Consider Homo erectus …

Championed as the earliest commonly found human species and, apart from anatomically modern humans (AMH), the most widespread through Africa and Eurasia. It also endured longer (~1.75 Ma) than any other hominin species, appearing first in East Africa around 2 Ma ago, the youngest widely accepted fossil – found in China – being around 250 ka old. The ‘erects’ arguably cooked their food and discovered the use of fire 1.7 to 2 Ma ago. The first fossils discovered in Java by Eugene Dubois are now known to be associated with the oldest-known art (430 to 540 ka) The biggest issue surrounding H. erectus has been its great diversity, succinctly indicated by a braincase capacity ranging from 550 to 1250 cm3: from slightly greater than the best endowed living apes to within the range of AMH. Even the shape of their skulls defies the constraints placed on those of other hominin species. For instance, some have sagittal crests to anchor powerful jaw muscles, whereas others do not. What they all have in common are jutting brow ridges and the absence of chins along with all more recently evolved human species, except for AMH.

This diversity is summed up in 9 subspecies having been attributed to H. erectus, the majority by Chinese palaeoanthropologists. Chinese fossils from over a dozen sites account for most of the anatomical variability, which perhaps even includes Denisovans, though their existence stems only through the DNA extracted from a few tiny bone fragments. So far none of the many ‘erect’ bones from China have been submitted to genetic analysis, so that connection remains to be tested. Several finds of diminutive humans from the Indonesian and Philippine archipelagos have been suggested to have evolved from H. erectus in isolation. All in all, the differences among the remains of H. erectus are greater than those used to separate later human species, i.e. archaic AMH, Neanderthals, Denisovans, H. antecessor etc. So it seems strange that H. erectus has not been split into several species instead of being lumped together, in the manner of the recently proposed Homo bodoensis. Another fossil cranium has turned up in central China’s Hubei province, to great excitement even though it has not yet been fully excavated (Lewis, D. 2022. Ancient skull uncovered in China could be million-year-old Homo erectus. Nature News 29 November 2022; DOI: 10.1038/d41586-022-04142-00; see also a video). Chances are that it too will be different from other examples. It also presents a good excuse to consider H. erectus.

Cranium of a Chinese Homo erectus, somewhat distorted by burial, from a site close to the latest find. (Credit: Hubei Museum, Wuhan, China)

The complications began in Africa with H. ergaster, the originator of the bifacial or Acheulean multi-purpose stone tool at around 1.6 Ma (see: Flirting with hand axes; May 2009), the inventor of cooking and discoverer of the controlled use of fire. ‘Action Men’ were obviously smarter than any preceding hominin, possibly because of an increase of cooked protein and plant resources that are more easily digested than in the raw state and so more available for brain growth. The dispute over nomenclature arose from a close cranial similarity of H. ergaster to the H. erectus discovered in Java in the 19th century: H. erectus ergaster is now its widely accepted name. In 1991-5 the earliest recorded hominins outside Africa were found at Dmanisi, Georgia, in sediments dated at around 1.8 Ma (see: First out of Africa; November 2003) Among a large number of bones were five well-preserved skulls, with brain volumes less than 800 cm3 (see: An iconic early human skull; October 2013). These earliest known migrants from Africa were first thought to resemble the oldest humans (H.habilis) because of their short stature, but now are classified as H. erectus georgicus. They encapsulate the issue of anatomical variability among supposed H. erectus fossils, each being very different in appearance, one even showing ape-like features. Another had lost all teeth from the left side of the face, yet had survived long after their loss, presumably because others had cared for the individual.

The great variety of cranial forms of the Asian specimens of H. erectus may reflect a number of factors. The simplest is that continuous presence of a population there for as long as 1.5 Ma inevitably would have resulted in at least as much evolution as stemmed from the erects left behind in Africa, up to and including the emergence of AMH in North Africa about 300 ka ago. If contact with the African human population was lost after 1.8 Ma, the course of human evolution in Africa and Asia would clearly have been different. But that leaves out the possibility of several waves of migrants into Asia that carried novel physiological traits evolved in Africa to mix with those of earlier Asian populations. From about 1 Ma ago a succession of migrations from Africa populated Europe – H. antecessor, H. heidelbergensis, and Neanderthals and then AMH. So a similar succession of migrants could just as well have gone east instead of west on leaving Africa. Asia is so vast that migration may have led different groups to widely separated locations, partially cut-off by mountain ranges and deserts so that it became very difficult for them to maintain genetic contact. Geographic isolation of small groups could lead to accelerated evolution, similar to that which may have led to the tiny H. floresiensis and H. luzonensisdiscovered on Indonesian and Philippine islands.

