Panthera spelaea, also called the cave lion (or sometimes the steppe lion), was an extinct species of big cat that lived in Eurasia and northwest North America during the Pleistocene era. Studies of ancient DNA show that it was closely related to modern lions (Panthera leo) but genetically different. Scientists believe the two species separated about 500,000 years ago.

The oldest fossils of this species (sometimes classified as Panthera fossilis or P. spelaea fossilis) in Eurasia are about 700,000 years old. This animal is likely an ancestor of the American lion (Panthera atrox). It lived from Western Europe to eastern Beringia in North America and was a top predator in the mammoth steppe ecosystem, sharing its habitat with other large carnivores like cave hyenas, with whom it sometimes competed.

The cave lion looked similar to modern lions, with yellowish-grey fur. Drawings in caves suggest that male cave lions may not have had thick manes like modern male lions. It is unclear whether they lived in groups like modern lions, but some scientists think they were mostly solitary, like tigers today.

Cave lions lived at the same time as Neanderthals and modern humans. People used their fur for clothing, and modern humans painted them in art.

Cave lions went extinct around 13,000 years ago during the end of the Pleistocene era. The exact reason for their extinction is unknown, but possible causes include changes in climate, fewer prey animals, and competition with other predators and humans.

Research history and taxonomy

In 1774, the Zoolithenhöhle cave near the village of Burggaillenreuth in Bavaria, southern Germany, was studied by Johan Friedrich Esper. He noticed that the cave contained bones from animals that no longer exist. In 1810, a fossil skull from the cave was named Felis spelaea by Georg August Goldfuss. This skull may be from the Last Glacial Period.

During the early to mid-1800s, scientists studied the remains of Panthera spelaea and found its body shape was most similar to lions, tigers, and jaguars. A study by W. Dawkins and W. Sandford in 1868 concluded that P. spelaea was most closely related to modern lions. At the end of the 1800s, M. Tschersky reported the first remains of P. spelaea from Siberia but mistakenly thought they were from tigers. In the 1900s and early 2000s, P. spelaea was often considered a subspecies of the modern lion, named Panthera leo spelaea. However, some scientists believed it was more closely related to tigers, suggesting it might be a subspecies of tigers, Panthera tigris spelaea. Genetic studies from 2004 and later confirmed that P. spelaea was most closely related to modern lions. Physical studies also showed that P. spelaea had unique skull and tooth features, supporting its classification as a distinct species.

In 2001, a subspecies named Panthera spelaea vereshchagini was proposed for seven specimens found in Siberia and Yukon. These specimens had smaller skulls and teeth than average P. spelaea. Before 2020, genetic tests using ancient DNA did not find evidence that this subspecies was distinct. However, analysis of mitochondrial DNA from 31 cave lions showed two separate groups. One group lived in western Europe, and the other was found only in Beringia during the Pleistocene. Because of this, the Beringian group is now considered a distinct subspecies, P. s. vereshchagini.

Lion-like cats first appeared in Tanzania’s Olduvai Gorge about 1.7 to 1.2 million years ago. These cats spread to Europe and Asia from East Africa during the Early to Middle Pleistocene, leading to the species Panthera fossilis. The oldest widely accepted P. fossilis fossils in Europe are about 700,000 to 600,000 years old, such as those found in Pakefield, England. Some older fossils from Western Siberia may be as old as 1 million years, and a 2024 study suggested P. fossilis was present in Spain about 1 million years ago. Scientists have debated whether P. fossilis was a separate species or a subspecies of P. spelaea. Genetic evidence shows that P. spelaea and modern lions split into separate groups about 1.85 million years ago. However, nuclear DNA studies suggest that modern lions and Eurasian fossil lions interbred until about 500,000 years ago, but no further interbreeding occurred after 470,000 years ago. Some scientists have questioned these younger dates because they do not match the fossil record well.

