Europe - Part II: The Human Dimension
So long as there has existed within our species a divide between the settled and the nomad, the farmer and the hunter, “civilised” and “primitive”, there has been the concept of the noble savage. At one with nature, an extension of his environment, the ideal of the earth-born man, living as one with his surroundings, is an ancient one. Yet what the idea has in potency, it lacks, perhaps regrettably, in reality. Only in more recent times have we begun to grasp the extent to which even small, uncomplicated societies have shaped their surroundings. From artificially fanning wildfires (22) to the decimation of island ecosystems (7), humans have been a catalyst for ecosystem change for millennia. From this realisation arises an obvious question: how long has this been the case? When did humanity first make the leap from peculiar ape to demiurgic agent, and, to reach the topic of this article, what role, if any, did humanity have to play in the extinctions of the ancient past? In the previous entry in this series, we laid the groundwork for our exploration of the megafaunal extinctions and surveyed the available evidence for a climatic cause. Now, we shift our focus, from geology and climatology, to archaeology.
The evidence for human hunting
A traditional argument against a human cause for the Pleistocene extinctions has been the apparent paucity of archaeological evidence. Two questions therefore must be addressed—the first, what is our current evidence, and the second, what sort of evidence should we expect?
The most well-known formulation of a human-hunting-hypothesis is Paul S. Martin’s “Overkill” or “Blitzkrieg” hypothesis, which explains the megafaunal disappearances by envisioning rapid waves of extinction following human arrival new continents (Martin. P.S. 1973). Donald K. Grayson & David J. Meltzer (2003) provided a well-known rebuttal to Martin’s Overkill Hypothesis, stating that a perceived paucity of archaeological evidence of human exploitation discredited Martin’s claims. This was followed up by the same authors in 2015 (16) and 2020 (17). Their specific case is based on North America, but generally relevant to Europe as well. Two key points may be extracted: The first, that evidence for kill-sites and related artifacts is scant for mammoths (Mammuthus primigenius/columbi) and mastodonts (Mammut americanum), and entirely lacking for the remaining megafauna. For this reason, any assertion that hunting was the cause of the extinctions is pure speculation, with no empirical foundation. The second key claim is that many extinct genera have not been dated to human arrival, disappearing from the fossil record millennia before the earliest estimates of human arrival. Both of these points, along with the rest of their claims, have been contested (9), and a complete analysis would require a substantial detour into North American material. Nevertheless, they are reasonable considerations in any context, and worth applying to Europe.
Figure 1. Paul S. Martin (1928-2010). American geoscientist and palaeontologist, and originator of the overkill-hypothesis.
Terms of use: This image is licensed under a Creative Commons Attribution 2.5 Generic license. It is attributed to Thomas R. van Devender. The image is unedited and the original can be found here
Whatever the paucity of evidence of hunting in North America, perceived or otherwise, the situation is somewhat different in Europe. There is substantial evidence of human exploitation of large herbivores during the ice age. Bosch et al. (2011) shows strong evidence for human-connected accumulations of megafauna-bones in the Gravettian-culture tradition of the Upper Palaeolithic (50,000-10,000 years ago). Here, a bone-deposit, located near a campsite and containing mostly mammoth, but also woolly rhinoceros (Coelodonta antiquitatis), horse (Equus ferus ferus) and more, shows butchery marks on the remains. Scavenging seems ruled out, since the total lack of animal gnaw-marks indicates exclusively human access to the animals. It has been suggested that growing populations may have driven people to hunt larger, more difficult game(1). Meanwhile, several studies provide evidence of exploitation of proboscideans by Neanderthals in the Middle Pleistocene(19)(26).
Pleistocene humans often focused intensively on particular species. This can be seen in Meiendorf, Germany, where more than 99% of mammal-remains were from reindeer (Rangifer tarandus ) (2). In that case, the reindeer enjoyed favourable climatic conditions at the time, and their populations could be replenished. It is not hard to see, however, how a similar intensity of exploitation might prove fatal for a more vulnerable and isolated population, such as those of the temperate faunas. The evidence provided by Bosch et al. (2011) in particular seems to rule out a reduction of human-megafauna interactions to mere scavenging. It does, nonetheless, remain true that the quantity of confirmed kill-sites is not overwhelming. Based on these remains alone, it would be overbold to claim overkill as the only, obvious, explanation. That humans hunted large animals in Europe is difficult to deny, but how intensive was this hunting?
