited from Greece in the first place with a small endogamous caste of Levantine elites leading them, or if they were brought in by the Phoenicians later on as a caste of maritime people subordinate to the Phoenicians who ultimately rose to become the dominant caste in Punic society as the Bronze Age Phoenician maritime empire fell apart.

The ancient DNA samples come almost entirely from the time period at and after the Punic region lost close contact with the Phoenicians of the Levant.

It is possible that Levantine Phoenicians and Greek/North African/Iberian peoples co-existed in the Punic region but were basically genetically distinct endogamous castes, and that the Phoenician ancient DNA from this later period is mostly absent from the sample because Phoenicians continued to cremate their dead, rather than because they had been replaced, while the other caste that had substantially Greek ancestry buried their dead at this point. (The Bronze Age Greeks also mostly cremated their dead at the point in time when Indo-Europeans conquered them and converted them to an Indo-European language.)

This linguistic data can help us weigh which of the possible narratives to explain the ancient DNA is most plausible.

The fact that the Punic people spoke a Phoenician language, rather than Greek or Latin, however, despite their lack of significant Levantine Phoenician genetic ancestry, suggests that the ancestors of the Punic people with Greek ancestry underwent a language shift from Greek to Phoenician due to elite dominance by a Levantine Phoenician elite.  

If ancestors of the genetically Greek Punic people had replaced the Levantine Phoenician people by simply conquering them, we would have expected the Punic people to speak a language related to Greek rather than a North Semitic language (that is a close linguistic cousin of Hebrew and Arabic).

Yet, the lack of admixture between the caste whose members had any Levantine Phoenician ancestry, and the caste that is mostly Greek in genetic ancestry tends to disfavor the presence of the non-Levantine caste in the earliest Bronze Age founding period of Carthage. This inference is particularly strong in light of that fact that the Phoenicians did have some admixture with the indigenous North Africans who proceeded them in Carthage and the vicinity.

It is more plausible that the primarily Greek caste became part of Punic society in the roughly two and a half entry long time period from the mid-sixth century BCE, when Levantine influence declined as their cities fell under the control of the Neo-Assyrian and Neo-Babylonian empires, to the fifth and fourth centuries BCE, when Carthage subsequently came into conflict with Greek city-states. 

Before that, these Phoenician colonies were probably just Levantine Phoenicians and indigenous North Africans. It also seems likely that this demographic shift took place at the early end of this quarter millennium time period, allowing the dominant-subordinate status of the respective castes to emerge before the conflicts with the Greek city-states reached their high water mark.

Another possibility is that part of what keep a Levantine Phoenician caste distinct and endogamous from a caste with an ancestral Greek core, is that the Levantine Phoenician caste spoke Punic, while the caste with an ancestral Greek core spoke some dialect of Greek as their primary language, but didn't interact with the outside world much because the Levantine Phoenicians were the ruling caste of the Punic world, even though they made up only a modest percentage of the total population. 

This would have some similarities to the situation in medieval Finland while it was under Swedish rule, where power was held by Swedish speakers for centuries, even though most of the people spoke Finnish as their primary language, but with less genetic admixture between the two linguistic groups.

A Hypothetical Narrative Of Afro-Asiatic Origins

The Afro-Asiatic language family's origins are a hard nut to crack. This is a plausible proposal from Robert Bench.

Did Homo Sapiens Arise As A Hybrid?

In addition to being highly model dependent and unsupported by ancient DNA data, and hence somewhat speculative, this is really a less revolutionary proposal than it seems. 

Modern humans still arise in Africa (including a hypothetical admixture event that gives rise to the new species ca. 290,000 years ago). Basically, it is just proposing that in addition to the Neanderthal admixture shared by all non-Africans, and the Denisovan admixture that took place in the first generation of modern humans to reach Asia, there was an 20% admixture event from Homo erectus involving all modern humans associated with the emergence of the new species.

In their model, the 80% source, probably Homo heidelbergensis is also ancestral to Neanderthals and Denisovans, evolves from from Homo erectus about 1,500,000 years ago and suffers a severe bottleneck period, while the 20% Homo erectus ancestry was exclusive to Homo sapiens and was probably initially a larger percentage as a result of an admixture event in Africa about 290,000 years ago. The Homo erectus ancestry percentage was reduced in percentage over time due to its inferior selective fitness in most parts of the Homo erectus genome that have an impact on phenotypes (i.e. that have any actual discernible effect).

The evolutionary path leading to the rise of modern humans is full of twists and turns, and the latest surprise reveals that our species likely sprung forth from two ancient intermingling populations. A new study has confirmed that these groups first diverged from each other around 1.5 million years ago and later merged back together 300,000 years ago, initiating a genetic mixing event that culminated with the birth of modern humans.

