Europe - part III: Into the Holocene
This is the third installment in our series on the European extinctions. The first entry, exploring the ecological background and potential climatic role in the extinctions, can be found here. The second entry, focusing on the arrival, culture and impact of modern humans to the continent, can be found here.
The Dawn of the Holocene
The end of the last Ice Age had enormous repercussions for the landscape of Europe. It has traditionally been posited that there existed during the glacial periods a series of refugia in the southern regions of Europe. Here, the temperate, heat-craving species survived, and later spread out after the end of the glacial. Newer studies, however, have indicated that this picture lacks nuance, and that some trees (particularly hardy, boreal species such as goat willow, Salix caprea, and silver birch, Betula pendula) had potential ranges far greater than hitherto assumed (36). Regardless of the exact tree-cover on the northern steppes, it remains clear, however, that the general picture was one of open habitat, and that for the vast majority of taxa, including the entirety of the temperate flora, the southern reaches of the continent did indeed constitute a vital refuge. This all changed with the onset of the Holocene. The southern flora had made tentative northward thrusts already during the Pleistocene, when conditions allowed it (33), but not until the Holocene did this forward march become permanent.
We must not make the mistake of envisioning this as one event—a sudden transformation at the crossing of a temporal boundary. The onset of the Holocene was a protracted, chaotic, even torturous process. It seemed firmly underway with the coming of the Bølling-Allerød interstadials (12.4-10kya), as the ice retreated, the skies warmed, and southern species, humans among them, rushed into the depopulated north (33). Then, circa 11kya, came suddenly the Younger Dryas, and the ice returned, and the young forests died, and the efforts of those first human settlers came to nothing. This was far from the first cold-snap in the glacial period, yet things had changed. The celestial rhythms that dictated the ebb and flow of the ice ages, the Milankovitch Cycles (8), decreed this to be the start of a new interglacial. Against this, the various gas-cycles and meteorological phenomena of Earth could only delay. Within a millennium, the Younger Dryas ended, and the Holocene proper began (19). The ice was not so quick to relent, however, and even the first few millennia of the Holocene bear witness to repeated glacial advances in the Alps (6). Yet the general trend from the Dryas on was one of consistent warming, however often temporarily punctuated.
Fig 1. Glaciers in the Swiss Alps. Today one of the last remnants of the great continental glaciers of the Pleistocene, their march and retreat during the Early Holocene tells a story of turbulent change.
Terms of use: This image is licensed under a Attribution 2.0 Generic. It is attributed to Trey Ratcliff, and the original can be found here. The image is unedited.
The exact nature of the Early Holocene European landscape has been much debated over the years. The traditional perspective has been that Europe was originally coated more-or-less from coast-to-coast by a dense growth of primeval forest, its thick canopy broken only sporadically by clearings and mountains. In more recent times, an alternative hypothesis has been presented, claiming that Europe was originally primarily an open steppe-or-savanna like landscape, resembling many parts of Africa today (30). This explanation, most often termed the “Vera-hypothesis” after its original proponent, Frans Vera, builds on the idea that dense populations of megafauna would hinder the development of forests, and that as a result, open landscape-conditions would prevail (7). This image has been criticised by certain authors, among them Mitchell, F. J. G. (2004), who shows that concentrations of AP (arboreal pollen) from the early Holocene consistently constituted over 60%, a value heavily connected with dense woodland. Only AP-levels under 50% indicate open landscapes (20). Newer studies, however, have added more support for at least an attenuated form of the Vera-hypothesis—truly dense, closed-canopy woodland seems to have covered no more than 60% of most areas, with the remainder divided between semi-closed forests, savanna and open fields (29). This is a substantially higher percentage of climax forest than either today or during the previous interglacials, yet still far from a uniform blanket. Furthermore, the exact percentage of forest-cover appears to have varied notably from region to region, with Central Europe, for one, having been substantially more densely forested than the Atlantic seaboard (28).