A key question which must be answered before this can be determined is what sort of evidence we should expect. How many kill sites should we expect to find, assuming a primarily human driver of extinctions. Martin already addressed the low number of kill-sites (8), noting that a brief interval between human colonisation and extinction would necessarily imply a paucity of surviving sites, due to low preservation rates. Though stated in the context of North America, this seems readily applicable especially to the temperate refugias in southern Europe, where most of the interglacial, Eemian fauna had become restricted to a few isolated areas supporting temperate conditions. Though this situation had occurred numerous times before, nearly the entire temperate fauna suddenly vanishes circa 50,000ky BP., very close to the estimated date of human arrival (3). Not only did this extinction occur very quickly, but the species affected were already restricted to a limited geographical areas, and presumably enduring in relatively small populations. Thus, we would not expect to find many, if any preserved remains (8). Indeed, the fact that we find so little evidence of interaction between humans and the refugia-fauna heavily indicates that said fauna either went extinct at, or just prior to human arrival, but not long after. A period of prolonged coexistence would be expected to produce evidence of exploitation, even if merely low intensity, as it has elsewhere. That these species should endure 9 consecutive glacial-interglacial cycles over a million years, each time seeing their ranges massively contract, expand, then contract again, only to suddenly and mysteriously die out within a couple thousand years of human arrival seems quite the extraordinary coincidence.
Fig 2. A group of human hunters, having tracked and brought down a large woolly rhinoceros (Coelodonta antiquitatis), must now defend it from a pack of hungry hyenas (Crocuta crocuta). On the far left hand is an early domestic dog or proto-dog (Canis lupus familiaris). Scene set circa 20kya, during the Last Glacial Maximum.
Terms of use: Artwork by Hodari Nundu and Commissioned by The Extinctions
Karen Lupo (2016) discusses the cost/benefit analysis of hunting large game, and concludes, that the hunting of megafauna is rarely justifiable in terms of energy expenditure vs energy gain. On this basis, she suggests that interpretations of prehistoric peoples as megafauna-specialists may be mistaken. It is known, however, that early humans undoubtedly made substantial use of proboscidean remains. Pleistocene cultures throughout Europe used mammoth bones to construct dwellings (13), while members of at least the Acheulian culture used elephant bones for the construction of handaxes (27).
Even beyond these, one may readily conceive of other, non-meat-related reasons early humans may have had for hunting mammoths, from cultural/spiritual to the use of mammoth-wool as clothing. Another paper (18), studying the Yana Late Pleistocene site in Siberia, finds only sporadic hunting of mammoths by the Yana. The same authors, however, note in a follow-up study (20), that the hunting here is primarily for the gathering of ivory, not meat. It could be argued, then, that even if Karen Lupo (2016) is correct in regards to the cost/benefit analysis, that would not necessarily preclude a heavy focus on megafauna-hunting for other reasons.
Whether it is true that hunting megafauna for meat is unprofitable is itself debatable, however. Karen Lupo (2016) documents that elephant-hunting in the modern Congo-Basin is the exclusive domain of specialised men called tûmas. These are relatively few, making out only 4 of every 12-15 hunter, and though prestigious, are clearly not the norm of their culture. Yet it is highly questionable whether the conditions of modern hunters in the tropical Congo are comparable to those of Pleistocene peoples in glacial Europe. The Congo is a rich environment, in which alternative food-sources are abundant, the need for heating and clothes is minimal, and building-materials are omnipresent. Such was not the case on the desolate, treeless steppe, where vast herds of megafauna would have constituted the bulk of the region’s biomass.