The study, published in Nature Genetics, completely rewrites the story of humans. Scientists have long believed that Homo sapiens first appeared in Africa somewhere between 200,000 years and 300,000 years ago, having descended from a single ancestral lineage. The idea of genetic admixture flips the script, however, showing that human origins are much more complex than previously thought.

The researchers . . . tapped into modern human DNA from the 1000 Genomes Project, an international catalog filled with human genomes from a variety of populations. The research team created a computational algorithm called cobraa, which was designed to represent the event of an ancestral population splitting and rejoining. . . . 

With this method, they were able to produce a structured model that displayed two ancestral populations breaking apart in ancient times. In the years after this divergence, one of the populations experienced major fluctuations in size.

“Immediately after the two ancestral populations split, we see a severe bottleneck in one of them — suggesting it shrank to a very small size before slowly growing over a period of one million years,” said co-author Aylwyn Scally from the University of Cambridge’s Department of Genetics, in a statement. “This population would later contribute about 80 percent of the genetic material of modern humans and also seems to have been the ancestral population from which Neanderthals and Denisovans diverged.”

The second population, meanwhile, contributed 20 percent to the genetic makeup of modern humans. The researchers found that many of the genes this group passed along to humans were not located near regions of the genome corresponding to gene functions; this could reflect a concept called purifying selection, which is the process of natural selection filtering out harmful mutations. However, the researchers believe that some of the genes from the second population may have still been integral to brain development in modern humans.  . . .

An element of mystery still surrounds the identity of these ancestral populations. The researchers point to Homo erectus and Homo heidelbergensis as potential answers since they were present in Africa around the time of the genetic admixture, but further research is needed to match genetic ancestors with fossil groups.

Nature Genetics. A structured coalescent model reveals deep ancestral structure shared by all modern humans

From Discover Magazine. The abstract of the open access paper states:

Understanding the history of admixture events and population size changes leading to modern humans is central to human evolutionary genetics. Here we introduce a coalescence-based hidden Markov model, cobraa, that explicitly represents an ancestral population split and rejoin, and demonstrate its application on simulated and real data across multiple species. 

Using cobraa, we present evidence for an extended period of structure in the history of all modern humans, in which two ancestral populations that diverged ~1.5 million years ago came together in an admixture event ~300 thousand years ago, in a ratio of ~80:20%. Immediately after their divergence, we detect a strong bottleneck in the major ancestral population. 

We inferred regions of the present-day genome derived from each ancestral population, finding that material from the minority correlates strongly with distance to coding sequence, suggesting it was deleterious against the majority background. Moreover, we found a strong correlation between regions of majority ancestry and human–Neanderthal or human–Denisovan divergence, suggesting the majority population was also ancestral to those archaic humans.

Humans In African Jungles 150,000 Years Ago

Modern humans existed in wet tropical forests of West Africa about 150,000 years ago, contrary to the believe that humans could only survive in this kind of ecological environment through trade with farmers.

Humans emerged across Africa shortly before 300 thousand years ago (ka). Although this pan-African evolutionary process implicates diverse environments in the human story, the role of tropical forests remains poorly understood. 

Here we report a clear association between late Middle Pleistocene material culture and a wet tropical forest in southern Côte d’Ivoire, a region of present-day rainforest. Twinned optically stimulated luminescence and electron spin resonance dating methods constrain the onset of human occupations at Bété I to around 150 ka, linking them with Homo sapiens. Plant wax biomarker, stable isotope, phytolith and pollen analyses of associated sediments all point to a wet forest environment. 

The results represent the oldest yet known clear association between humans and this habitat type. The secure attribution of stone tool assemblages with the wet forest environment demonstrates that Africa’s forests were not a major ecological barrier for H. sapiens as early as around 150 ka.

Eslem Ben Arous, et al.,"Humans in Africa’s wet tropical forests 150 thousand years ago" Nature (February 26, 2025).

Early Homo Erectus In Spain And Where It Fits In The Larger Narrative

Overview

Anthropologists have found partial Homo erectus remains in Spain from 1.1-1.4 million years ago, adding to 1.97 million year old Homo erectus remains in Grăunceanu, Romania, and 1.77-1.85 million year old Homo erectus remains in Dmanisi, Georgia. 

Homo erectus first appears in Africa. Outside of Africa, Homo erectus remains are most often found in Indonesia and China, dating from around 108,000 years ago in Southeast Asia, back to about 70,000 years after this species evolved in Africa. 

