A World of Lions

Lions (Panthera leo) are arguably the most iconic species alive today. A staple among African megafauna, lions are loved around the world for their grace and prowess, often featured as a symbol of power and leadership in many human cultures, and are prominent in countless stories and works of art. As apex predators and keystone species, they are also crucial in maintaining populations of large herbivores such as zebras, buffalo, and different antelope species.[1] However, a major misconception today is that lions are an exclusively African species. While lions originally evolved in Africa and are most common on that continent today, they once inhabited every continent except for Australia and Antarctica as recently as 11,000 BC, making them the most widespread terrestrial mammal at the time.[2] And while lions may no longer inhabit these regions, they still endure within the human cultures that they inspired, and are missed in the ecosystems they left behind.

From Wikimedia: an artist’s reconstruction of an American lion (Panthera leo atrox)

The genus Panthera originated in Asia during the late Pliocene epoch, which resulted in the divergence of the big cats, with the ancestors of tigers and snow leopards remaining in Asia while the ancestors of leopards, jaguars, and lions migrated west towards Europe and Africa.[3] In Africa, lions and leopards began to take their current forms. While leopards were similar to other cats in their morphology and behavior, lions were unique not only for their larger size, but also for their social behavior, a rare phenomenon among wild cats. Their lifestyle of living in groups enabled them to take on larger prey, live in larger territories, and raise their young with higher survival rates, allowing them to dominate most environments that they encountered[4](lions are usually absent from tropical rainforests, making the expression “king of the jungle” a bit of a misnomer[5]). These cats eventually made their way out of Africa and entered Europe and Asia, where they evolved into larger forms (possibly in response to colder climates), and where they preyed upon a new assortment of species, including reindeer, horses, and aurochs (the wild predecessor to most domestic cattle breeds). Lions were not the only large predators to thrive in prehistoric Europe. In addition to contemporary carnivores such as wolves, brown bears, and lynx and wolverines, other large carnivores also prowled the ancient European landscapes, including cave hyenas (an extinct subspecies of the extant African spotted hyena), leopards, as well as extinct taxa such as Homotherium and the Eurasian cave bear (Ursus spelaeus). As the glacial cycles of the Pleistocene progressed, these species became a part of the mammoth steppe, a massive grassland ecosystem that stretched from Britain, across Eurasia and Beringia,[6] all the way into the Yukon. This massive biome, which has since been replaced by boreal forests and tundra, supported large populations of big herbivores, including woolly mammoths, woolly rhinoceroses, bison, musk ox, Irish elk (Megaloceros giganteus), and many other species both living and extinct.

During glacial maximums when sea levels were low, lions traversed their way into North America, where they evolved into their largest form, the American lion (Panthera leo atrox). Weighing up to 351 kilograms, 2.5 meters in length (not including the tail), and up to 1.2 meters tall at the shoulder, the American lion was one of the largest felines to ever exist, and was approximately the same size as Smilodon populator, the contemporary South American saber tooth cat.[7] While people view lions as a tropical species today, it is evident that in prehistoric times they, like brown rats or ospreys, had a cosmopolitan distribution, meaning that they were present in almost all of the earth’s terrestrial environments, and were not restricted to any one ecosystem or biome. However, another African species, us modern humans, were quick to eliminate lions in much of their territory. It’s unlikely that humans directly preyed upon lions (other than for possible ceremonial reasons, as the Massai people of east Africa used to do[8]), but modern humans, with their newly developed hunting technologies, were able to decimate populations of many large herbivores, especially in the Americas where most megaherbivores became extinct following the arrival of humans. Without suitable prey populations to sustain them, lions soon became extinct in the Americas, northern Europe, and Siberia.[9] However, in regions where species had already adapted to other hominids, such as the neanderthals or homo erectus, and where local species of megafauna had gone extinct more gradually, humans reflected their view of lions in their art. Eurasian cave lions (Panthera leo spelaea) are depicted on the walls of Chauvet cave in southern France, in a scene where some researchers believe they are hunting.[10] Another example of the adoration that humans held for lions is seen in the lion man of the Hohlenstein stadel, a sculpture carved from mammoth ivory from approximately 40,000 years ago, depicting a lion-human hybrid, and is the oldest known animal carving in the world.[11]

Despite their disappearances from the Americas and much of Eurasia during the Pleistocene epoch, lions were still widespread inside and outside of Africa during much of the Holocene and in historical times. Lions were still present in Spain during the early Holocene, and were present in in Ukraine as recently as 3,000 BC. Lions were especially common in Greece for thousands of years, and became a cultural icon in antiquity, featured prominently in the folklore and mythology in the region. However, as a result of a growing human population and increasingly complex society, lions were persecuted both for sport and for the protection of livestock. Lions disappeared from Europe by 100 ad, and with them a once-integral part of European nature.[12] Over the following centuries, lions disappeared from more and more areas. Lions were no longer present in the Caucasus until the 10th century, and had vanished from turkey and Syria by the mid-19th century. The last lions of North Africa disappeared by the 1940s, and lions had been eradicated from Iran by the 1940s.[13] In India, where lions were once widespread, heavy hunting pressure following British colonization led to the depletion of lions around the country until just a single population remained in the Gir national forest. This population represents the last population of wild lions outside of Africa, although their numbers are on the increase.[14] The species hasn’t fared well in Africa either. Lions are now critically endangered in west Africa,[15] and remain at risk and in decline in much of eastern Africa where they are persecuted for their attacks on livestock, either through direct killings, or through the poisoning of animal carcasses that they feed on, which also damages populations of many other carnivorous animals.[16] They are also overhunted by wealthy foreigners and poached for traditional medicines in certain African and Asian markets.