A diagram by Tseng et al. (2014) shows the evolutionary relationships of several big cats, including Panthera spelaea (cave lion) and Panthera atrox (American lion).

The arrival of Panthera (spelaea) fossilis in Europe was part of a major change in animal life during the Early-Middle Pleistocene. Many species that lived before this time, such as the giant hyena Pachycrocuta and the sabertooth cat Megantereon, became extinct. After P. fossilis arrived, the lion-sized sabertooth cat Homotherium and the "European jaguar" Panthera gombaszoegensis became much rarer and eventually died out in Europe during the late Middle Pleistocene. Competition with lions may have contributed to their extinction.

Specimens that are between P. fossilis and Late Pleistocene P. spelaea are called the subspecies P. s. intermedia. The transition from P. fossilis to P. spelaea shows a decrease in body size and changes in skull and tooth shape. Genetic studies of fossil lion remains show that the American lion is closely related to P. spelaea and likely separated from it when an early P. spelaea population became isolated south of the Cordilleran Ice Sheet. Early estimates suggested this split occurred about 340,000 years ago, but later studies suggest it happened around 165,000 years ago. This timing matches the first appearance of P. spelaea in Eastern Beringia (now Alaska and nearby regions) during the Illinoian period, about 190,000 to 130,000 years ago.

Description

Carvings and cave paintings of cave lions found in the Lascaux and Chauvet Caves in France date to the Upper Paleolithic period. A drawing in the Chauvet Cave shows two cave lions walking together. The lion in the front is slightly smaller than the one in the back, which has a scrotum and no mane. These paintings suggest that male cave lions did not have manes or had very small ones.

Early cave lions, classified as Panthera (spelaea) fossilis during the Middle Pleistocene, were much larger than later cave lions and modern lions. Some of these early lions were estimated to be 2.5–2.9 meters (8.2–9.5 feet) long, 1.4–1.5 meters (4.6–4.9 feet) tall at the shoulder, and weighed 400–500 kilograms (880–1,100 pounds). This made them among the largest cats that ever lived. Later cave lions, known as Panthera spelaea spelaea, were smaller but still large compared to modern cats. They were estimated to be 2–2.1 meters (6.6–6.9 feet) long and 1.1–1.2 meters (3.6–3.9 feet) tall at the shoulder. Over time, cave lions gradually became smaller until they went extinct. The last cave lions were about the size of small modern lions, measuring 1.2–1.3 meters (3.9–4.3 feet) long, 70–75 centimeters (2.30–2.46 feet) tall at the shoulder, and weighing 70–90 kilograms (150–200 pounds).

Panthera spelaea had a longer and narrower muzzle compared to modern lions. Their cheekbones were strongly curved, and their teeth had differences in shape, including preparastyles on the carnassial teeth. Like modern lions, female cave lions were smaller than males. Compared to earlier Panthera (spelaea) fossilis, later Panthera spelaea spelaea had larger incisor teeth, narrower and flatter canines, and narrower upper and lower third and fourth premolars with different tooth shapes. Their eye sockets were larger, and their muzzles were slightly narrower. Their nasal area was proportionally narrower, while the areas behind the eye sockets and near the ears were wider. Their ear bones were also more inflated.

In 2016, hair found near the Maly Anyuy River was identified as cave lion hair through DNA testing. Comparing this hair to modern lion hair suggests cave lions had fur similar in color to modern lions but slightly lighter. Cave lions likely had a thick, dense undercoat of yellowish-to-white wavy downy hair and a smaller layer of darker guard hairs, which may have helped them survive the cold Ice Age climate. Juvenile cave lions probably had yellow fur, while adult cave lions likely had grey fur.