Homo erectus went extinct in most of the world around 1,000,000 years ago, but persisted longer in Southeast Asia and possibly in East Asia, and relict populations of Homo erectus probably admixed with Denisovans at some point when both species existed. A major population bottleneck described below, probably took place in Homo erectus starting around 930,000 years ago, but it didn't result in the complete extinction of the species. Homo erectus was probably extinct by the time that modern humans first ventured beyond South Asia (not long after the Toba eruption ca. 75,000 years ago). It is plausible that the Toba eruption, followed by first contact with modern humans, may have led to the final extinction of Homo erectus, to the final extinction of H. floresiensis and H. luzonensis, and also to the extinction of Denisovans over most of their range (with the last relict Denisovans in Tibet probably going extinct in connection with their contacts with modern humans in this remote place).

We know that Homo erectus evolved in Africa rather than Eurasia, because that is where the species that Homo erectus evolved from, mostly likely H. habilis, but possibly some other African archaic hominin, was located at the time, and not just because the oldest Homo erectus remains are found there.

The oldest identified H. erectus specimen is a 2.04 million year old skull, DNH 143, from Drimolen, South Africa, coexisting with the australopithecine Paranthropus robustus. H. erectus dispersed out of Africa soon after evolution, the earliest recorded instances being H. e. georgicus 1.85 to 1.78 million years ago in Georgia and the Indonesian Mojokerto and Sangiran sites 1.8 to 1.6 million years ago.

(The quoted Wikipedia summary hasn't been updated to reflect the Romanian discovery announced earlier this year.)

Half a million years and a few hundred meters away from this site, there are Homo antecessor remains, from a time when Homo erectus had gone extinct in Europe, almost 700,000 years before Homo erectus went extinct in Asia.

The New Discovery

ATE7-1 fossil face (right) with mirrored 3D model (left). Credit: Maria D. Guillén / IPHES-CERCA / Elena Santos / CENIEH

When the global timeline passed one million years ago, more than half the span of hominin presence in Eurasia had already passed by. The earliest archaeological evidence in Eurasia is more than two million years old—found in places like Shangchen, China, and the Dawqara Formation of Jordan. Just this year Grăunceanu, Romania, joined the list of early archaeological traces of hominins in Europe, dating to an estimated 1.97 million years ago.

Still, I think about the threshold of one million years ago quite often. The number of sites in Eurasia with hominin evidence before one million years ago has grown quite large. It would have been hard to imagine this in 1990, when many scientists wondered if any sites in Eurasia were really older than this. Today there are many. And yet, the number of sites with fossils of hominins is quite a lot smaller than the number with stone artifacts or cutmarked animal bones. Most are in China or Indonesia, in addition to the exceptional site of Dmanisi, Georgia.

In western Europe there may be only two such sites, both in Spain: Sima del Elefante and Barranco Léon.

This week Rosa Huguet and collaborators have reported on a significant new addition to this very humble record. In work at Sima del Elefante in 2022, excavators uncovered a fragmentary facial skeleton, designated as ATE7-1. The estimated age of this fossil face is between 1.4 million and 1.1 million years ago. The new fossil joins two other hominin fossils from this cave deposit, within the same range of ages, a finger bone and a fragment of the front portion of a mandible with several worn teeth, ATE9-1. These fossils have been previously published, the mandible in 2008.

None of these fossils provide much to go on. Huguet and coworkers compared the facial anatomy of ATE7-1 with fossil faces attributed to Homo erectus from Dmanisi, Georgia, and Sangiran, Indonesia. They also compared the face to fossils from Gran Dolina, Spain, attributed to Homo antecessor. This site is located only a few hundred meters from Sima del Elefante but represents hominins and stone artifacts from around 780,000 years ago—as much as a half million years or more later than Sima del Elefante.

The ATE7-1 face is more like most H. erectus faces than either is like the later Gran Dolina fossils.

From John Hawks.

Context

Where does this discovery fit in the larger narrative of archaic hominin evolution?

Neanderthals, Denisovans, and modern humans (i.e. Homo sapiens) all share a Homo erectus ancestor and probably also at least one intermediate archaic hominin ancestor that evolved from Homo erectus.

The oldest archaeological evidence of modern humans, which is, order of magnitude consistent with age estimates for the most recent common ancestor of all modern human uniparental Y-DNA and mtDNA lineages, is about 300,000 years ago in Africa. Modern humans first left Africa around 125,000 to 100,000 years ago, and did so via the Middle East rather than Iberia. But the lion's share of non-African modern humans appear to be descended from a later wave of modern human expansion out of Africa about 50,000-74,000 years ago, with the lion's share of that wave closer to 50,000 years ago than 74,000 years ago. Neanderthal populations largely stalled this expansion into Europe until about 40,000 years ago. One or more of the hominin populations of Southeast Asia, and the jungles of Southeast Asia, probably stalled modern human expansion via the Southern route into Asia until around the time of the Toba eruption around 74,000 years ago (with the eruption possibly weakening these barriers and possibly also creating a reason for the modern humans of South Asia to expand to the Southeast).

