By Gabriele Russo1, Annemieke Milks2, Dirk Leder3, and Thomas Terberger3,4
1 Senckenberg Centre for Human Evolution and Palaeoenvironment, Eberhard Karls University of Tübingen
2 University of Reading
3 Lower Saxony State Office for Cultural Heritage (Niedersächsisches Landesamt Für Denkmalpflege)
4 University of Göttingen
Felids have always played a primary role in the history of humanity, ranging from small domestic companion cats to formidable predators like lions, which have unfailingly evoked fascination and respect. We have shared most of our evolutionary history with lions, which has likely been partly shaped by their presence (Brantingham 1998; Stiner 2012).
The earliest ancestors of modern lions evolved in Africa, with the first fossils dating back to nearly 4 million years ago; these fossils originate from sites familiar to prehistoric archaeologists, such as Laetoli and Olduvai Gorge in Tanzania (Werdelin and Dehghani 2011). Around 700,000 years ago, some descendants of these lions migrated to Europe (Lewis, Pacher and Turner 2010), and subsequently evolved into the cave lion (Panthera spelaea) by 400,000 years ago. This species spread across Eurasia until its extinction around 13,000 years ago (Stuart and Lister 2011).
Throughout the same extended period, various hominin species also inhabited Eurasia. The earliest interaction between humans and lions is documented in Western Europe, specifically in level TD10-1 of the Gran Dolina site at Atapuerca (Blasco et al. 2010). Here, the remains of lions butchered by Middle Pleistocene hominins approximately 350,000 years ago were discovered. Increasing evidence of human-predator interactions becomes evident with the arrival of our species (Homo sapiens) in Europe during the Upper Paleolithic, approximately 40,000 years ago. These foragers introduced a new dimension to the relationship between humans and lions, embracing both indirect expressions like portable and rock art (E.g., Wehrberger 1994; Conard 2003; Clottes and Azéma 2005) and direct engagements. This includes the utilization of lion body parts for tools or ornaments (e.g., Vanhaeren and d’Errico 2006; Cueto et al. 2016) as well as an intensified exploitation of cave lions (e.g., Kitagawa et al. 2012; Wojtal et al. 2020). However, the evidence of interactions with lions in the long period between the site of Gran Dolina and the Upper Paleolithic—usually defined as the Middle Paleolithic—remained overlooked and was often disregarded as occasional exploitation due to interspecies competition. See Figure 11. During this time, Western Europe was inhabited by Neanderthals (Homo neanderthalensis), an extinct human species closely related to our own. While during the 20th century Neanderthals were usually depicted as primitive, less intelligent and brutish, in recent years, increased evidence has overturned this view (see Wragg Sykes 2020; Romagnoli, Rivals and Benazzi 2022). Our recent study sheds light on a previously-unexplored aspect of our extinct relatives: their dynamic relationship with one of the most formidable predators with which they shared territory: the cave lion.
Figure 11. Paleolithic sites with direct evidence of human-cave lion interaction in the form of subsistence activity, culture (i.e., rock or mobile art), or both. 1 Einhornhöhle, 2 Siegsdorf, 3 Gran Dolina, 4 Caverna delle Fate, 5 Chez-Pinaud (Jonzac), 6 Cueva de Bolomir, 7 Le Portel, 8 Grotte du Renne, 9 Hohlenstein-Stadel, 10 Vogelherd, 11 Hohle Fels, 12 La Garma, 13 Chauvet, 14 Peyrat (Saint-Rabier), 15 Pair-non-Pair, 16 La Gravette, 17 Grotte Duruthy, 18 Pavlov I, 19 Dolní Věstonice I, 20 Trois-Frères, 21 Grotte de la Vache (Ariège).
We recently reported the discovery of bone remains bearing cut marks from a lion’s paw, which were part of a lion pelt dating back over 190,000 years and the oldest direct evidence of a large predator being hunted and killed in human history (dated to 48,000 years ago) in Scientific Reports (Russo et al., 2023). Here, we give an overview of that research.