Possibly as few as 20,000 lions are left in the wild, down from the estimated 450,000 individuals in the 1940s.[17] However, lions as a species are still listed as vulnerable by the international union for the conservation of nature (IUCN), and there is reason to be optimistic for the future of lions. Conservation successes in southern African countries like Namibia, Botswana, and South Africa have allowed lion populations to increase,[18] and as more people begin to realize the ecological and cultural importance of the animals, some societies have begun to change their ways to accommodate them, such as the recent decision to reintroduce lions to Rwanda,[19] or the changes made by the Massai people of Kenya and Tanzania, who once hunted lions as a way to prove masculinity, but now have devoted much of their time to protecting lions.[20]

From Wikimedia: An Asiatic lion (Panthera leo persica). Today, they are only found in India.

While it is undoubtedly important to conserve lions in their current ranges, is it right for us humans to only allow them to inhabit a fraction of the range that they lived in during the Pleistocene? Those who advocate for Pleistocene rewilding believe that species such as lions should be reintroduced to their prehistoric ranges, not only as a conservation strategy for that species, but also to help restore ecological processes that went missing after that species disappeared. A compelling case for restoring lions in North America would be to control populations of feral horses. Horses, which were reintroduced approximately 500 years ago after being extinct in North America for thousands of years, are known to aid grass seed dispersal and to increase the diversity of native grass species, but when their populations are too robust, they can cause soil compaction, which can lead to erosion.[21] Lions, which naturally prey upon equids such as zebras and wild asses in Africa, would play an important role in regulating the populations of horses and other large ungulates.[22] However, given public attitudes toward the reintroductions of large native predators, such as wolves or grizzly bears, it is unlikely that this will happen in the near future.

Despite the dismal outlook in North America, the situation in Europe may be more suitable. Despite being a smaller continent with a larger population than North America, the natural setting in Europe has rebounded in recent years due to a mass migration of people from rural areas to urban areas, allowing many species to reclaim their prior territories. Populations of moose, red deer, and other large ungulates have expanded. Multiple organizations have reintroduced wild horses, European bison, and wild cattle to national parks and wild areas throughout Europe, and strict protections for carnivorous animals have allowed animals such as brown bears, wolves, and lynx and wolverines to reclaim their old territories.[23] In some areas, species that have not inhabited Europe since prehistoric times have been reintroduced, including fallow deer, musk oxen, and water buffalo, and are now considered part of the natural setting. As it stands, there are even tentative (albeit controversial) plans to restore lions to the Far North of Siberia.[24] Given that lions disappeared from Europe much later than some other taxa currently being restored, and that many of its old prey species are extant in Europe, there is a compelling argument for its return.

While there are potential ecological benefits to restoring lions outside of their current and historical ranges, there is also a feeling of wonder that comes with the prospect of doing so. Lions, with all of their grandeur, are an irreplaceable part of nature, and embody the spirit of wilderness itself. So while us humans may have forgotten about them, we still very much live in a world of lions.

Notes

  1. ALERT, “The Ecological Role of Lions,” LionAlert.
  2. C. R. Harington, “Pleistocene remains of the lion-like cat (Panthera atrox) from the Yukon Territory and northern Alaska,” Canadian Journal of Earth Sciences.
  3. Ji H. Mazak, “Oldest Known Pantherine Skull and Evolution of the Tiger,” PLOS One.
  4. Kristin Nowell, “Wild Cats,” IUCN/SSC Cat Specialist Group.
  5. While lions generally avoid rainforests, they have been known to inhabit them, as reported by Mongabay in 2012.
  6. During glacial maximums during recent ice ages, the sea separating Asia and North America receded, leaving a low altitude plain in its absence, known as Beringia.
  7. American Lion,” Yukon Beringia Interpretive Center.
  8. The Massai, a pastoral culture, were freverently protective of their livestock against lions, and the act of slaying a lion was viewed as a rite of passage from boyhood to manhood.
  9. Joachim Burger, “Molecular phylogeny of the extinct cave lion Panthera leo spelaea,” Molecular Phylogenetics and Evolution.
  10. Panel of the Lions,” Bradshaw Foundation.
  11. Martin Bailey, “Ice Age Lion Man is world’s earliest figurative art sculpture,” The Art Newspaper.
  12. R.S. Sommer, “Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review,” Journal of Zoology.
  13. Past and present distribution of the lion inNorth Africa and Southwest Asia,” The Asiatic Lion Information Centre.
  14. Jeremy Hance, “Asiatic lion population rises by 27% in five years,” Mongabay.com.
  15. Brian Clark Howard, “Lions Approach Extinction in West Africa,” National Geographic.
  16. Lion Poisoining– An Urgent Issue,” Living with Lions.
  17. Declining Lions,” National Geographic Big Cats Initiative.
  18. Panthera leo,” IUCN Red List.
  19. Clement Uwiringiyimana, “Rwanda brings lions back to safari park, plans for rhinos,” Reuters.
  20. Meghan Dunn, “Transforming lion killers into ‘Lion Guardians,'” CNN.
  21. Stacey D. Ostermann-Kelm, “Impacts of feral horses on a desert environment,” BMC Ecology.
  22. Josh Donlan, “Pleistocene Rewilding: An Optimistic Agenda forTwenty-First Century Conservation,” Advanced Conservation Strategies.
  23. Stefanie Deinet, “Wildlife Comeback in Europe,” Rewilding Europe.
  24. Martin W. Lewis, “Pleistocene Park: The Regeneration of the Mammoth Steppe?,” GeoCurrents.