The oldest Neanderthal remains are about 430,000 years old. Neanderthals were moribund by 40,000 years ago (with modern human Cro-Magnon people entering Europe around the same time that Neanderthals became extinct and overlapping with them for periods of a thousand or two thousand years or so in any one place), with the final relict population going extinct around 29,000 years ago. The leading explanations for Neanderthal extinction include a wave of volcanic eruptions, climate change, and the growing superiority of modern human hunter-gatherers due to their cultural evolution (e.g. stone technologies and the domestication of dogs) and/or genetic evolution. The range of Neanderthals extended from Northern Wales to the Middle East to South Asia and the Altai Mountains. There was significant Neanderthal admixture with modern humans, probably around 50,000-100,000 years ago (the latest estimates tend to favor a more recent date) in the vicinity of the Middle East or Iran (leaving a DNA legacy in all non-African modern humans), and there was also a more modern admixture with Altai Neanderthals (leaving a DNA legacy in Asian modern humans). Non-Africans today have up to 2% Neanderthal DNA, with Asians having a little more than Europeans, although ancient DNA from modern humans in ancient Eurasia, much closer to Neanderthal admixture sometimes have higher percentages of Neanderthal admixture. Neanderthals had bigger brains than modern humans, but also a more static material culture and less diverse range of hunting prey heavily concentrated around large megafauna (suggesting reduced brain plasticity and less ability to adapt culturally rather than genetically), with modern humans also relied on a wider array of smaller prey like rabbits, smaller birds, fish, and other seafood. At the time of first contact with modern humans, the effective population size of Neanderthals was about ten times smaller than the effective population size of modern human Cro-Magnons, and the effective Neanderthal effective population size ranged from about 3,000-12,000 throughout their existence and was fractured into multiple more or less isolated regional subpopulations.

Wikipedia says this about the extinction of Neanderthals:

The extinction of Neanderthals was part of the broader Late Pleistocene megafaunal extinction event. Neanderthals were replaced by modern humans, indicated by the near-complete replacement of Middle Palaeolithic Mousterian stone technology with modern human Upper Palaeolithic Aurignacian stone technology across Europe (the Middle-to-Upper Palaeolithic Transition) from 41,000 to 39,000 years ago. Iberian Neanderthals possibly persisted until about 35,000 years ago, modern human expansion perhaps impeded by the Ebro River. Neanderthals in Gibraltar may have survived as late as 28,000 years ago at Gorham's Cave. The dating of these late Iberian sites is contested.

Historically, the cause of extinction of Neanderthals and other archaic humans was viewed under an imperialistic guise, with the superior invading modern humans exterminating and replacing the inferior species.

When sapiens began to expand and spread, he eliminated the other contemporary races [including Neanderthals] just as the white man drove out the Australian aborigines and the North American Indians.

— Ernst Mayr, 1950

The assimilation of Neanderthal populations into modern human populations had long been hypothesised with supposed hybrid specimens, and was revitalised with the discovery of archaic human DNA in modern humans. Similarly, the Châtelperronian industry of central France and northern Spain may represent a culture of Neanderthals adopting modern human techniques, via acculturation. Other ambiguous transitional cultures include the Italian Uluzzian industry, and the Balkan Szeletian industry.

Aside from competition with modern humans, Neanderthal extinction has also been ascribed to their low population as well as the resulting mutational meltdown, making them less adaptable to major environmental changes (specifically Heinrich event 4) or new diseases.

The admixture between modern humans and Neanderthals went in both directions. And, some of the late archaeological tool cultures of Neanderthal, which coincide with the arrive of modern humans in Europe, may reflect the increased brain plasticity of hybrid Neanderthal-modern human individuals.

Denisovans (named after the cave in the Altai where the type remains were discovered) probably existed from at least 285,000 years ago to about 25,000 years ago, general in Asia to the east of the Neanderthal range from Altai and Tibet to Southeast Asia, and overlapping with the Neanderthal range in the Altai region. High altitude adaptation DNA admixed from Denisovans are found in Tibetans. Trace levels of Denisovan admixture are found in mainland Southeast Asia and East Asia, and in island Southeast Asia up to the Wallace Line. Modern humans with Australian aboriginal ancestry or Papuan ancestry or Filipino negrito ancestry have substantial Denisovan ancestry (up to 6%) in addition to their Neanderthal ancestry (up to 2%). Presumably, the Denisovan-modern human admixture whose legacies exist in Australian aborigines, Papuans, Filipino negritos, and mainland Southeast Asians and East Asians must have occurred around the time of first contact between the first wave of modern humans in Asia around 50,000 to 75,000 years ago, and was then greatly diluted by subsequent waves of modern human migration west of the Wallace line in Asia. Also, Denisovans presumably went extinct within a thousand or two thousand years or so of first contact with modern humans (which took place much later in Tibet than almost everywhere else).