An Age of Man
The advent of the Holocene saw also, more-or-less synchronously, the advent of agriculture in the Levant (2). Though slow in reaching Europe, no technology, save perhaps the discovery of fire, would ultimately have greater consequences for the continent and its denizens. It would be folly to attempt in this article a full exploration of the technological, ethnic, and demographic transformations that occurred across the period. That would be, and is, the matter of multi-volume books. A summary sketch, however, may be provided:
The span between the end of the Pleistocene and the coming of agriculture saw not so much technological advancement—though this did occur—as it did technological and economic change. The world of the Holocene was a very different one from that of the last glacial period, for reasons both climatic and human-caused. Technological changes often correspond to oscillations in climatic and (accordingly) environmental conditions, indicating that the increased stresses and opportunities of changing conditions may have been recurring motivators of innovation and cultural transformation (34). Such changes included everything from the gradual adaptation through Europe of pottery (24), to innovations in the construction of arrowheads, spear-tips and more (22).
When discussing the impacts and advances of human cultures, it is useful to employ archaeological chronologies. Specifically, we see from the Late Pleistocene to Mid-Holocene the process of 3 great archaeological epochs, the Palaeolithic (ending in the European context circa 15kya), the Mesolithic (ending in Europe circa 8kya), and the Neolithic, which ended around 1700 BC, at the dawning of the Bronze Age. A central factor impacting the Palaeolithic-Mesolithic transition was the declining availability of large game. The hunting of vast herds of megafauna had been, for the Pleistocene Europeans, one of if not the single most important source of resources, from food to tools to construction-materials. This changed with the coming of the Holocene. As the pool of large animals gradually thinned, and those species that survived grew less numerous, new sources of food and raw material became necessary. The decline in both the abundance of hunted game and of long-distance hunts is archaeologically traceable, gradually growing more pronounced as the Holocene progressed (22). Mesolithic populations grew increasingly disparate and coast-oriented, with seafood gaining a growing importance. This included in many sites more than 50% of protein-uptake—in some places upwards of 90% (26). By the late Mesolithic, fossils from England evidence an exploitation of shellfish so intensive that it caused notable population-declines, and the virtual disappearance of intertidal molluscs over a certain age (18). This should not of course be taken as meaning that Mesolithic populations lost interest in big game-hunting. Where possible, they still pursued, felled and consumed the remaining species of megafauna. Yet by the end of the Mesolithic, the populations of these animals had declined to such an extent that they quite simply could not constitute the mainstay of most tribes’ economies.
Fig 2. A reconstructed round-house of the early Mesolithic. Like most Mesolithic settlements, it is erected by the coast.
Terms of use: This image is licensed under a Attribution 2.0 Generic. It is attributed to Andrew Curtis, and the original can be found here. The image is unedited.
We may see in this the inklings of a dark spiral for game-populations: It is plausible, even likely, that human hunting was the main instigator of the initial megafaunal collapses, but as their populations declined, conditions only worsened. Their populations reduced, they could no longer maintain through grazing and browsing the open habitats that best suited their species, so that even as their numbers fell through hunting, the environment around them grew ever more unfavourable. This was a disadvantage, not only for them but for human hunters as well, for whom hunting in dense woodland was more difficult than in open country. The result for humans was a lifestyle increasingly centred on the coasts and on fishing, but for the aurochs, elk and tarpan, such was not an option. Furthermore, there enters here the dark horse of demographic stochasticity: all populations are subject to random variation, be it from adverse seasons, outbreaks of disease, or other such factors. Yet where a large and varied population might weather such turbulence—or at worst replace the impacted populations by newcomers from elsewhere—the stakes are much worse for populations which are small, shrinking and isolated. Inbreeding becomes a risk, poor genes may proliferate, and any local disaster becomes potentially existential. Even as man gradually shifted his focus from the great game to sea, the game itself slowly faded into dark refuges and, in many cases, extinction.
