The Lost Rhinos of Europe
Introduction
When it comes to ‘charismatic megafauna’, few animals more thoroughly exemplify the concept than the rhinoceroses. In Africa, they number among the renowned Big Five, in Asia they are recorded already in the seals of the Indus Valley civilisation (Konasukawa, A. & Koiso, M. 2018), and across the world there are few groups that can rival them for fame or recognisability. From the Western perspective, a core part of this appeal has always been of an exoticising sort—the rhinoceros is the perfect fanciful, foreign animal, so simultaneously bizarre yet impressive as to form an ideal fixture for any bestiary or menagerie. That is to say, the rhinoceros grew so famed in Europe, so familiar, paradoxically precisely because it was not found there. And yet, this has not always been the case.
The woolly rhinoceros (Coelodonta antiquitatis) is a well-known creature today, immortalised, in the proper heroic fashion, long after its demise. It has become emblematic of the ice age world, an icon alongside the mammoth and the saber cat, if not, perhaps, quite on their level. Yet if the woolly rhino is, perhaps, a somewhat second-rate figure in the ice age world, what is even less known, indeed almost forgotten, is its relatives—the other rhinos of ancient Europe. Some were even stranger, others more abundant, and all were crucial elements in any proper understanding of the continent’s original nature. All of these species are fascinating, and all worth understanding, wherein the object of this article.
Origins and Affinities
The family Rhinocerotidae has a venerable history, counting over 100 attested species over 50 million years and across Africa, Eurasia, North, and Central America (Liu et al. 2021). Since their Eocene origins—most probably in Asia, though possibly in North America (Bai et al. 2020)—rhinoceroses have come in a myriad shapes and niches, including some of the largest land-mammals to have ever existed (Deng et al. 2021). Despite this, the earliest known European rhinoceros is not attested until the early Miocene, in the form of the cursorial species Lartetotherium sansaniense (Pandolfi et al. 2022). Rhinoceros diversity peaked around to the Miocene-Pliocene boundary, only to crash dramatically in the ensuing cooling, and the slow lead-up to the glacial cycles. Some subfamilies, such as the long-lasting Aceratheriines, were lost entirely, whilst others endured, but at a reduced diversity (Pandolfi et al. 2022) (Liu et al. 2021). In North America, rhinoceroses vanished entirely, the last species attested from the early Pliocene, thus ending an evolutionary history leading back to the Eocene (Wang, B. 2016).
Even so, their distribution remained wide across Afro-Eurasia all the way through the Pliocene and Pleistocene. The Pliocene species Coelodonta thibetana, whose origins on the high Tibetan plateau may have given it some advantage in the cooling world, radiated throughout Eurasia as the glaciations took hold, with a succession of descendant species finally giving rise in eastern Europe to Coelodonta antiquitatis, the woolly rhinoceros, which would become the cold climate rhino par excellence (Yuan et al. 2023). By the Late Pleistocene its range covered nearly the entire Eurasian steppe biome, from its westward extremities in ancient Doggerland to the far reaches of Siberia. It never, however, managed the leap into the Americas, where its ancestors may once have originated.
Figure 1. Artist’s reconstruction of the Late Pleistocene woolly rhinoceros, Coelodonta antiquitatis.
Terms of use: This image is licensed under a Creative Commons Attribution-ShareAlike 4.0 International. It is attributed to Benjamin Langlois. The image is unedited and the original can be found here.
In Europe, the rhinoceros fauna came from the Early Pleistocene to be shaped by two genera—Stephanorhinus and Elasmotherium. These alone would populate the continent until the entry of Coelodonta towards the latter end of the period. The elasmotheres were a strange and deeply genetically divergent linage of enormous, one-horned rhinoceroses, whose subfamily, the Elasmotheriinae, had split from all other living rhinos already by the end of the Eocene (Kosintsev et al. 2018). Throughout the Pleistocene they only skirted the edges of the European continent, spreading at times south through Ukraine into the regions above Crimea, but persisting by the Late Pleistocene only in European Russia (Stefaniak et al. 2023). The final species, Elasmotherium sibiricum, was once thought to have vanished already circa 200kya, but is now known to have persisted until only 39kya, if not later, thus rendering it a part of the general megafaunal extinctions which swept the terminal Pleistocene (Kosintsev et al. 2018).