 Another aspect of the Asian continent is its unsurpassed range of altitude, latitude and climate zones. Its ecologically diversity offers a multitude of food resources, and both climate and elevation differences pose a range of potential stresses to which humans would have had to adapt. The major climate cycles of the Pleistocene would have driven migration across latitudes within the continent, thereby mixing groups with different physical tolerances and diets to which they had adapted. Equally, westward migration was possible using the Indo-Gangetic plains and the shore of the Arabian Sea: yet more opportunities for mixing between established Asians and newly arrived African emigrants.

Seven thousand years of cultural sharing in Europe between Neanderthals and modern humans

Two years ago material excavated from the Bacho Kiro cave in Bulgaria revealed that anatomically modern humans (AMH) had lived there between 44 and 47 ka ago: the earliest known migrants into Europe. Bacho Kiro contains evidence of occupancy by both Neanderthals and AMH. This discovery expanded the time over which Europe was co-occupied by ourselves and Neanderthals. The latter probably faded from the scene as an anatomically distinct group around 41 to 39 ka, although some evidence suggests that they lingered in Spain until ~37 ka and perhaps as late as 34 to 31 ka in the northern Ural mountains at the modern boundary of Europe and Asia. For most of Europe both groups were therefore capable of meeting over a period of seven to eight thousand years.

Aside from interbreeding, which they certainly did, palaeoanthropologists have long pondered on a range of tools that define an early Upper Palaeolithic culture known as the Châtelperronian, which also spans the same lengthy episode. But there have been sharp disagreements about whether it was a shared culture and, if so, which group inspired it. Evidence from the Grotte du Renne in eastern France suggests that the Neanderthals did abandon their earlier Mousterian culture to use the Châtelperronian approach early in the period of dual occupancy of Europe.

Dated appearances in France and NE Spain of Neanderthal fossils (black skulls), Châtelperronian artefacts (grey circles) and proto-Aurignacian artefacts (white squares) in different time ‘slots’ between 43.4 and 39.4 ka. (Credit: Djakovic et al., Fig. 3)

Igor Djakovic of Leiden University in the Netherlands , Alastair Key of Cambridge University, UK, and Marie Soressi, also of Leiden University have undertaken a statistical analysis of the geochronological and stratigraphic context of artefacts at Neanderthal and AMH sites in France and NW Spain during the co-occupancy period (Djakovic, I., Key, A. & Soressi, M. 2022. Optimal linear estimation models predict 1400–2900 years of overlap between Homo sapiens and Neandertals prior to their disappearance from France and northern Spain. Scientific Reports, v. 12, article  15000; DOI: 10.1038/s41598-022-19162-z). Their study is partly an attempt to shed light on the ‘authorship’ of the novel technology. The results suggest that the Châtelperronian (Ch) started around 45 ka and had disappeared by ~40.5 ka, along with the Neanderthals themselves. Early AMH artefacts are known as proto-Aurignacian (PA) and bear some resemblance to those of Châtelperronian provenance. The issue revolves around 3 conceivable scenarios: 1. the earliest AMH migrants brought the PA culture with them that Neanderthals attempted to copy, leading to their Ch tools; 2. Neanderthals independently invented the Ch methodology, which AMH adopted to produce PA artefacts; 3. both cultures arose independently.

Djakovic and colleagues have found that the data suggest that the proto-Aurignacian first appeared in the area at around 42.5 ka. Maps of dated human remains and artefacts for six 400-year time ranges from 43.4 to 39.4 ka show only Neanderthal remains and Châtelperronian artefacts from the earliest range (a in the figure). Two sites with proto-Aurignacian artefacts appears in NW Spain during the next ‘slot’ (b) then grow in numbers (c to e) relative to those of Châtelperronian provenance, which are not present after 40 ka (f) and neither are Neanderthal remains. These data suggest that local Neanderthals may have made the technological breakthrough before the appearance of the AMH proto-Aurignacian culture, which supports scenario 2 but not 1. They also suggest that the sudden appearance of Ch in France and Spain and the abandonment of earlier Neanderthal artefacts known as Mousterian could indicate that the Ch culture may have been introduced by Neanderthals migrating into the area, perhaps from further east where they may have been influenced by the earliest known European AMH in Bulgaria: i.e. tentative support for 1 or 2.

However, well documented as Djakovic et al.’s study is, it considers only 17 sites across only a fraction of Europe and a mere 28 individual artefacts each from Neanderthal and AMH associations (56 altogether). More sites and data are bound to emerge. But the study definitely opens exciting new possibilities for cultural ‘cross fertilisation’ as well as the proven physical exchange of genetic material: the two seem very likely to go hand-in-hand. Seven thousand years (~350 generations) of mutual dependence on the resources of southern Europe surely signifies too that the initially distinct groups did not engage in perpetual conflict or ecological competition, as with small numbers of both one or the other would have been extinguished within a few generations.