The exact path from Homo erectus to modern humans, Neanderthals, and Denisovans (and possibly other now extinct archaic species derived from Homo erectus) is a matter of ongoing investigation and debate.

Denisovan mtDNA diverged from that of modern humans and Neanderthals about 1,313,500–779,300 years ago; whereas modern human and Neanderthal mtDNA diverged 618,000–321,200 years ago. Krause and colleagues then concluded that Denisovans were the descendants of an earlier migration of H. erectus out of Africa, completely distinct from modern humans and Neanderthals.

However, according to the nuclear DNA (nDNA) of Denisova 3—which had an unusual degree of DNA preservation with only low-level contamination—Denisovans and Neanderthals were more closely related to each other than they were to modern humans. Using the percent distance from human–chimpanzee last common ancestor, Denisovans/Neanderthals split from modern humans about 804,000 years ago, and from each other 640,000 years ago. 

Using a mutation rate of 1×10^−9 or 0.5×10^−9 per base pair (bp) per year, the Neanderthal/Denisovan split occurred around either 236–190,000 or 473–381,000 years ago respectively. Using 1.1×10^−8 per generation with a new generation every 29 years, the time is 744,000 years ago. Using 5×10^−10 nucleotide site per year, it is 616,000 years ago. Using the latter dates, the split had likely already occurred by the time hominins spread out across Europe. 

H. heidelbergensis is typically considered to have been the direct ancestor of Denisovans and Neanderthals, and sometimes also modern humans. Due to the strong divergence in dental anatomy, they [i.e. Denisovans] may have split before characteristic Neanderthal dentition evolved about 300,000 years ago.

The more divergent Denisovan mtDNA has been interpreted as evidence of admixture between Denisovans and an unknown archaic human population, possibly a relict H. erectus or H. erectus-like population about 53,000 years ago. Alternatively, divergent mtDNA could have also resulted from the persistence of an ancient mtDNA lineage which only went extinct in modern humans and Neanderthals through genetic drift. Modern humans contributed mtDNA to the Neanderthal lineage, but not to the Denisovan mitochondrial genomes yet sequenced. The mtDNA sequence from the femur of a 400,000-year-old H. heidelbergensis from the Sima de los Huesos Cave in Spain was found to be related to those of Neanderthals and Denisovans, but closer to Denisovans, and the authors posited that this mtDNA represents an archaic sequence which was subsequently lost in Neanderthals due to replacement by a modern-human-related sequence.

The intermediate species that is the most recent common ancestor of Neanderthals, Denisovans, and modern humans probably arose not long after a genetic bottleneck which has been inferred from modern DNA. This genetic bottleneck probably occurred in the clade of H. erectus which is ancestral to modern humans. As one secondary source explaining this notes:

Between 930,000 and 813,000 years ago, something nearly ended humanity before it even began. A mysterious bottleneck reduced the human breeding population to just 1,280 individuals, pushing our ancestors to the brink of extinction for an astonishing 117,000 years. 

Scientists have long puzzled over a gap in the African and Eurasian fossil records, and now, a team of researchers may have found the answer. Using a groundbreaking method called FitCoal, they analyzed the genomes of 3,154 modern humans to reconstruct ancient population sizes. What they found was staggering. Nearly 99% of early humans vanished, likely due to extreme climate events such as glaciations, severe droughts, and the collapse of ecosystems.

The world was changing. Glaciation, extreme droughts, and collapsing ecosystems made survival nearly impossible. Food sources vanished, and so did most of our ancestors. Those who remained – just a tiny fraction of the original population – fought to endure in a harsh and unpredictable environment. 

But against all odds, they survived. And in doing so, they may have changed the course of human evolution forever. Scientists believe this bottleneck could have led to the merging of two ancestral chromosomes, forming what we now know as chromosome 2 – a key feature that separates modern humans from other primates.

Around 813,000 years ago, the climate began to shift. Our ancestors may have mastered fire, allowing them to cook food, stay warm, and fend off predators. Populations rebounded, and from that tiny group of survivors, the future of humanity was born. 

This discovery reshapes our understanding of human history, and raises new questions. Where did these survivors live? How did they overcome such extreme conditions? Did this struggle push human intelligence to evolve faster?