Far more important in the European context was the genus Stephanorhinus, which by the end of the Pleistocene would furnish two of the continent’s 4 species of rhinoceros, and the only two widely distributed throughout the temperate zone. These were more “conventional” rhinoceroses, similar in morphology to surviving species, but of very confused taxonomy. Any discussion of the genus is complicated by Stephanorhinus’ uncertain nature, with most studies until recently finding it to be paraphyletic (meaning that it fails to include all descendants of its last common ancestor) (Pandolfi, J. 2023b). Two species, Stephanorhinus jeanvireti and S. etruscus, are recorded from the Early Pleistocene, and may have emerged from the Pliocene genus Pliorhinus, which otherwise disappears around the epoch boundary. Though related, the exact affinities of S. jeanvireti and S. etruscus vis a vis each other and the remainder of the genus is murky. S. etruscus was the more long-lasting of the two, surviving into the Middle Pleistocene, where it was replaced by yet another relative, S. hundsheimensis, possibly arriving from the east (Pandolfi, L. 2023a). By the Late Pleistocene, the European population of S. hundsheimensis, or a closely related one, had in turn given rise to two species: S. hemitoechus, the narrow-nosed rhinoceros, and S. kirchbergensis, the Merck’s rhinoceros (Pandolfi, L. 2023b) (Stefaniak et al. 2023). Though the newest phylogenetic data recovers both of these Late Pleistocene species as each other’s closest relatives, this has long been a topic of confusion, not helped by the fact that the Merck’s rhinoceros seemingly only appears in Europe relatively late, with its earliest fossil evidence from Early Pleistocene China (Pandolfi, L. 2023b). The narrow-nosed rhinoceros, meanwhile, is first attested somewhat later from the Mediterranean region. Molecular evidence indicates that the closest extant relative both Stephanorhinus and Coelodonta is the Sumatran rhino, Dicerorhinus sumatrensis (Pandolfi, L. 2023b).
Figure 2. Skull of Stephanorhinus etruscus, kept at the Musee d'Histoire Naturelle, Paris.
Terms of use: This image is licensed under a creative commons attribution-ShareAlike 3.0 Unported. It is attributed to user Ghedoghedo. The image is unedited and the original can be found here
Thus, by the final millennia of the Pleistocene, of the nine remaining species of rhinoceros in the world, four were found in Europe—the giant elasmothere, E. sibiricum, the woolly rhinoceros, C. antiquitatis, and the two species of Stephanorhinus, the Merck’s and narrow-nosed rhinoceroses. Europe, then, was as much a hotspot of rhinoceros diversity as Africa or Asia.
Rhinos of the Wood and Pasture
Though the woolly rhino is the most famous extinct rhinoceros of Europe—and the only species most people are liable to know—it was not the continent’s most characteristic one, where temperate conditions prevailed. Had one visited nearly any part of Europe during the last interglacial, it is the two species of the genus Stephanorhinus, the Merck’s and narrow-nosed rhinoceroses, that our theoretical time-traveller would have been overwhelmingly most likely to encounter, and it is these two species that would likely populate the continent today, had they not been lost in the extinctions that swept the Late Pleistocene and early Holocene.
The Merck’s rhinoceros was the largest known species of Stephanorhinus, greater in size both than its narrow-nosed contemporary, as well as all earlier recorded relatives (Stefaniak et al. 2023). A tall, long-legged animal with a high head-posture, this, together with its dentition, as well as isotope signatures, indicates strongly a diet oriented primarily around browsing (E.N. Van Asperen, R.-D. Kahlke. 2015) (Stefaniak et al. 2021). It occupied a very broad range across Eurasia and is attested from as far as arctic Yakutia in Russia, where it inhabited a forest-grassland mosaic with a cold, moist climate, similar to the region today (Kirillova et al. 2017). In England, it is attested during cool-climate conditions, co-existing during the late Middle Pleistocene with species of mammoth and wild horses (Stuart, A. J. 1976). Unlike the woolly rhinoceros, however, the Merck’s rhinoceros was not a specialist of chilly environs, and it is attested in England also as part of definitively temperate assemblages, co-existing there with fallow deer (Dama dama), roe deer (Capreolus capreolus), the European elephant (Palaeoloxodon antiquus) and other more southerly elements (Rivals, F. & Lister, A. 2016). The Merck’s rhinoceros is also referred to as the “forest rhinoceros” in literature (Rivals, F. & Lister, A. 2016), a name which is not perhaps entirely un-apt, though the species possessed a flexible diet, and was not restricted to densely forested conditions (E.N. Van Asperen, R.-D. Kahlke. 2015). Isotope analyses from the enamel of teeth indicate that, like the narrow-nosed rhinoceros, it occupied primarily open woods and moist grassland, with some possible signatures of closed-canopy conditions as well, though this may be an artifact of preservation (Stefaniak et al. 2021).