 See also: Devlin, H. 2022. Neanderthals and modern humans may have copied each other’s tools. The Guardian, 13 October 2022; Davis, N. 2020. Humans and Neanderthals ‘co-existed in Europe for far longer than thought’. The Guardian, 11 May 2020.

Wider traces of the elusive Denisovans

We know that when anatomically modern humans (AMH) arrived in Asia they shared the landscape with ‘archaic’ humans that had a much longer pedigree. In 2010 an individual’s little-finger bone dated to around 30 to 49 ka old was found in the Denisova Cave in central Siberia (at 50°N). It yielded a full genome that was distinctly different from those of AMH and Neanderthals (see: Other rich hominin pickings; May 2010). Four other fossils found subsequently in the Denisova Cave contained similar DNA. Checking the DNA of living humans and fossil Neanderthal remains revealed that the newly discovered human group had interbred with both. In the case of AMH, segments of Denisovan DNA are found in the genomes of indigenous people living in East and South Asia, Australia, the Pacific Islands and the Americas, at levels of 0.2%, rising to 6% in Melanesian people of Papua-New Guinea. But such introgressions have not been found in Europeans (but see below), suggesting that the Denisovans were restricted to Asia.

There have been suggestions that at least some of the ‘archaic’ human remains found widely and abundantly in China may have been Denisovans; although they might equally be of Homo erectus. But none of the Chinese fossils have been subjected to gene sequencing – those found in caves outside tropical and sub-tropical climates might retain DNA just as well as Neanderthal and even older remains in temperate Europe. Yet a partial lower jaw discovered in a cave on the Tibetan Plateau (at 35°N) did yield proteins that had close affinities to those recovered from Siberian Denisovans. Now similar analyses have been performed on an abnormally large molar found in a cave in Northern Laos, showing that it too is most likely to be from a young (as suggested by its being little worn), possibly female (it lacks male-specific peptides), Denisovan. The locality lies at about 20°N, far to the south of the other two Denisovan sites (Demeter, F. et al. A Middle Pleistocene Denisovan molar from the Annamite Chain of northern Laos. Nature Communications, v. 13, article 2557; DOI: 10.1038/s41467-022-29923-z). Sparse as the evidence is, Denisovans were able to tolerate climate differences across 30 degrees of latitude.

A probable Denisovan molar from 164 to 131 ka old cave sediments in northern Laos. (credit: Demeter, et al.; Fig. 2)

The Wikipedia entry for Denisovans is a mine of additional information. For instance, detailed analysis of the roughly 5% of their genome that indigenous people of New Guinea carry suggests that the two groups may have interbred there as late as 30 ka. Since Both New Guinea and Australia were until 8 thousand years ago part of the Sahul landmass when sea level was low during the last ice age, these inferences add tropical occupancy to the Denisovan range. Does this suggest that Papuans and indigenous Australians migrated with Denisovans, or had the latter crossed the sea from Timor earlier and independently, after moving from Asia by ‘hopping’ from island to island through eastern Indonesia? There is a possibility that Denisovans could even have survived in Sahul until as late as 14.5 ka. Even more odd, modern Icelandic people are unique among Europeans in having detectable traces of Denisovan DNA. However, rather than having been directly shared between Denisovans and ancestral Scandinavians – a possibility – it may have been carried by Neanderthal-Denisovan hybrids migrating westwards from Siberia with whom the Icelanders’ ancestors interbred. There are other interesting points in the Wikipedia entry. One is that the consistently lower Denisovan ancestry in living East Asians compared with people of Oceania, may indicate two separate waves of eastward migration by AMH. The latter may have arrived first, had greater contact with Denisovans and then moved on across seaways to remain isolated from the later migrants.

Finally, something that puzzles me as a non-geneticist. If both Denisovans and Neanderthals died out as genetically distinct groups tens of millennia ago how could the genetic traces of interbreeding with AMH have been retained at such high levels until the present; i.e. through thousands of generations? Each of us carries a 50% deal of genes from our parents. Then with each subsequent generation the proportion is diluted, so that we inherit 25% from grandparents, 12.5 % from great-grandparents and so on. Yet Papuans still have 5 to 6 percent of Denisovan DNA: much the same holds for Europeans’ Neanderthal heritage. Does such a high level of retention of this ancestry suggest that a large proportion of the earliest migrating AMH individuals stemmed from generation to generation interbreeding on a massive scale? Did the ‘newcomers’ and ‘locals’ band eventually together almost completely to merge genetically, or am I missing something … ? Probably