First and Last Men
A question that may arise, in considering our prior references to megafauna-exploitation by Neanderthals and their ancestors, is why they nonetheless failed to cause any comparable extinctions. The first, and simplest, answer is to point out that they were hardly as efficient hunters as Cro-Magnons (5). What particular quality in modern humans distinguished us from other animals has been a subject of enduring debate across science and philosophy, but it may be ascertained that it was absent in Neanderthals. Indeed, as victims of the human expansion, an argument might be put forward for viewing them (in an ecological context), not so much as precursors of humanity, but instead as members of the megafauna, lost alongside the sabretooth and mammoth. This view seems backed up by recent research (28), which finds that Neanderthal extinction occurred rapidly between 50-35ky. This was apparently unconnected to any dramatic climatic events, yet coincides well with the continent’s first settlement by modern humans and the extinction of the southern megafauna. We must not, however, go too far in de-humanising the Neanderthals; though not fully ‘human’ in the sense we now know it, they were nevertheless capable of abstract, symbolical thinking (39), and had in fact been increasing in technological complexity right up until their sudden demise (40). Although we may consider them, in a palaeoecological context, as constituents of the broader European megafauna, it seems very likely that the extinction of the Neanderthals had, at the time and on the ground, the character of a war more so than any sort of hunt.
And yet, that undeniable, if ineffable, difference of quality between Homo sapiens and Homo neanderthalensis remains. Overwhelming evidence indicates that the first populations of Cro-Magnon settlers in Europe were up to 10 times larger than those of the Neanderthals they replaced (French, J. F. 2015). During the Aurignacian period, the population of Neanderthals in Southwestern France may have been between 80-1300 individuals, whilst the population of early Cro-Magnons ranged from 800 to over 12,000 (42). Despite low levels of interbreeding (41), the contest of the two species was not even. This is not to imply that Neanderthals are necessarily irrelevant in the conversation. The co-evolution hypothesis has been put forward as an explanation as to why more megafauna seems to have survived in regions with long histories of hominid presence. It is a subject warranting a deeper exploration in its own article. The point is rather that their impact cannot be equated with that of later humans. The Neanderthals had evolved side-by-side with the continent’s megafauna, were less efficient hunters than their eventual replacements, and subsisted in far smaller populations than our earliest ancestors. They were not alien to the old megafauna, but part of it, and their fate was one and the same.
It may be noted, furthermore, that the conventional view of ‘stone age humans’, even as pertains to our own species, is hardly an accurate one. Time and again, the image seen in portrayals of the Pleistocene is one of small bands of fur-clad hunters, aggressively jabbing at large animals with pointed sticks. While Pleistocene hunters did indeed use spears, and furs, these were not the only tools at their disposal, and the furs were hardly mere rags. Our earliest evidence of bows and arrows anywhere in the world comes from Africa, and goes back more than 50,000 years (29). In Europe, the earliest known remains, from Mannheim-Vogelstang in Germany, date to circa 17.5-18kya. That this should represent the actual period when bows were first adopted in Europe seems unlikely, and owing to the nature of artifact-preservation, it is entirely possible that the very first hunters in Europe were already able archers. Similar considerations must be made regarding another, greatly underappreciated tool—the spear-thrower, also called the atlatl. Able to greatly enhance the distance and velocity of the dart or spear, our earliest direct evidence for this weapon comes from Dordogne, France, dating to 17.5kya (29). The fact that the weapon was seemingly in use in Australia at roughly the same time, as well as circumstantial evidence from that continent dating back 42ky, points, however, to a far more ancient origin for the atlatl. Even the blowgun might conceivably have been employed, being known from both Asia and the New World, including into the temperate reaches of North America (30). No reports of such weapons are known from Europe, but preservation even in the regions they are known from is exceedingly scanty. Even if present, they would hardly have been aids in the downing of giant megafauna, yet they give an added nuance to the diversity of tools that may have been available to our Pleistocene ancestors.