The paper that is the basis for this account is Wangjie Hu, et al., "Genomic inference of a severe human bottleneck during the Early to Middle Pleistocene transition" 381(6661) Science 979-984 (August 31, 2023). Its abstract materials state:

Editor’s summary 

Today, there are more than 8 billion human beings on the planet. We dominate Earth’s landscapes, and our activities are driving large numbers of other species to extinction. Had a researcher looked at the world sometime between 800,000 and 900,000 years ago, however, the picture would have been quite different. Hu et al. used a newly developed coalescent model to predict past human population sizes from more than 3000 present-day human genomes (see the Perspective by Ashton and Stringer). The model detected a reduction in the population size of our ancestors from about 100,000 to about 1000 individuals, which persisted for about 100,000 years. The decline appears to have coincided with both major climate change and subsequent speciation events. —Sacha Vignieri 

Abstract 

Population size history is essential for studying human evolution. However, ancient population size history during the Pleistocene is notoriously difficult to unravel. In this study, we developed a fast infinitesimal time coalescent process (FitCoal) to circumvent this difficulty and calculated the composite likelihood for present-day human genomic sequences of 3154 individuals. Results showed that human ancestors went through a severe population bottleneck with about 1280 breeding individuals between around 930,000 and 813,000 years ago. The bottleneck lasted for about 117,000 years and brought human ancestors close to extinction. This bottleneck is congruent with a substantial chronological gap in the available African and Eurasian fossil record. Our results provide new insights into our ancestry and suggest a coincident speciation event.

The proposed climate event was part of the Mid-Pleistocene Transition. Some key aspects of this, in places where Homo erectus reached, were as follows:

Europe

In Europe, the MPT was associated with the Epivillafranchian-Galerian transition and may have led to the local extinction of, among other taxa, Puma pardoides, Megantereon whitei, and Xenocyon lycaonoides. The prevalence of ungulates adapted for grazing increased in the Mediterranean region after the "0.9 Ma event". The northern North Sea Basin was first glaciated during the MPT. The increased intensity of transgressive-regressive cycles is recorded in northern Italy.

Asia

The cooling brought about by the MPT increased westerly aridity in the western Tarim Basin. East Asian Summer Monsoon (EASM) precipitation declined. Grasslands expanded across the North China Plain as forests contracted.

During the MPT, the Indian Summer Monsoon (ISM) decreased in strength. In the middle of the MPT, there was a sudden decrease in denitrification, likely due to increased solubility of oxygen during lengthened glacial periods. After the MPT, the Bay of Bengal experienced increased stratification as a result of the strengthening of the ISM, which resulted in increased riverine flux, inhibiting mixing and creating a shallow thermocline, with stratification being stronger during interstadials than stadials. Paradoxically, variability in Δδ18O in the Bay of Bengal between glacials and interglacials decreased following the MPT.

Africa

In Central Africa, detectable floral changes corresponding to glacial cycles were absent prior to the MPT. Following the MPT, a clear cyclicity became evident, with interglacials being characterised by warm and dry conditions while glacials were cool and humid.

According to one of the leading papers on the 0.9 Ma Event, closely associated with the Homo erectus genetic bottleneck:

The Early-Middle Pleistocene Transition (EMPT) (ca. 1.4–0.4 Ma) represents a fundamental transformation in the Earth's climate state, starting at 1.4 Ma with a progressive increase in the amplitude of climatic oscillations and the establishment of strong asymmetry in global ice volume cycles. The progressive shift from a 41kyr–100kyr orbital rhythm was followed by the first major build-up of global ice volume during MIS 24-22, the so-called “0.9 Ma event”. The Vallparadís Section (Vallès-Penedès Basin, NE Iberian Peninsula) is one of the few Pleistocene series in Europe that spans the onset of the transition (from 1.2 to 0.6 Ma), thus representing a pivotal array of localities to investigate the effect of glacial dynamics on environmental conditions in Southern Europe. Here we inspect the effects of the EMPT on terrestrial ecosystems by examining the dietary adaptations (through dental meso- and microwear patterns) of fossil ungulates from the Vallparadís Section dated before and after the “0.9 Ma event”. Results show a steady presence of open grasslands before MIS 22 and more humid conditions at MIS 21. Both before and after MIS 22, a consistent presence of ungulates with long-term patterns that point to a grazing or grass-rich mixed feeding behaviour is observed, while noticeably, short-term patterns point to increased seasonality right after the “0.9 Ma event” glacial period. This increment of seasonality may have had an important effect on the Mediterranean habitats leading to recurring changes in the quality of plant resources available to large herbivores, which in response periodically adopted more mixed feeding behaviours widening their dietary breadth to consume also sub-optimal food items during adverse seasons.

In particular, during this event, global ice volumes increased substantially, and the Northern Hemisphere experienced increased seasonality and aridity, and surface sea temperatures in the North Atlantic reached their lowest values during the EMPT at this time. Also, grasslands expanded across the North China Plain as forests contracted.

