Species Fact: Lions


This beloved wild cat needs no introduction.

However, it might surprise you to hear that the King of Beasts has a lot in common with Fluffy, the king of our hearts (or bane of existence, if you’re allergic to house cats).

For instance:

  • In both species, related females group together in a social unit. With Fluffy, it’s called a colony, not a pride.
  • Both lionesses and she-cats help out with birth and then share nursery duties when raising their cubs and kittens, respectively.
  • Lions and tomcats vocalize at the right frequency to be heard far away. However, size does matter: other lions will hear that roar 3-6 miles away, while a tomcat’s yowl is only audible to other domestic cats within about a quarter mile.

Of course, there are also some very obvious differences.

Scientific name:

Panthera leo (Linnaeus 1788). Most of us know the African lion (P. leo leo), but there is also an Asiatic lion (P. leo persica) in India.



Physical statistics:

These figures are from the Cat Specialist Group website (see source list), except where noted. The data range is so wide because lions are much bigger than lionesses.

African lion:

  • Weight: 240 to 600 pounds.
  • Height at the shoulder: 3 to 4 feet tall (Sunquist and Sunquist)
  • Body length: 4-1/2 to 8 feet.
  • Tail length: 2 to a little over 3 feet.
  • Coat: No patterns, except in cubs and occasional faint belly markings in some adult females; color generally light tan to silvery gray to yellowish red or dark brown. Underside is generally a paler version of the overall coat color. (Sunquist and Sunquist)
  • Average litter size: 1-4 cubs.
  • Average life span: 12 to 16 years.

Asiatic lion (difference highlighted):

  • Weight: 240 to 420 pounds.
  • Body length: 4-1/2 to 8 feet.
  • Tail length: 2 to a little over 3 feet.
  • Average litter size: 1-4 cubs.
  • Average life span: 16-18 years.

Features unique to lions:

  • Mane and tail tassel: Whether it’s yellow, brown, or reddish brown, the mane usually starts to grow as a young lion matures, at around age 3-1/2; then it darkens and thickens with age. (Sunquist and Sunquist) The Asiatic lion’s mane isn’t as impressive as its African counterpart, but in cooler climates this subspecies does grow a thick mane. (Loveridge et al., 2010b) The Asiatic lion has the longest tail tassel, though. (It also typically has a thick fold of belly skin, unlike African lions.)
  • The pride: Feral house cats form colonies, and related females of all cat family species tend to live close to each other. However, only the lion is this social. It’s hard even to generalize about pride structure, since numbers of cats and other details vary from place to place and even from pride to pride. Adult lionesses (all of them related) are the core members. They average 4 or 5 per pride in India’s Gir National Park and up to 18 out on the Serengeti. (Sunquist and Sunquist) Lionesses typically are accompanied by either growing cubs or subadults who are learning the fundamentals of hunting and pride living. Adult males come and go (2 to 6 per pride in Gir, on average, and 1 to 7 in Serengeti lions, per Sunquist and Sunquist), with an average tenure of 24 to 36 months (Cat Specialist Group), though some male coalitions last for years — hey, these are cats. Just like Fluffy, lions don’t always follow rules that have been tentatively put together by scientists trying to understand these beautiful animals. Lion range size varies a lot, too, and not just because of prey abundance, although that’s certainly a major factor. A Serengeti pride may own 10 to 20 square miles, for example, while elsewhere on the plain another pride occupies 150 square miles or more. (Ewer) As for sharing their space, it depends on what source you believe: prides are either highly territorial or, per other references, pride ranges overlap everywhere except in the central zone that each group calls home. It’s probably a little bit of both. Again — cats.
  • The roar: According to Kitchener et al. (2010), sonograms show that only the lion has a structural call series. Leopards and jaguars just show a few parts of this pattern. Tigers do the main call and grunt, but they do not roar, and neither do snow leopards.