Like the Merck’s rhinoceros, the smaller narrow-nosed rhinoceros was also something of a mixed feeder, though tending primarily towards grazing and inhabiting more open habitats than its larger relative (Rivals, F. & Lister, A. 2016). This is indicated both by its dentition, as well as its short legs and low head-position. In this, it was more specialised towards an open-habitat lifestyle than earlier species of Stephanorhinus, forming the latest stage of an adaptive trajectory towards grassier, less wooded conditions (Rivals, F. & Lister, A. 2016). Due to its ecology and distribution into grassland regions, the narrow-nosed rhinoceros has also been termed the “steppe rhinoceros” in the past—this is, however, a fairly unhelpful name, as all four rhinoceros species of the European Late Pleistocene occupied portions of the steppe as part of their ranges (Stefaniak et al. 2023). Both the so-called “forest” and “steppe” rhinoceroses had substantial overlaps in range and habitat, occupying a rich temperate-climate mosaic of floodplains, savannas and woodland (E.N. Van Asperen, R.-D. Kahlke. 2015), wherefore it seems prudent to simply avoid these terms entirely. It is interesting, in this context, to note that both the narrow-nosed and woolly rhinoceros share skeletal structures similar to that of the modern white rhinoceros (Ceratotherium simum), a grazer, while the mandibles of the Merck’s rhinoceros resemble those of the black rhinoceros (Diceros bicornis), a browsing species (Stefaniak et al. 2021). The niche partitioning in ancient Europe between the two species of Stephanorhinus thus appears to have resembled that seen between species of rhinoceros in present-day Africa.
Both the Merck’s and narrow-nosed rhinoceros had broad distributions in Central and Eastern European during the last interglacial, where they are recorded from Slovakia, Czechia and Poland (Stefaniak et al. 2023). Further west, England, Germany and most of the countries in south-western Europe provide Late Pleistocene attestations of Stephanorhinus (Pandolfi et al. 2017) (E.N. Van Asperen, R.-D. Kahlke. 2015). These species, then, were both widely distributed, ecologically and climatically flexible and by all indications highly successful, raising two obvious questions—the cause and timing of their extinctions.
Figure 3. Eemian scene showcasing a herd of narrow-nosed rhinoceroses (Stephanorhinus hemitoechus) crossing a river in autumn, somewhere in ancient Central Europe. In the foreground can also be seen Eurasian otters (Lutra lutra) and a European dipper (Cinclus cinclus).
Terms of use: Artwork by Hodari Nundu and Commissioned by The Extinctions
A substantial degree of uncertainty has long surrounded the exact progression of their disappearance, owing to patchy fossil records. The Merck’s rhinoceros was widely distributed throughout Asia until as late as 40kya and is recorded from one site in China around the Last Glacial Maximum, circa 25-30kya (Stefaniak et al. 2023). This latter record plausibly represents a very late relict population, and the situation in Europe is unclear: most probably, the species survived the onset of the last glacial period, the Weichsel (Stefaniak et al. 2023), retreating south into the warmer refugias alongside the remainder of the continent’s temperate fauna. When, precisely, it vanished from these regions is not known, however. It is even possible, though by its very nature difficult to prove, that it had entirely vanished from the continent following the Eemian (the last interglacial), and that Europe only ever formed part of its natural range while interglacial conditions prevailed. Either way, an investigation of its cousin, the narrow-nosed rhinoceros, may perhaps be fruitful in shedding some light by comparison.