It was against this backdrop that we see the introduction of one of the most transformative technologies in European history, both for its people and its landscape: agriculture. At the end of the Mesolithic, the character of the continent was still one primarily marked by dark, climax forests—in places almost unbroken, in others, as we have noted, the dominant element in a mosaic. Over the next few millennia, all of this changed (28) (30). The process of deforestation was not one that occurred all at once, but rather, through a long, protracted decline, which did not reach its zenith until historical times. There were pauses, even reversals, but all were temporary (28). In the Alpine region, the earliest stages of the Neolithic show only small, scattered fields, still islands in a sea of woodland, as evidenced by the continued presence in cereal stores of plants characteristic of forest verges and clearings (30). Over the course of the period, these woodland-signatures decline, and by its end have been mostly replaced by weeds evidencing a firmly open habitat. In the millennia following agriculture, the percentage of Europe covered by temperate woods fell from 70 to 63%. By the early modern period, it had reached 45%, and today it is only 41% (28). The dates of these changes do not correspond neatly across the continent, with some regions experiencing the most dramatic bouts of deforestation as late as the terminal Bronze Age (5). The reasons for this are various, and most probably tied into the general complexity of human culture and demographics. An example of how ethno-political dynamics can directly transform the landscape may be given through one particular prehistoric revolution: The Indo-European invasion of Denmark. This event can be pin-pointed through both genetics, archaeology, and pollen-records (13).
Beginning around the dawn of the 3rd millennium BC, Western Europe, including Jutland, experienced a series of incursions by a new people, hailing ultimately from the eastern Pontic-Caspian steppe. These people, termed in the north-central European context the Corded Ware Culture, were the earliest introducers of the Indo-European languages. They brought with them not merely their language, but also their culture and lifestyle, which in contrast to the preceding Neolithic-farmers was one of nomadic pastoralism, rather than sedentary agriculture. The consequences of this were profound. Today, western Jutland is exceptionally open and sandy compared to the rest of Denmark, dominated in historical times by windswept heathland. The region was always defined by sandy soils—a consequence of ancient glacial advances—but it was not always open and treeless. Even through the Neolithic, the west of Jutland was in fact one of the more densely wooded parts of Denmark, precisely because the poor, sandy ground made it unappealing ground for agriculture. When the Indo-Europeans arrived, however, this changed. The stark, marginal land was inopportune for wheat and barley, but perfect for pastoral herding. Quite remarkably, these Bronze Age invaders proceeded to engage in what can only be described as an act of terraforming, cutting, burning, and grazing the woodlands away on a massive scale, until they had, in western Jutland, effectively recreated the steppe-conditions of their Ukrainian homeland (13). The consequences of this ancient transformation continued to define the landscape until only a century ago.
Fig 3. A shepherd on the great heath of Jutland, by artist Frederik Vermehren. Though today an ancient and iconic landscape, the creation of the vast heath was the product of human actions, of deliberate cutting and burning, carried out by the first Indo-Europeans in Denmark.
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We should not be too swift in registering this decline as a case of unmitigated environmental destruction. As noted earlier, closed-canopy vegetation in the Early Holocene was unusually high compared to earlier interglacials, likely an artifact of the megafaunal extinctions, which resulted in an absence of necessary ecological functions that would otherwise have limited forest-growth. The Early European farmers were not, however, engaging in some charitable act of Pleistocene rewilding, reconstructing the temperate savanna for the benefit of the remnant Eemian megafauna. Although such species as the aurochs and elk might have been expected to benefit from the process of deforestation, in fact they declined throughout the period, and for a simple reason—the coming of agriculture brought also livestock, and the only cattle welcome on the new savanna were those of a domestic pedigree (30). Though it benefited a host of meadow-birds and insects (38) (14), the creation of the European pastoral system heralded less, not more habitat for most of the wild megafauna. It is interesting to note that what was initially a fairly strong south-north gradient of open to closed habitat had been, by around the birth of Christ, firmly replaced by one running west-to-east. During the Atlantic period, circa 7kya, southern Iberia was overwhelmingly the most grass- and heath-dominated landscape in Europe. By the year 0, it was the British isles (9). This appears quite clearly to reflect a shift from an ecologically contingent vegetation-cover to a landscape shaped by the conditions for agriculture and pastoralism.