Where found in the wild:

Presently, the 400 or so individuals in and around Gir Park, Gujarat State, India, are the only known wild population of Asiatic lions. They used to be found throughout southwestern Asia. (Cat Specialist Group)

African lions are found south of the Sahara in various habitats, mainly savanna lands that support numerous prey animals (preferably medium- to large-sized hoofed plant-eaters, though lions will eat almost anything). They like open spaces as well as woodland that gives them enough shelter and cover for hunting. Historic records show that P. leo once was widespread throughout all of Africa and much of Europe, the Middle East, and southwestern Asia. It’s hard to estimate how many wild lions live in Africa now, but the total might be around 20,000 to 30,000, mostly in southern and eastern Africa, with more than half of them in conservation areas. (Cat Specialist Group; Macdonald and others, 2010a)

Closest cat-family relatives:

Most studies of the big cats show that lions are a little more closely related to jaguars and leopards than they are to tigers and snow leopards. (Christiansen; Kitchener et al., 2017)

Famous real-life lions:



How lions hunt and eat:

Lions form prides so they can “gang up” and catch more prey, right?

Actually, the jury is still out on possible connections between lion society and hunting.

Like all cats, lions are stalk-and-ambush hunters, though they certainly can take larger prey than Fluffy can.

This is why, in addition to the typical killing bite to the back of the neck, lions (and other big cats) also are skilled in using either a throat- or snout-covering bite that suffocates their hapless victim, sometimes without even breaking its skin.

As for catching prey, lionesses have been seen using any of three different group hunting techniques (Ewer):

  1. Fanning out and flushing prey out of cover
  2. Circling around and flushing the prey toward another lioness who’s waiting in ambush
  3. Herding prey into a cul-de-sac (don’t laugh — where there is little cover, hunters of all sorts must know their terrain very well, or perish)

Even these clever groups only have a success rate of about 30%, tops. (Sunquist and Sunquist)

Perhaps, instead, having a pride helps lions defend their food. (Macdonald and others, 2010a)

Scavenging is a big problem for lions, who operate out in the open (usually right after sunset and before dawn, and perhaps also around midnight).

Hyenas, for instance, immediately know when there’s been a kill. And if a large enough pack gathers, hyenas will chase lionesses away from a carcass.

But if the pride’s lion or lions are around, these scavengers will just have to wait for leftovers. (Macdonald and others, 2010a)

The down side to this is that one carcass only goes so far. Competition is fierce during lion feeding, and the youngsters often get pushed aside, though Mom tries to make room for them when possible.

How they reproduce:

Let’s go with the domestic-cat comparison here. (Numbers are from Ewer unless otherwise noted.)

Domestic cats can have 2 litters a year, with breeding seasons focused around October through January and from July to September.

Lions will mate at any time of year, but once pregnant, a lioness doesn’t come into heat again until her cubs mature, about 24 months after birth.

House cat pregnancies last 65 days; for lions, that’s 105 days or so.

A newborn kitten only weighs around 3 ounces, and its eyes stay closed for up to 20 days.

A newborn lion cub, of course, is heavier — about 2-1/2 to 3 pounds — and its eyes generally are open at birth, or soon afterwards.

Lion cubs are often spotted, and these spots sometimes fuse into stripes. The marks generally fade away as the cat matures. (Sunquist and Sunquist)

Domestic cats are usually weaned by 8 weeks and independent from Mom at around 6 months of age. This whole process takes about four times as long for lion cubs. During those 2 years of raising Junior, the lioness does not come into heat again.

Unless there’s a pride takeover.

Subadult males disperse, that is, they leave their family group and make their own way in the world. Lions are almost unique in that their nomad males form coalitions. (Cheetahs do something similar, but not to such an extent, and unlike lions, cheetah coalitions don’t defend females, per Macdonald et al., 2010a.)

For a while, male coalition members hone their hunting and other survival skills, but when they’re about 4 years old, they’ll start looking for a pride.

This leads to some intense confrontations.



Sometimes the resident wins. Sometimes the newcomers take over, and when they do, they drive out all the subadult males and, sadly, kill all the cubs so that they can sire their own offspring.

Almost a third of all cub deaths on the Serengeti are from such infanticide. (Macdonald and others, 2010a)

It’s brutal, but the fittest lion is always the one who gets to pass along his genes.

Interactions with people:

Where to start? We have always loved and feared lions. Check out Sunquist and Sunquist for some fascinating historical details.

Basically, as Loveridge et al. (2010a) note, there are two very different kinds of interaction between lions and people.

On the one hand, lions are cultural icons, as well as tourist attractions and conservation symbols.