As regards weaponry and hunting-aids, a point distinctive, yet necessary to discuss, is that of the domestic dog. The earliest remains confidently identified as domesticated date back to circa 12k BC, and hail from Bonn-Oberkassel, Germany (36). Recent palaeogenomic evidence, however, indicates that domestication began as far back as 23kya, in Siberia, from where dogs later dispersed both east and west, entering Europe an indeterminate time later, and crossing with the first human settlers into the Americas (371). Tentative remains from various parts of Europe may push this date all the way back to 30kya, but these are controversial, and may in any case represent “proto-domestications” or failed attempts (38). It can scarcely be denied that the advent of domestication, of which dogs were the first product, would have had a profound impact on the way in which humans interacted with their environment, both in what they could derive from it, and how they could control it. The dog, as an implement in hunting, would have only solidified our ancestors’ position as the hunters par excellence. If any Neanderthals met them, however, it was the very first dogs greeting the very last of their kind. The dating does not appear to overlap, and so though significant in the history of our species, the dog does not offer an explanation for the collapse of Homo neanderthalensis.
Figure 3. Painting of a hand from the Cave of Aurignac in southwestern France. Paintings and objects from the caverns date to upwards of 45,000 BP, making them pieces of the oldest culture attributed to modern humans in Western Europe.
Terms of use: This image is licensed under a creative commons attribution-ShareAlike 3.0 Unported. It is attributed to Daniel Villafruela. The image is unedited and the original can be found here
There exists a general underestimation of the diversity and sophistication of non-agricultural peoples, past and present. Though nomadic hunters, both modern and prehistoric, are often considered as dwelling in tiny bands of a few dozen at most, camps and villages of 100 to even upwards of 250 individuals are known from multiple peoples across the world. Seasonal gatherings among some groups can last months and reach numbers in the thousands (43). Even more dramatic are the societies of the settled foragers, documented from across the world, whose societies can number many thousands, spanning at times vast geographical areas (44) (43). Capable of everything from unitary monarchies to complex secret societies, historical settled foragers have constructed wooden houses, monumental structures and even equipped themselves with sophisticated shields and armour (32) (44) (43) (31). Settled and nomads alike, hunter-gatherers are known to have engaged in large-scale landscape-alteration through artificial wildfires (33) (34), and though data from Iberia at least indicates the arrival of modern humans here did not precipitate a detectable increase in wildfires, this does not preclude the alteration of fire-regimes at a local level, nor speak for the remainder of the continent (35). A deeper exploration into these various points would necessitate a tangent beyond the scope of the article, but the point is straightforward: Palaeolithic humans were almost certainly both more technologically sophisticated, socially organised and overall more numerous than is often conceived. The implications for this as goes the Pleistocene megafauna are clear.
The End-glacial extinctions
Having addressed the evidence of human hunting, and the strong if circumstantial case for a human driver of the temperate fauna’s extinction, it must nevertheless be noted that the remaining European megafauna survived considerably longer. The ice age fauna coexisted with modern humans for circa 30,000 years, only to quickly go extinct come the dawn of the Holocene. From this follows an intuitive narrative: The refugia-species were indeed lost due to human hunting at a time of contracted ranges and inopportune conditions, but the remainder of Europe’s megafauna was a casualty of the climate. This is, at face value, a nuanced assessment. There is overwhelming evidence that climatic fluctuations do indeed raise extinction rates (24), and the prolonged period of human-megafauna coexistence in Europe seemingly precludes any sort of “Blitzkrieg” scenario. Furthermore, the coincidence of the decline of at least some members of the glacial megafauna with the BAIC and other end-Pleistocene warming events seems solid (15).
Unfortunately, there are several critical issues raised by this explanation. If the end of the Weichsel had such catastrophic ecological effects, why not the end of the Saalian, or any of the preceding 8 glacials? Why not the onset of the Weischel-glaciation, which would surely have been a much more ecologically destructive event than its ending? Vast amounts of previous habitat, even of northern steppe-species such as the mammoths and saiga antelopes, would certainly have been destroyed by the onset of the glaciation. It is an unfortunate truth that very few, if any, of the studies arriving at a climate-driven extinction include this broader picture.