We must be somewhat cautious in describing the transformations that progressed throughout the Holocene. Nothing in nature is a constant, save change itself. Disentangling which processes were natural, and which were man-driven, is not always entirely simple (3). Temperatures have shifted through the Holocene, and even events such as the rapid decline of elms observed during the Neolithic (3) could, as modern experience shows, be the result of natural pathogens. Nevertheless, the general pattern is clear, often accompanied by strong archaeological evidence, and all-together unprecedented in Earth’s history. Though Man’s influence was felt even in the Late Pleistocene, from the half-way point of the Holocene it became all-pervading. Europe’s landscape became a human landscape.
The Pleistocene Continued?
Having progressed this far in our exploration of the Holocene, it is perhaps time to ask a simple yet fundamental question: does the Holocene exist? That may seem a ridiculous question—of course it does, we have just been discussing it! The dawn of the Holocene is demarcated, neatly, by the retreat of the glaciers and the warming of the planet and the gradual rise and dominance of Man. This is not, per say, wrong, yet it is a narrative which suffers from a critical lack of scope. What we commonly term the ‘Ice Age’ was in truth only the latest cold period in a series of oscillations between glacial advances and retreats, which has been ongoing for over a million years. There were ice ages before the Ice Age, and ‘Holocenes’ before the Holocene. Not accounting for the impacts of manmade climate-change, the current warm period will likely last for a few more millennia before ultimately descending back into another long age of glaciers. Discomforting as the notion may be, this era of relative warmth we have so artificially segregated from the remainder of the Pleistocene is not, in fact, geologically or climatically unique. What then of its extinctions? We have discussed already in the first two parts of this series those species that were lost in the Pleistocene—of the megafauna, the majority. Yet there were losses just before the conventional Holocene-boundary, and, as we will now see, losses just and long after.
It is a fascinating point that many species typically counted as ‘Pleistocene’ did in fact survive, even locally in Europe, well into the Holocene. This includes such famous taxa as the enormous Irish elk (Megaloceros giganteus), potentially the woolly rhinoceros (Coelodonta antiquitatis), and definitely the woolly mammoth (Mammuthus primigenius) (16) (17). Megaloceros remains dated to circa 7,5 ka are known from as far west as Maloarkhangelsk, near the Ukrainian border, and mammoths persisted in European Russia and the Baltics as late as 9,8 ka. Dholes, the so-called ‘whistling dogs’, appear to have survived into at least Iberia until the early Holocene (27). The Holocene record of lions in Europe is relatively well-known, yet taxonomically confusing, owing to an uncertainty regarding whether finds pertain to late-surviving cave lions or (as seems more likely), a dramatic Early-Holocene colonisation-event by Asiatic lions out of Anatolia (32). At any rate, they were present in Mesolithic Spain and Italy, with remains dated respectively 7-8 and 5-7 BC, as well as Hungary and Ukraine, where they persisted into the Bronze Age, circa 3000 BC. The collapse of their range occurred gradually throughout the Holocene, running first from the west to the east, then the north to south. By the 1st millennia BC, they were restricted to the southern Balkans, and by the birth of Christ, the last holdout of European lions was in the mountains of northern Greece (32).
Among the most adaptable of palearctic animals, the Eurasian elk (Alces alces) ranges today from dense boreal forests to parkland tundra, from mountain slopes to boggy moors and even into farmland (11). Though restricted to the colder reaches of the world, their resilience within these habitats is remarkable—during the Pleistocene, they reached as far north as within 350 km of the ice-front, to the very limits of the forest steppe (31). It is therefore not surprising that they were among the very first large mammals to recolonise Northern Europe following the glacial retreat, becoming by the preboreal (10.3-9kya) the most abundant large ungulate in the Baltics (31). Yet by the Boreal period (8–5.7kya), elk had begun to decline in the now-isolated Britain, vanishing shortly after. They remained common in the Alpine region into the Mesolithic and Neolithic, but were declining by the latter. (31). By the dawn of the Bronze Age, they had likely already vanished from southwestern France, and by the 1st century BC, the animals are reported by Julius Caesar as occurring in Germania, but not in Gaul (31).