On the other, they prey on livestock and on us (particularly in Tanzania, where almost 600 people were killed and hundreds more injured between 1990 and 2004, the last dates I could find data on; from 1978 to 1991, Gir lions killed 28 and injured 165 people, mainly near the preserve rather than in it). And we overexploit lions, steal their prey for bushmeat, and break up their habitat.

For most readers, this talk about predation and man-killing are intellectual problems, but for anyone living near a preserve, especially a subsistence agriculturalist who really can’t afford to lose livestock and is afraid to let the children outdoors to play, they are crises.

Many people, especially in India for cultural reasons, try to work with conservationists and other stakeholders, but it’s very difficult to get everyone on board. And this is vital — it only takes one aggrieved human with weapons and/or poison to wipe out an entire lion pride.

Red-listed?

  • African lions: Vulnerable.
  • Asiatic lions: Endangered.

Fossil relatives:

Lions have the best fossil history of all the big cats, yet their evolution is still mostly a mystery.

Biomolecular techniques date the origin of all big cats to at least 10 million years ago, but the earliest widely accepted fossils are much younger. One of the best known big-cat fossils, from Laetoli, in Africa, is less than 4 million years old.

And it might have been related to modern lions. No one knows for sure. This big cat certainly was lion-sized, but the material is broken up and not well preserved. (Werdelin and Dehghani)

The oldest unequivocal fossil lion is less than 2 million years old and comes from a very familiar place: Olduvai Gorge.

Two ancient species you might have heard of are the cave lion (Panthera spelea) and the American lion (Panthera atrox or P. leo atrox).

Both evolved during the Pleistocene epoch, with Spelea the older of the two, but not everyone agrees that they were lions. They may well have been, but the debate is ongoing. (Bocherens, Christiansen, King, Werdelin and others)

Lions_group_Chauvet_Cave
We have eyewitness descriptions of Spelea from around 30,000 years ago. Note the lack of a mane: was this a group of females or was Spelea mane-less? (Source)

Spelea prowled the mammoth steppe — an open biome that was sort of like a savanna, but during the Ice Age and with bears and other creatures you’re not likely to ever see in Africa.

This big cat also was around during the last interglacial period, which was a little warmer than the one we’re currently in; like modern lions, it was apparently very flexible in terms of habitat.

Spelea fossils have been found from England to China’s 400,000-year-old Peking Man site.

Isotopes suggest that its favorite food may have been reindeer, but during this age of gigantism, Spelea was apparently only an apex predator in areas where it was too cold for other large carnivores. (Bocherens)

And when the reindeer headed towards the Arctic at the end of the last Ice age, Spelea did not follow. It went extinct some 11,000 years ago.

320px-California_Sabre-tooth_&_American_Lion_01
Atrox (right) and one of the other cats in the neighborhood — Smilodon, the sabercat. (Joe Mabel, via Wikimedia, CC BY-SA 3.0)

Fossils of Atrox, the “American lion,” were first found in Natchez, Mississippi, in the late 1850s. It soon became apparent that this big cat had roamed all over North America during the late Pleistocene, roughly 50,000 years ago.

Some top paleontologists consider Atrox a lion (see references in King); others think it was more closely related to the jaguar, another American cat. (Christiansen)

The problem is that carnivores are much fewer in number than their prey, and they tend to die in conditions that promote decay rather than fossilization.

No wonder that most of the cat family’s evolutionary history probably hasn’t made it into the fossil record! (Johnson et al.)

Whatever their evolutionary history, modern lions symbolize wild nature today. Their numbers in India, while low, have doubled over the last 10 years. In Africa, conservationists, governments, and local people are working out ways to protect Leo and the human beings who must live alongside him (and her).

Certainly it’s a challenge. But a hallmark of both cats and people is never shrinking from a challenge.

A terribly beautiful challenge . . .




Featured image: Derek Keats, CC BY 2.0


Sources:

Bauer, H.; Packer, C.; Funston, P.F.; Henschel, P.; and Nowell, K. 2016. Panthera leo (errata version published in 2017). The IUCN Red List of Threatened Species 2016: e.T15951A115130419.

Bocherens, H. 2015. Isotopic tracking of large carnivore palaeoecology in the mammoth steppe. Quaternary Science Reviews, 117: 42-71.