Distribution and habitat

During the Last Glacial Period, P. spelaea lived in a continuous population across the mammoth steppe, stretching from Western Europe to northwest North America. It was found in many regions, including the Iberian Peninsula, Italian Peninsula, Southeast Europe, Great Britain, Central Europe, the East European Plain, the Ural Mountains, most of Northeast Asia (as far south as Northeast China, Japan, and possibly the Korean Peninsula), and across the Bering land bridge into Alaska, Yukon, and possibly Alberta. The cave lion lived at a wide range of elevations, with remains discovered over 2,000 metres (6,600 feet) above sea level in the European Alps and in Buryatia in Northern Asia. However, they likely did not live in mountainous areas all year.

The cave lion probably lived mainly in open areas such as grasslands and steppes, though it also lived in open woodlands. During the Last Glacial Period, it was often found in cold environments, but the species also lived in temperate areas, such as parts of Europe during the Last Interglacial/Eemian.

Paleobiology

P. spelaea was an important species in the mammoth steppe ecosystem, acting as one of the main top predators along with the gray wolf, cave hyena, and brown bear. Many bones from P. spelaea were found in caves, where bones from cave hyenas, cave bears, and Paleolithic tools were also discovered. Although they are called "cave lions," they likely used caves only rarely or not at all, living in areas without caves. Studies of their ecology suggest they were active during the day.

Isotopic analysis of bone collagen from remains in Europe and East Beringia shows that reindeer were a major part of cave lions' diets during the Last Glacial Period. They also hunted cave bear cubs and may have targeted adult cave bears sometimes. Research on the ratio of male to female cave lions suggests that healthy males likely hunted cave bears. Other studies of European remains indicate that cave lions ate wild horses, woolly mammoths, and cave bears. They may have hunted hibernating bears in mountain caves during winter. Bite marks on straight-tusked elephant bones in Germany, dating to the Last Interglacial, were likely made by cave lions scavenging.

Other possible prey included giant deer, red deer, muskox, aurochs, wisent, steppe bison, and young woolly rhinoceros. Cave lions likely competed for food with European leopards, cave hyenas, brown bears, gray wolves, short-faced bears, the sabertooth cat Homotherium, and Beringian wolves. Some cave lion bones found in cave hyena dens were from confrontations over carcasses. Many of these remains show no signs of being eaten, suggesting cave hyenas avoided eating cave lions, as seen in conflicts between spotted hyenas and African lions. Cave lion cubs and weak or sick adults were also found in cave hyena dens, likely due to cave lions hunting cave bears or stealing prey from hyenas. Some caves, like Zoolithen Cave, had many male cave lion bones, suggesting males may have entered caves to kill hyenas. Bite marks on cave hyena skulls may have been caused by cave lions. Isotopic analysis suggests cave lions and hyenas may have hunted different prey due to competition.

Cave lions may have hunted smaller brown bears but often lost their kills to larger ones. Unlike tigers, cave lions lived alongside large, strong wolves, which were serious competitors. After MIS 3, smaller and dwarf cave lions became more vulnerable to wolf attacks, especially during MIS 2. Like hyenas, wolves likely limited cave lions' access to large prey and outcompeted them after the Last Glacial Maximum.

Cave lion cubs likely lived in dens during their early lives, as modern lion cubs do, and were probably raised only by females, like living Panthera species. Whether cave lions lived in groups like modern lions is unclear. Some scientists think they were solitary, based on the lack of manes in Paleolithic art, while others argue that group living may have existed before manes evolved in modern lions. Differences in canine size between males and females suggest cave lions may have lived in groups or larger prides, but evidence is not enough to confirm this. Experts warn that sexual dimorphism alone cannot determine social behavior, as leopards are also highly dimorphic but live alone.

Isotopic analysis by Hervé Bocherens and colleagues suggests cave lions may have been solitary, as their diets changed after cave hyenas disappeared and they faced competition. In mountain areas and high altitudes, cave lions may have been solitary or hunted in pairs, similar to modern lions. Modern lions hunt in prides below 1,500 meters but are solitary or hunt in pairs at higher altitudes. This behavior is supported by the 1:1 male-to-female ratio found in Moravian Karst remains.