A similar picture can be painted for the wisent, or European bison (Bison bonasus). They are confidently recorded from much of northern and eastern Europe during the early Holocene, as far north as Östergötland in Sweden and northern Belarus (4). Whether they ever occurred in Britain is unclear, complicated mostly by the generally uncertain origins of the wisent prior to the Holocene. The earliest skeletons identified as B. bonasus date to the Early Holocene, and the species is thought to have emerged from the complex of steppe bison-populations (Bison priscus) that spanned the Late-Pleistocene Palearctic (4). Steppe bison are indeed known from Britain during the Late Pleistocene, so the question of whether or not ‘wisents’ proper were ever present in the region may perhaps be regarded as ultimately arbitrary (1). They had, at any rate, disappeared from the area by the Early Holocene. By 7000 years ago, they have disappeared from southern Scandinavia, and by 3000 seemingly from France. In the north-east, they famously persist for far longer, constituting in the Polish-Lithuanian region up to 20% of ungulate bones even into the Late Pleistocene. Though unknown from Italy, there exists also a solid if confusing record of wisents from the Balkan peninsula, as far south as the mountains of Thrace (23) (12). Here, they were known as Paeonian oxen, as well as bubalus, bisones, bonasus and, in the tongue of the Paeonians, monapos (12). The confusion stems from the fact that Ancient (and Medieval) authors frequently failed to distinguish adequately between aurochsen and bison, using the names of the two species interchangeably. Nevertheless, a combination of in places quite specific descriptions, as well as archaeological evidence, makes it clear that wisents did indeed range at one time all the way to the borders of Greece, persisting perhaps in the southeastern Balkans as late as the 7th century AD (23). By this time, however, they had already been rare in the region for millennia, as were they in Central Europe, where by the Mid-Late Holocene, aurochs-bones outnumber those of wisents 1:9 (4). Only in northeastern Europe and parts of the Caucasus were they locally abundant into the Middle ages, when at last these final holdouts began to collapse, so that by the dawn of the Modern period the species was rapidly nearing extinction.
Fig 4. A group of hunters find themselves caught in a bison-stampede of their own making. The scene takes place in ancient Thrace, circa 200 BC, very near the site of modern bison-reintroductions carried out by Rewilding Europe.
Terms of use: Artwork by Hodari Nundu and Commissioned by The Extinctions
We have focused thus far on large mammals, and indeed they were both the most numerous and significant victims of Holocene extinctions, yet not the only. Europe’s avifauna was resilient, but even here do we find losses. The last Great Auks (Pinguinus impennis) famously vanished only a scant few centuries ago, in the 19th century, yet by then they had already been confined to the scattered isles of the North Atlantic for millennia. This, however, was already a mere relic of their former distribution, and as recently as 3000 years ago, great auks could still be found along the coasts of the Netherlands. During the Mesolithic, they nested along the northern and eastern coasts of Denmark, and their distribution in the early Holocene spanned the North Sea (15). Dalmatian pelicans (Pelecanus crispus), today confined to Southeastern Europe, were widespread in the North Sea region until the Mid-Holocene, circa 5000 years ago, and Roman authors as recently as 50 AD describe pelicans living in the Low Countries (21). Another total extinction is witnessed just on the cusp of historical times, as the large crane Grus primigenia survived in north-western Europe into the Bronze Age (10).