Materials and Methods
We conducted an examination of cave lion remains from two distinct sites. The first set was obtained from Einhornhöhle (in German, Einhornhöhle means “Unicorn Cave”), a cave situated in central Germany within the low Harz mountain range which borders the North European Plain. Three paw bone elements, including two toe bones where the claws were attached, were recovered. These cave lion bones originated from layer H, located in a gallery about 30 meters away from the prehistoric cave entrance, underlying a deposit dated to approximately 190,000 years ago, giving layer H a minimum age. One of the toe bones exhibited cut marks, which were analyzed using 3D microscopy to identify the nature of the incisions.
The second set of cave lion bones constitutes a partial skeleton from a single individual unearthed at the open-air site of Siegsdorf in southeast Germany (Bavaria), at the feet of the Alps. This well-known skeleton, previously documented in other publications (e.g., Gross 1992; Barnett et al. 2016) and directly dated to around 48,000 years ago (Rosendahl and Darga 2004), displayed cut marks on various bones, including ribs, hip, and hind bones. During the re-examination of these bones, we identified human-induced modifications associated with the lion kill, previously unreported and henceforth referred to as “hunting lesions.” To reconstruct the events leading to the creation of these marks, we employed 3D microscopy to measure the modifications, along with forensic reconstructions utilizing micro-computed tomography scans and digital models to depict the ballistic trajectory.
Results
The scores on the lion paw bone from the Einhornhöhle exhibit a straight trajectory and taper at both ends, features typically associated with cut marks created by stones – lithics – with a cutting edge during the butchering process of a carcass. To confirm this inference, we employed 3D microscopy, which revealed additional microscopic features. These features not only align with typical cut marks but also suggest that the tool used to create them was intentionally modified – retouched – indicating it was not a simple lithic flake. See Figure 12. Similar artifacts were also discovered in the same layers where the lion remains were found.
Figure 12. Cave lion remains from Einhornhöhle Area 1. Phalanx III and close-up view × 30 and × 500 magnifications of the cut marks, and the other unmodified bones.
The cut marks were precisely located in correspondence with the tendon attachment. This, coupled with the fact that there were only paw elements from a cave lion in layer H, suggests that they resulted from the skinning process and that the targeted final product was the pelt with paws and claws still attached. The bones represented all that remained from a lion pelt. The partial skeleton of the Siegsdorf cave lion exhibited cut marks that were already documented. These cut marks are textbook examples, easily visible to the naked eye and microscopically distinct from a few other traces on the bones that naturally occurred after exposure to environmental elements.
One of the ribs showed a partial perforation previously associated with carnivore gnawing in earlier publications. However, typical carnivore bites create two teeth marks on each side of the bone, formed as the jaw closes, while the perforation on the lion rib exhibited a single mark. To explore the hypothesis of a carnivore bite, we compared the measurements of the puncture on the rib to punctures on bones created by modern large carnivores, including lions, hyenas, bears, and wolves. The modification on the lion proved to be both larger and deeper than any of the other carnivore bite marks.
We then examined the possibility that humans were responsible for this perforation by comparing it to perforations created by spears and javelins, defined as hunting lesions, including both archaeological and experimental data. This examination revealed that the puncture on the lion’s rib not only perfectly aligned within the range of hunting lesions but was also more closely aligned with punctures created by wooden-tipped spears. See Figure 13. The ballistic reconstruction further revealed that the lion must have been lying on its right side at the moment of the fatal stab.
Figure 13. On the left: Siegsdorf lion skeleton with distribution of observed anthropogenic modifications. Elements highlighted in gray represent those that were originally unearthed. On the right: Details of the puncture on Siegsdorf ’s cave lion rib.
While examining the skeleton, we uncovered previously-unreported bone modifications: notch-like damage on the ribs and one vertebra. These differed from carnivore-induced damage and from natural modifications on the same elements. Their location and morphology more closely resemble another type of hunting lesion called drag marks (so-called as they result from the violent impact of a projectile hitting bone transversally; (Duches et al. 2016). Few studies explore this category of hunting lesion, and none analyze drag marks created by Neanderthal technologies such as wooden spears/javelins or stone-tipped spears. However, computer tomography revealed these modifications were made when the animal carcass was fresh, suggesting a connection to either lion hunting or processing.