Unlike the Merck’s rhinoceros, the narrow-nosed rhinoceros is quite well-attested from the last glacial period, owing perhaps to its more grazing-centred diet, which would have been less impacted by the drier, more open conditions of the time (Pandolfi et al. 2017). In France and Monaco, remains of S. hemitoechus are dated chronologically to circa 70-60kya (Pandolfi et al. 2017). In Spain, remains from Abric Romani near Barcelona have been radiometrically dated to circa 44.5 and 47.1kya, while another Spanish site finds a date between 45-42kya. In Italy, the narrow-nosed rhinoceros appears to have co-occurred with the woolly rhinoceros between circa 69-41kya, though factors of preservation make this exact dating problematic. The latest apparent evidence of any European Stephanorhinus seems to come from Portugal, where remains have been found in the Gruta da Figueira Brava cave. No direct dating has been done on the rhinoceros fossils here, but a human tooth from the cave was dated to circa 24kya, while a molar from the European elephant, P. antiquus, was dated to 31-30kya (Pandolfi et al. 2017). This is in and of itself interesting, though tentative, as the molar from Gruta da Figueira Brava is also the very latest evidence of P. antiquus, and indeed Portugal/south-western Iberia appears to have constituted a very late refugium for a number of temperate megafauna, of which the narrow-nosed rhinoceros was evidently a part. As to the causes of the final extinction, other than noting the peculiarity of a range-collapse in such an evidently successful species, the case of the two Stephanorhinus taxa cannot be disentangled from the broader conversation surrounding the end-Pleistocene extinctions, and the roughly contemporaneous collapse of Europe’s temperate megafauna.
The Furtive Elasmothere
A proper discussion on the natural history and eventual extinction of Elasmotherium cannot be undertaken here, since, as discussed, these events were overwhelmingly focused in Central Asia, whilst the genus’ presence in Europe was always of a decidedly peripheral nature. Nevertheless, some space may be dedicated to a brief sketch of its occurrences on the European fringe.
Elasmotherium sibiricum, the only Late Pleistocene representative of its lineage, was indisputably the largest rhinoceros of its time, with an estimated average weight of around 3.5 tons, and a potential maximal limit of 4-5 tons (Rivals et al. 2020), which if realised would place it in the same weight class as an adult African bush elephant (Loxodonta cyclotis) (Laurson, B., Bekoff, M. 1978). It reached 2 meters at the withers, and around 4-5 meters in length, with the second largest head of any rhinoceros (Rivals et al. 2020). This was famously adorned with an enormous domed protuberance over the nasal region, evidently coated in keratin. This anatomical feature has traditionally been interpreted as the base of a truly gigantic horn, giving the species its characteristic “unicorn” look. In 2021, however, a study was released that challenged this notion, and concluded through careful reconstruction that the structure could not possibly support a great horn, but was a much smaller, more rounded structure, protecting an enlarged nasal cavity. This, they speculate, may have enhanced the sense of smell, or even provided a sort of sound-amplification, allowing them to emit far more reverberating calls (Titov et al. 2021). No detailed follow-up has yet been carried out, such that the exact reconstruction of Elasmotherium’s horn is, at present, something of an open question.
Figure 4. Skeletal reconstruction of Elasmotherium sibiricum, rendering the nasal horn according to the traditional interpretation.
Terms of use: This image is licensed under a Creative Commons Attribution-ShareAlike 4.0 International. It is attributed to Ivan Iofrida. The image is unedited and the original can be found here.
Animals of the genus Elasmotherium are known from Europe since the Early Pleistocene, though initially in the form of three more ancient forms, E. peii, E. caucasicum and E. chaprovicum. These earliest Pleistocene records are from the southern Ukraine and Moldova and are particularly centred in the region above the Sea of Azov. At the end of the Early Pleistocene, E. peii and E. chaprovicum become confined to European Russia, while E. caucasicum becomes dominant in Ukraine (Stefaniak et al. 2023). All three species vanish during the Middle Pleistocene and are replaced by E. sibiricum, which from this period onwards is only attested in European Russia, with conditions in Ukraine seemingly growing unsuitable for the genus (Stefaniak et al. 2023). The elasmotheres appear to have been hyper-specialised grazers, with associated isotope values that indicate a habitat of dry steppe and even desert (Kosintsev et al. 2019). It is most probably this particular niche that, combined with competition from the better-adapted woolly rhinoceroses, prevented them from following the greater part of the mammoth steppe fauna west into ice age Europe, where conditions even at the hight of the glacial advance were far lusher and moister than their habitual haunts in the Eurasian interior.
When, precisely, the last populations of Elasmotherium vanished from the edge of the European continent is unclear, but the genus appears to have gone globally extinct sometime around 40kya, about 20 millennia prior to the onset of the Last Glacial Maximum (Stefaniak et al. 2023). It is worth nothing, however, that it is difficult to pin-point the exact date of extinction for any prehistoric species, particularly ones with patchy fossil records, and so this gap plausibly lies within the margin of error.