Relationship with humans

Neanderthals and modern humans both lived with cave lions. At Einhornhöhle in Germany, scientists found cave lion hand bones in layers of dirt and rock that are at least 190,000 years old. These bones may have come from a cave lion pelt that Neanderthals removed the skin from and brought to the site. In Scladina Cave in Belgium, bones from cave lions that are about 130,000 years old were used by Neanderthals to make tools for shaping or fixing stone objects. At Siegsdorf in Germany, a cave lion skeleton from about 48,000 years ago has a hole in one of its ribs. This hole may have been caused by a wooden spear pushed into the lion’s chest by Neanderthals. Other bones from the same lion show marks from cutting tools, suggesting Neanderthals later cut up the body for meat or other uses.

When modern humans arrived in Europe during the Upper Paleolithic period, they created art of cave lions. They painted cave lions on walls, carved their shapes into bones, and made sculptures, such as the Löwenmensch figure from Hohlenstein-Stadel cave in Germany. This figure has the body of a human and the head of a lion and is about 41,000 to 35,000 years old. Some cave lion teeth with holes may have been worn as decorations. In southern Italy, stones with carvings of cave lions were found. Like Neanderthals, modern humans used cave lion pelts, as shown by hand bones found at La Garma in Spain from about 16,800 years ago. Some bones from cave lions in layers of dirt and rock from the Upper Paleolithic, such as the Aurignacian and Gravettian periods in southern Germany, show cuts and signs of being made into tools. This might mean modern humans hunted cave lions, but evidence of this is rare.

Extinction

Scientists used a method called radiocarbon dating to learn that the species became extinct around the same time across its entire range during the final thousands of years of the Late Pleistocene, about 14,000 to 15,000 years ago. It may have survived for about 1,000 more years in the far eastern part of North America. This timing matches the start of a warm period called the Bølling–Allerød Interstadial, which led to the breakdown of the mammoth steppe ecosystem. The exact reason for its extinction is not certain, but it may have been caused by changes in the environment, such as the shift from open areas to forests, changes in the number of prey animals, and human activity. It is hard to determine which of these factors was most important. Competition with wolves may have played a major role in its extinction. Between 47,000 and 18,000 years ago, cave lions experienced a population bottleneck, which greatly reduced their genetic diversity. This likely happened partly because of unstable climate conditions. The extinction of this species was part of a larger event at the end of the Pleistocene, during which most large land mammals worldwide disappeared, including many large carnivores.

Mummified specimens

In 2008, a well-preserved adult cave lion was found near the Maly Anyuy River in Chukotka Autonomous Okrug. The specimen still had some clumps of hair remaining.

In 2015, two frozen cave lion cubs were discovered near the Uyandina River in Yakutia, Siberia, buried in permafrost. The cubs were estimated to be between 25,000 and 55,000 years old. Research suggests the cubs were likely less than a week old when they died, as their baby teeth had not fully grown in. Evidence also shows the cubs were hidden in a den until they were strong enough to follow their mother, similar to modern lions. Scientists believe the cubs were trapped and killed by a landslide. The lack of oxygen underground prevented their bodies from decaying, which helped preserve them. A second expedition to the site was planned for 2016 to search for a third cub or the cubs' mother.

In 2017, another frozen specimen, believed to be a male cave lion cub, was found on the banks of the Tirekhtyakh River in Yakutia, a tributary of the Indigirka River. This cub was thought to be about one and a half to two months old when it died. In 2018, another preserved cub was found 15 meters (about 50 feet) away. This cub was estimated to be about one month old when it died approximately 50,000 years ago and was thought to be a sibling of the 2017 cub. However, carbon dating showed the cubs lived about 15,000 years apart. The female cub was estimated to have lived 28,000 years ago, and the male cub 43,448 years ago. Both cubs were well preserved, though with some damage. The female cub was considered the best-preserved Ice Age animal discovered to date.