Discussion and Conclusion
The presence of only lion paw bones at the Einhornhöhle suggests that the lion was processed elsewhere. Simultaneously, the absence of other evidence makes it unlikely that the claws were worn as pendants or used as aesthetic components for clothing. Instead, it is more likely that they were part of a pelt introduced to the cave. See Figure 14.
Figure 14. Diagram with the interpretation of the chain of events involving the cave lion pelt from Einhornhöhle: The lion was skinned elsewhere, and its pelt was brought into the cave, eventually being abandoned.
Soft skins or pelts seldom survive in prehistoric contexts. However, their presence at archaeological sites can be deduced through indirect evidence, such as skinning marks on bones, tools associated with hide work, micro-traces left on stone tools, or even the latitude and environmental context of the site. Such evidence is present in the Paleolithic record as far back as 500,000 years ago or earlier (Rodríguez, Willmes and Mateos 2021); worked skins are usually thought to have been used as clothing or bedding elements to cope with the climate.
Zooarchaeological records indicate that Neanderthals exploited various animals, including large carnivores like bears and wolves (e.g., Gómez-Olivencia et al. 2018; Romandini et al. 2018). Lions were also on the menu, even for pre-Neanderthals (Blasco et al. 2010). Therefore, it is unsurprising that these hominins utilized the skin of this large predator. The uniqueness of the lion pelt from Einhornhöhle lies in the fact that this has never been demonstrated before in the entire archaeological record, including for contemporaneous H. sapiens in Africa, who also coexisted with lions. Additionally, the deliberate retention of aesthetic elements such as claws in this skin is a novel aspect. This detail is crucial as it underscores the care and effort Neanderthals invested in preserving these elements in the skin, a behavior not recorded for any other exploited species.
The lion holds meaning in many modern African indigenous cultures (Ontiri et al. 2019) and also carries great symbolic value in modern pop culture. The same was true for Upper Paleolithic H. sapiens. The ivory sculpture of a lion man found at the Swabian cave of Hohelnstein-Stadel (Germany), dated to about 40,000 years ago, is a well-known example of this (Wehrberger 1994). Therefore, we argue that the careful processing and use of lion pelt with claws from Einhornhöhle represent evidence of the capacity of Neanderthals to engage culturally with large predators.
Figure 15. Diagram depicting the interpretation of the chain of events involving the cave lion skeleton from Siegsdorf: The lion was either actively hunted using javelins to disadvantage it, then received a coup de grâce with a wooden spear; or it was ambushed and killed while asleep, butchered at the kill site without bone breakage, and subsequently, the carcass was abandoned.
On the other hand, the skeleton of the lion from Siegsdorf offers a unique snapshot of a Neanderthal day. See Figure 15. The lion was either actively hunted, using javelins – that created the drag marks – to disadvantage it and then received a coup de grâce with a wooden spear, creating the partial puncture. Alternatively, it may have been ambushed and killed while asleep. Regardless of the hunting method, the lion was subsequently carefully butchered, eviscerated, and left at the site without breaking the bones.
Neanderthals were skilled hunters and employed sophisticated techniques to pursue their prey. They strategically used spears both for impaling and throwing by targeting their prey’s vital areas (e.g., Boëda et al. 1999; Gaudzinski-Windheuser et al. 2018). Archaeological data suggests that Neanderthals were selective in their hunting practices, focusing on specific species and individuals at particular sites and times while opportunistically hunting any creature they could capture on other occasions (see Romagnoli, Rivals and Benazzi 2022 for a comprehensive review).
However, the active hunting of a large predator such as a cave lion by these hominins was never documented before and was previously associated only with the technological proficiency of H. sapiens. The remains of the cave lion from Siegsdorf represent, therefore, not only a testament to Neanderthal hunting abilities but also the earliest evidence thus far of a large predator hunt in human history.
Our study adds a new dimension to the understanding of Neanderthal culture. The interactions of these hominins with large predators included the cultural use of lion body parts and the ability to hunt them, a behavior initially attributed exclusively to our species.
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