Cat Specialist Group. 2019. African lion. http://www.catsg.org/index.php?id=108 Last accessed March 5, 2019.

___. 2019. Asiatic lion. http://www.catsg.org/index.php?id=113 Last accessed March 5, 2019.

Christiansen, P. 2008. Phylogeny of the great cats (Felidae: Pantherinae), and the influence of fossil taxa and missing characters. Cladistics, 24(6): 977-992.

Craft, M. E. 2010. Ecology of infectious diseases in Serengeti lions, in Biology and Conservation of Wild Felids, ed. Macdonald, D. W., and Loveridge, A. J., 263-281. Oxford: Oxford University Press.

Ewer, R. F. 1998. The carnivores. Cornell University Press.

Figueiró, H. V.; Li, G.; Trindade, F. J.; Assis, J.; and others. 2017. Genome-wide signatures of complex introgression and adaptive evolution in the big cats. Science Advances, 3(7), e1700299.

Johnson, W. E.; Eizirik, E.; Pecon-Slattery, J.; Murphy, W. J.; and others. 2006. The Late Miocene Radiation of Modern Felidae: A Genetic Assessment. Science, 311:73-77.

King, L. M. 2012. Phylogeny of Panthera, Including P. atrox, Based on Cranialmandibular Characters. Electronic Theses and Dissertations. Paper 1444. http://dc.etsu.edu/etd/1444

Kitchener, A. C.; Van Valkenburgh, B.; and Yamaguchi, N. 2010. Felid form and function, in Biology and Conservation of Wild Felids, ed. Macdonald, D. W., and Loveridge, A. J., 83-106. Oxford: Oxford University Press.

Kitchener, A. C.; Breitenmoser-Würsten, C.; Eizirik, E.; Gentry, A.; and others. 2017. A revised taxonomy of the Felidae: The final report of the Cat Classification Task Force of the IUCN Cat Specialist Group. https://repository.si.edu/bitstream/handle/10088/32616/A_revised_Felidae_Taxonomy_CatNews.pdf

Li, G.; Davis, B. W.; Eizirik, E.; and Murphy, W. J. 2016. Phylogenomic evidence for ancient hybridization in the genomes of living cats (Felidae). Genome Research, 26(1): 1-11.

Loveridge, A.; Wang, S. W.; Frank, L.; and Seidensticker, J. 2010a. People and wild felids: conservation of cats and management of conflicts, in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 161-195. Oxford: Oxford University Press.

Loveridge, A. J.; Hemson, G.; Davidson, Z.; and Macdonald, D. W. 2010b. African lions on the edge: reserve boundaries as ‘attractive sinks’, in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 283-304. Oxford: Oxford University Press.

Macdonald, D. W;, Mosser, A.; and Gittleman, J. L. 2010a. Felid society, in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 125-160. Oxford: Oxford University Press.

Macdonald, D. W.; Loveridge, A. J.; and Nowell, K. 2010b. Dramatis personae: An introduction to the wild felids, in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 3-58. Oxford: Oxford University Press.

Macdonald, D. W.; Loveridge, A. J.; and Rabinowitz, A. 2010c. Felid futures: crossing disciplines, borders, and generations, in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 599. Oxford: Oxford University Press.

O’Brien, S. J., and Johnson, W. E. 2007. The evolution of cats. Scientific American, 297 (1): 68-75.

Sunquist, M. and Sunquist, F. 2002. Wild Cats of the World. Chicago and London: University of Chicago Press. Retrieved from https://play.google.com/store/books/details?id=IF8nDwAAQBAJ

Turner, A., and M. Antón. 1997. The Big Cats and Their Fossil Relatives: An Illustrated Guide to Their Evolution and Natural History. New York: Columbia University Press.

Werdelin, L.; Yamaguchi, N.; Johnson, W. E.; and O’Brien, S. J.. 2010. Phylogeny and evolution of cats (Felidae), in Biology and Conservation of Wild Felids, eds. Macdonald, D. W., and Loveridge, A. J., 59-82. Oxford: Oxford University Press.

Werdelin, L., and Dehghani, R. 2011. Carnivora, in Paleontology and Geology of Laetoli: Human Evolution in Context, Volume 2: Fossil Hominins and the Associated Fauna, Harrison, T., ed., 189-232. Springer, Dordrecht.



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