This hypothesis is model dependent, could be impacted by sources of systemic error, like the possible much later extinction of Homo erectus populations derived from the same source population, later hard genetic sweeps of Homo erectus source genes, the effective extinction of modern humans arising from other clades of Homo erectus at some much later time, a lack of consideration of Neanderthal or Denisovan genes in the analysis, and a complete lack of ancient Homo erectus genomes. 

Also, in understanding this narrative one has to recognize that genetics researchers call an "effective population" of 1,280 individuals could have involved a census population at any one time that was many times larger than that. And, this is still about five times as large as the effective population size of the founding population of the Americas, for example. So, the bottleneck wasn't quite as extreme as some popular accounts of it would imply.

But the oldest examples of the species Homo antecessor does first appear in Europe, shortly after this inferred bottleneck, and there are no Homo erectus remains in Europe during or after the time of this inferred bottleneck.

Homo antecessor (Latin "pioneer man") is an extinct species of archaic human recorded in the Spanish Sierra de Atapuerca, a productive archaeological site, from 1.2 to 0.8 million years ago during the Early Pleistocene. Populations of this species may have been present elsewhere in Western Europe, and were among the first to settle that region of the world, hence the name. The first fossils were found in the Gran Dolina cave in 1994, and the species was formally described in 1997 as the last common ancestor of modern humans and Neanderthals, supplanting the more conventional H. heidelbergensis in this position. H. antecessor has since been reinterpreted as an offshoot from the modern human line, although probably one branching off just before the modern human/Neanderthal split.

Despite being so ancient, the face is unexpectedly similar to that of modern humans rather than other archaic humans—namely in its overall flatness as well as the curving of the cheekbone as it merges into the upper jaw—although these elements are known only from a juvenile specimen. Brain volume could have been 1,000 cc (61 cu in) or more, but no intact braincase has been discovered. This is within the range of variation for modern humans. Stature estimates range from 162.3–186.8 cm (5 ft 4 in – 6 ft 2 in). H. antecessor may have been broad-chested and rather heavy, much like Neanderthals, although the limbs were proportionally long, a trait more frequent in tropical populations. The kneecaps are thin and have poorly developed tendon attachments. The feet indicate H. antecessor walked differently than modern humans.

H. antecessor was predominantly manufacturing simple pebble and flake stone tools out of quartz and chert, although they used a variety of materials. This industry has some similarities with the more complex Acheulean, an industry which is characteristic of contemporary African and later European sites. Groups may have been dispatching hunting parties, which mainly targeted deer in their savannah and mixed woodland environment. Many of the H. antecessor specimens were cannibalised, perhaps as a cultural practice. There is no evidence they were using fire, and they similarly only inhabited inland Iberia during warm periods, presumably retreating to the coast otherwise.

Meanwhile:

Homo heidelbergensis (also H. erectus heidelbergensis, H. sapiens heidelbergensis) is an extinct species or subspecies of archaic human which existed from around 600,000 to 300,000 years ago, during the Middle Pleistocene. Homo heidelbergensis was widely considered the most recent common ancestor of modern humans and Neanderthals, but this view has been increasingly disputed since the late 2010s.

In the Middle Pleistocene, brain size and height were comparable to modern humans. Like Neanderthals, H. heidelbergensis had a wide chest and robust frame.

Fire likely became an integral part of daily life after 400,000 years ago, and this roughly coincides with more permanent and widespread occupation of Europe (above 45°N), and the appearance of hafting technology to create spears. H. heidelbergensis may have been able to carry out coordinated hunting strategies, and consequently they seem to have had a higher consumption of meat.

It is debated whether or not to constrain H. heidelbergensis to only Europe or to also include African and Asian specimens, and this is further confounded by the type specimen (Mauer 1) being a jawbone, because jawbones feature few diagnostic traits and are generally missing among Middle Pleistocene specimens.

H. heidelbergensis was subsumed in 1950 as a subspecies of H. erectus but today it is more widely classified as its own species. H. heidelbergensis is regarded as a chronospecies, evolving from an African form of H. erectus (sometimes called H. ergaster).

At least three other archaic hominin species overlapped with hominins from the H. erectus era or later.

• Homo floresiensis – Extinct small human species found in Flores
• Homo luzonensis – Archaic human from Luzon, Philippines
• Homo naledi – South African archaic human species

H. floresiensis and H. luzonensis may have been regional variations of the same species and show similarities with each other. The most plausible theory of their phylogenetic position, in my view, is that both of them were sub-species of H. habilis, and may have left Africa, either independently, or together with either H. erectus, the Denisovan ancestor, or Denisovans themselves. H. floresiensis and Denisovans (and possibly the earliest modern humans to arrive there as well) may have co-existed on the island of Flores, Indonesia (which is past the Wallace line) at some point in  time. There are no remains of H. floresiensis, H. luzonensis, H. habilis, or any other archaic hominins before H. erectus disperses from Africa. 