A full survey of every Holocene extinction is beyond the scope of this article, but we could continue. Wild horses, for instance, maintained a broad distribution across North- and Central Europe during the early Holocene but were already then uncommon. Over the next millennia they declined, so that by the early Atlantic period, circa 7000 years ago, they had nearly vanished, occurring only in a few scattered and evidently isolated sites in Iberia, France, and northern Europe (35). The end of the Atlantic period and the coming of agriculture gave the species a second wind, as woodland clearances once again created grassland-conditions under which the species could thrive. That wild horses should have flourished in this period while the decline of the aurochs accelerated is an interesting point, and one which seems plausibly addressed by the presence, at this time, of domestic cattle, but not domestic horses. The aurochs competed with the first farmers’ livestock in a way wild horse as of yet did not. This is an important reminder of a point raised earlier in this article, that in dealing with humans and their impact on nature, cultural and economic dynamics matter just as much as environmental constraints. The advent of a new technology, the spread of a new people, the domestication of a new animal—all of these can produce sudden and drastic consequences for the natural world, not easily explicable if explored from a purely climatological and/or environmental lens.
We must end the accounts of species here, for the sake of brevity. Not even discussed are the Holocene island-extinctions, such as those that impacted Sardinia and Cyprus (37), nor the regional collapse of many marine populations of for instance walrus in historical times. The overall picture should by now be evident—extinctions, megafaunal and otherwise, were staggered throughout the Holocene. It is indeed true that there occurred between the Preboreal and the dawn of written history, that is, from circa 10,000-3000 BP, something of a lull in their rate, both locally and continent-wide. It is not true that this lull constituted a full pause, such that those losses which occurred in the Late Pleistocene, and those that happened in historical times can be neatly delineated from each other. Within this interval we see the final of the Irish elk, the loss from Britain and the south of Europe of the extant elk, and the collapse of lions in all of Europe save Greece. Bison vanished from Scandinavia during the early Holocene and were in evident steep decline through most of their range already by the Mid-Holocene. Aurochs seem generally to have been a more adaptable species, remaining widespread until the early Bronze Age, when growing human populations and intensifying land-use seems finally to have prompted widespread range-contractions (39). The Holocene saw the loss of seabirds, large grazers, apex predators and island dwarfs. Its course was not, perhaps, as defined by the overflood of extinctions as was the end-Pleistocene, but it was marred by them, and at a rate far above the usual. Like the Late Pleistocene, but unlike the entire Cenozoic before it, the Holocene losses saw no replacement, only vacancies, save for those general niches filled by our own domestic livestock, and the workings of our axes and ards.
Modernity
We come now to the final stage of our narrative, and with it, to the present day, and the years immediately preceding it. The Neolithic gave way to the Bronze Age, and the Bronze Age to the Iron. With the ending of the latter ended also prehistory as traditionally delineated, but the Holocene continued. Extinction continued. It is often said that we live in an age of extinctions, a statement which it is not our intention to dispute. What has hopefully been provided, however, is a measure of perspective. Over the course of this three-part series, we have traced the background, course and causes of the European extinctions, from first vanishings of the southern megafauna, 50,000 years ago, to the final vanishing of the great auk, less than 200 years ago.
We will not here attempt a deep exploration of the modern extinctions, though they pertain to the age of the Holocene. That is the material of another article. Of several. In part, this is due to our improved records, letting us follow more closely the exact causes and quantities of recent losses. In part, it is because the partial lull of extinctions seen through much of the Holocene has ended. Dramatic and consequential as the loss of the mammoth and the sabretooth was, the number of species now threatened is larger still by an order of magnitude. Slowly, persistently, the process of extinction has worked its way down the chain of life, starting with the spear and the lion, ending with pesticide and the meadow-bee. In time, we will explore the transformations of the last few centuries in far greater detail, the burgeoning of the human population, the advent of industry and the green revolution. For now, we have endeavoured to trace a line from the first coming of Man to his eventual dominance, from the initial vanishings in the European south, to the vast declines of species, great and small, that have swept and are still sweeping the continent. The phenomenon of extinction is not new, but it is forever.
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