Nevertheless, the continued survival of the ice age fauna from human arrival until the end-Pleistocene is a valid criticism against the human-hypothesis. Indeed, let us freely admit that it seems entirely certain the ending of the glacial, and the warming climate, were the primary reasons why extinctions occurred when they did. The correlation of the two events seems too thorough to deny. Yet this only reinforces the aforementioned question as to why none of the previous glaciations and/or deglaciations had any comparable effect. There may, however, be an answer which reconciles both points: Human distribution. Due to the harsh climates, vast ice-free regions in northern Europe and Asia were entirely, or almost entirely free of humans during the entirety of the Pleistocene (23). Even the great plain of Doggerland, the largest extension of steppe in ice-age Europe, was home to no more than 1-4 humans/100 square km . Several long periods saw most of the plain entirely abandoned by human hunters when the climate deteriorated near the Glacial Maximum. (23). The reason for the failure of early European hunters to wipe out the ice-age fauna between their arrival and the end-Pleistocene is thus given a pleasantly concise answer: They were simply absent from most of the fauna’s range. Even where humans did coexist with, and hunt, the steppe megafauna, such as in the relatively warm stretch of France and Italy between the Pyrenees and the Alps, any thinning of local prey populations could simply be reenforced by migrants from the vast, untouched lands to the north.
Figure 3. Europe during the end-glacial, after the retreat of the ice from the Doggerland plain and the formation of the Ancylus Lake. Vast swathes of the continent were still virtually uninhabited by humans until the very end of the Pleistocene.
Terms of use: This image is licensed under a Creative Commons Attribution-ShareAlike 4.0 International. It is attributed to Stoolddog13. The image is unedited and the original can be found here
This also explains, perhaps, the seemingly greater extinctions in the temperate European megafauna compared to its contemporaries in East-Asia. To this day, there persist large felines such leopards (10) and tigers (12) as far north as Korea and Siberia, whilst rhinoceroses survived as far north as Sichuan into historical times (21). Even an Asian relative of the European elephant—Palaeoloxodon namidicus, may have survived in China until only 3000 years ago according to one study (11). This discrepancy is somewhat odd, assuming a climatic cause in Europe, but quite readily explainable in light of a hunting-explanation. Whereas the temperate refugia in Europe were small and isolated, the East-Asian mainland is a contiguous landmass, allowing all of South Asia as an accessible refuge for the interglacial fauna. This is not to imply that no, or even few extinctions took place—that would be incorrect (25)—nor to begin a detour. Digressions can be didactic, but one must be careful not to conflate them for the point. It may suffice to say that mobility and ecological flexibility are both crucial mitigators of extinction-risk (24), and that such factors were poor in the European refugias, but rich—for the duration of the glacial—in the north.
Conclusion
The evidence for human overkill is generally circumstantial, but nevertheless significant. Though sceptical authors are correct in pointing out the low numbers of kill-sites, it is far from clear to what degree this is an issue. As Haynes G. (2007) points out, it can be nearly impossible to find remains from even present-day hunts between common prey and predators, and taphonomic factors mean that most sites are inevitably lost. Humans made heavy use of neighbouring megafauna on the mammoth-steppe, and it seems evident that this exploitation was at least in part active, not merely the product of scavenging. That such hunting-dynamics are different in modern hunter-gatherer societies is interesting, but perhaps not particularly meaningful: the modern Congo and glacial Europe are two very different places, and the necessities (and cultural factors) in the one are bound to differ from the other. Furthermore, the chronology of events betrays a noticeable overlap of human arrival and the first wave of extinctions. Even accounting for uncertainties in the dating, the coincidence is striking. The continuing survival of the glacial fauna for another 40,000 years, though an initially imposing issue, becomes easily resolved, even inevitable once the sparsity of human settlement in the north is accounted for.
In conclusion, it is probably impossible to prove with complete certainty a causal link between human activity and the megafaunal extinctions. Nevertheless, both through the coincidence of events and the total failure of competing hypotheses, it seems a strong and reasonable position in the face of the evidence. In the next part of this series, we will continue the narrative of the European extinctions, past the retreat of the ice and the heating of the world. Into the Holocene.
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