H. naledi was a South African archaic hominin species that flourished from 335,000 to 226,000 years ago, that was probably not directly ancestral to modern humans or any other non-African archaic hominins, but would have co-existed in time (and possibly space) with the earliest modern humans in Africa.

A November 6, 2024 post at this blog recapped some other possible non-African archaic hominins who existed at the same time that modern humans did: 

Notably the remains of the Red Deer Cave People of China from 14,000 years ago (a few thousand years before the start of the Holocene era) are genetically modern humans and are not archaic hominins despite some of their seemingly archaic features. See also here.

I am also inclined to think that they may yet be a small relict population of small archaic hominins in a remote Indonesian jungle on the island of Sumatra and perhaps Flores as well, where these cryptids, called Orang Pendek, locally, have been attested but not definitively confirmed to still exist. I discuss this further at this post.

Homo floresiensis (discovered in 2003) are commonly known as "hobbits" and have been found on the island of Flores. Their phylogeny is disputed, but I find the theory that they are an asian branch of H. habilis to be most convincing. H. luzonesis (discovered in 2007) is similar and contemporaneous, but found further east in the Philippines and is supported by a less complete archaeological record. Both of these diminutive species are found in association with late Pleistocene tools and "oriental fauna".

Personally, being more of a lumper than a splitter, I'm inclined to see H. floresiensis and H. luzonesis as sub-species variations of the same species ("race" within that species to use some outdated terminology), and likewise to see H. longi, H. juluensis, and Denisovans as sub-species variations of the Denisovan species. The Hualongdong archaic hominin fossils ... could be a hybrid individual, perhaps a Neanderthal-Denisovan hybrid individual (something that has precedent in a Denisovan cave DNA sample).

Academic anthropologists, in contrast, tend to be splitters, in part, because it is cool and career advancing to discover and name your own archaic species, in part because the data is so fragmentary that grouping different fragmentary remains in a clade presumes relationships between the remains that aren't solidly proven, and in part, because it is easy to underestimate how much morphological diversity is possible within a single species if populations of it exposed to different environmental conditions.

H. longi a.ka. "dragon man" dates to an earlier time period (still contemporaneous with modern humans in Africa) in China and Manchuria, was discovered in 1933, and has been hypothesized to be a sister clade to Neanderthals, Denisovans, and modern humans, and a descendant of the pre-modern human hominin species H. antecessor due in part to basal archaic features in the skull.

H. juluensis (literally "big heads") is contemporaneous H. longi, and beyond that time frame into the time frame of H. floresiensis and was discovered from 1976-1979 in China and Tibet. The authors assign this specimen along with Xiahe and Penghu fossils, to the Denisovan species (a sister clade to Neanderthals and modern humans) based upon comparisons of their fossil teeth and rough geographic proximity. H. juluensis is found in association with early Paleolithic tools and remains of Paleoarctic fauna. But they have larger brain cases than H. longi. A previous suggestions of the link between H. longi and the Denisovan species are discussed here and here at this blog. At least one Denisovan tooth has been found in Laos dated to 131,000 years ago.

The article also discusses the Hualongdong archaic hominin fossils that "date to the late Middle Pleistocene (~300,000 years BP) and display a mosaic of characteristics that cannot be easily fitted into any one lineage," although they are closer to H. longi and H. juluensis. This individual could be a hybrid between these two subspecies, with H. erectus, or with a Neanderthal who was far east of his usual range.

Prior to 2021, H. longi and H. juluensis tended to be classified as H. erectus (remains of which start to appear at a much greater time depth in Asia) or as archaic modern humans.

The Narmada and Maba partial skulls, especially the latter, are suggestively associated with Neanderthals by the article.

These Asian archaic species also overlap in time with the Southern African archaic hominin clade H. naledi which is a sister clade to the modern human ancestors and to the common ancestor of modern humans, Neanderthals, and Denisovans, but is not actually among our ancestors. As I explained at the link, this species "is basically a story from The Silmarillion of hominin evolution. It is entertaining, especially for hard core human evolution fans, but it doesn't really advance the plot."

A small number of papers reported genetic evidence in modern Africans of admixture with an archaic hominin "ghost species" in Africa, but subsequent papers have explained this "ghost species" signal as a methodological artifact that merely arises from population structure in early modern human Africans (see also here). But there may have been relict archaic hominins that did not admix with modern humans in Africa that were also contemporaneous with modern humans, at least, early on.

The question of whether behaviorally modern humans started showing advanced behavior around 70,000-50,000 years ago (at the dawn of the Upper Paleolithic era and close in time to the Out of Africa event for modern humans), was associated with an evolutionary leap in their brains is an open and unresolved question. See also here (addressing the question of what made modern humans genetically distinct from archaic hominins).