The Last Rhino Standing
Few extinct animals are as famous as the woolly rhinoceros. Like the woolly mammoth (Mammuthus primigenius), it shares in that quality of the paradoxical, the seemingly contradictory—an arctic rhinoceros, a woolly rhinoceros, when its living relatives are recognisable precisely as great, hairless beasts of the tropics. Nevertheless, there has also always been something of an obscurity about the animal—this, again, perhaps a kind of paradox. It is rarely absent from the ice age mural, but almost always lingering somewhat in the back and off to the side, rarely front and centre. Fittingly, maybe, its ecology has long been poorly understood, the causes and progress of its ultimate extinction dubious, and it is only in more recent years that we have come closer to a complex understanding (Tiunov, A. V & Kirilova, I. V. 2010) (Fordham et al. 2024).
As discussed, the Late Pleistocene woolly rhinoceros, Coelodonta antiquitatis, descended from the Asian Pliocene form Coelodonta thibetana, specifically through two intermediate species, Late Pliocene C. Nihowanensis and Early to Middle Pleistocene C. tologoijensis (Yuan et al. 2023). The evolutionary history of the genus saw an increasing broadening of its range until, by the Weichsel glaciation, it was found throughout essentially the entire cold-climate Eurasia, its range expanding and contracting with the cycling of glacials and interglacials (Fordham et al. 2024) (Stuart, A. J. 1976). It formed a characteristic component of the so-called Mammuthus-Coelodonta Faunal Complex, the “mammoth steppe fauna”, co-existing throughout most of its range with species such as the woolly mammoth, steppe bison (Bison priscus), wild horse (Equus ferus/przewalskii), saiga antelope (Saiga tatarica) and others (Kahlke, R-D. 2014) (Ma et al. 2021). In countries such as England, it appears in association with these species both before and after the last interglacial, vanishing from the records while temperate conditions prevail (Stuart, A. J. 1976).
The species inhabited primarily treeless landscapes of steppe and tundra (Stefaniak et al. 2021) (Tiunov, A. V & Kirilova, I. V. 2010), consuming both herbaceous plants and woody vegetation where present, the latter in the form of the dwarf shrubs characteristic of tundra environs. In places, these areas contained also elements of wetland, including evidence of peat bogs and marshy meadows, and woolly rhinoceroses have been found with teeth and intestines containing remains of aquatic plants (Stefaniak et al. 2021). Their diet appears to have varied seasonally, relying primarily on grazing during the summer half of the year, but potentially turning more towards shrubs in the winters (Tiunov, A. V & Kirilova, I. V. 2010).
Despite its broad distribution, the woolly rhinoceros does not seem to have been naturally abundant and was seemingly a slow disperser (Fordham et al. 2024). These factors would have naturally made subpopulations vulnerable to disruption and meant that recolonisation of suitable-but-vacant areas would not always have been rapid. The species’ decline began prior to the Last Glacial Maximum, particularly along the southern fringes of its range, where human densities were higher. Here, meat from woolly rhinoceroses may have provided up to 30% of the protein intake for some human populations (Fordham et al. 2024). As the Pleistocene drew unto its end, the major climatic disruptions, first of the Glacial Maximum, followed by the tumultuous onset of the Holocene, appears to have played a major role its metapopulation decline. Nevertheless, the changes at the end of the last glacial period were no more severe than those preceding the previous interglacial, and modelling indicates that even during the Eemian, large suitable habitats still remained throughout Eurasia (Fordham et al. 2024). The presence of a novel factor—human hunting, further destabilising the already struggling populations and inhibiting their ability to colonise new suitable habitat as the climate shifted—appears then to have played the ultimate deciding factor in the species’ disappearance.
Fig 5. 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
The European continent, however, was always a far extremity of its range. In the event, human pressures appear to have driven the species from most of the region even before climatic shifts rendered it environmentally unsuitable (Fordham et al. 2024). Nevertheless, it is doubtful whether the species would ever have persisted here into the Holocene. The very last individuals are attested from the north of the continent, circa 17-15kya; with their vanishing came the end of a continuous history of European rhinoceroses, dating back over 20 million years and spanning numerous diverse lineages. Even in their extinction, however, the impact of Europe’s rhinos is not altogether faded—it is a feature of megafauna, of all keystone species, that they leave an impact. The pressures they exert compel adaptive changes in the landscapes about them which may persist for many long ages, even in their wake. The very vegetation of Europe, the fruits dropped by its trees, the thorns with which they gird themselves, speak to an absence in the ecosystem, to the shadowy outlines of lost animals, still etched against the backdrop of its woods and acres.
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