"Big cat"? "Freeloader"? Let's take a look at the ancient history of the "king of the grassland"

There is an animal that is both familiar and unfamiliar to people. It is familiar because of its image features, which can be found in the writings and cultural relics of many civilizations and in many zoos. It is unfamiliar because people have misunderstandings about its living habits and distribution areas, and do not understand its genealogy and evolution.

It is the lion Panthera leo . Like the tiger ( Panthera tigris ), it is one of the largest cats on earth and a well-known top predator. Today, let's talk about the lion's family tree and recipes~

Modern lion, photographed by the author at Tianjin Natural History Museum

What is the relationship between lions, tigers and leopards?

Lions belong to the subfamily Felinae under the family Felidae of the order Carnivora - the tribe Panthera - the genus Panthera. The panther tribe separated from the cat family, which includes domestic cats, cheetahs, and manuls, about 12 million years ago.

The tiger, leopard ( Panthera pardus ), jaguar ( Panthera onca ) and snow leopard ( Panthera uncia ) are also its relatives in the Panthera genus. The most recent common ancestor of the five relatives was born more than 4 million years ago, and then split into two branches.

One is the evolutionary branch of Panthera in Eurasia, including tigers and snow leopards, and the other is the evolutionary branch of Panthera in Africa - lions, leopards and jaguars , which later came to other continents during the Ice Age, just like ancient humans.

Different skeletal anatomical features of cats and leopards. Source: Classification of Felidae - Molecular Systematics Framework and Fossil Evidence Author: Zhang Kai, Shen Xueli, Liu Kaichou, Jiang Hao, Jiang Zuoqigao DOI: 10.13859/j.cjz.202301001

3.7 million years ago, some areas in eastern Africa had already transformed from forests to relatively dry grasslands. At this time, the lion's ancestor, the proto-leopard ( Panthera principalis ), also walked out of the forest.

The average size of the original leopards during this period was no less than that of modern lions. 2.4-1.7 million years ago, the Shawi lion ( Panthera shawi ), which was also no less than the modern lion, appeared on the grasslands of East Africa.

In addition, the lion family also parted ways with the leopard 2.6 million years ago, gradually moving towards the modern lion , and began to compete with members of the saber-toothed tiger subfamily such as the Homotherium and Megantereon that occupied the African grasslands at the time.

Restoration of the Shashi lion, source: prehistoric-fauna.com

About 600,000 to 800,000 years ago, a group of lions that were more suitable for the grasslands that expanded during the Ice Age emerged and entered Eurasia. This was the fossil lion, which weighed up to 400 kilograms and was the largest cat in history.

After entering Eurasia, the fossil lion underwent morphological changes in its incisors and jaws, becoming a cave lion ( Panthera spelaea ) that was moderately reduced in size but could still grow to two to three hundred kilograms or even larger.

Cave lions were the kings of the north in Eurasia, Alaska, and western Canada during the Ice Age, and even Northeast China was part of their range. Scholars speculate that compared to modern lions, they may have lived alone or in small groups.

Comparison of the size of fossil lions, Homo sapiens and modern lions. Source: prehistoric-fauna.com

The green part is the late Pleistocene and historical distribution area of ​​modern lions, the blue part and the orange part are the distribution areas of brutal lions and cave lions respectively. Source: Wikipedia

Some fossil lions evolved into Panthera youngi in eastern Asia, which was not a tiger but a relative of modern lions. It once threatened the survival of Peking Man in Zhoukoudian, Beijing.

There was also a group of lions that expeditioned to the heartland of the North American continent and South America during the last few hundred thousand years of the Ice Age. These were the atrocious lions ( Panthera atrox ). Their average weight was slightly larger than that of the cave lions. Their English name was American lion, a real cougar.

The cougar ( Puma concolor ) we often talk about today is not a lion - it is an animal in the same family as the domestic cat and is very distantly related to the lion.

Where is the modern lion?

Comparison of the size of a brutal lion and a human, source: prehistoric-wildlife.com

Yang's tiger image source: cnfossil.com

Note: The modern lion Panthera leo and the above-mentioned Yang's tiger, cave lion, brutal lion and fossil lion can all be collectively called lions, but there is controversy in the biological community as to whether they are the same species. Of course, in addition to modern lions, other extinct lions are often called "maneless lions". This is related to their lack of manes. According to genomic speculation, the hair of modern lions evolved independently after the separation from maneless lions, which appeared about 320,000-190,000 years ago.

While on expeditions, lions also have a home-guarding force, which is the modern lion ( Panthera leo ). All existing lions were divided into two major branches about 250,000 years ago. One branch is mainly distributed in southwest Asia, central and western Africa (there are also some in Ethiopia and the Democratic Republic of the Congo in the east, and northern Africa is also their historical distribution area), which is the northern lion subspecies ( Panthera leo leo ).

The northern lion subspecies is the lion that Manjushri Bodhisattva sits on, and is also the prototype of the Sphinx. It is also the lion in Aesop's fables and the lion found from other places in the ancient Roman Colosseum - after all, the background of these stories is set in the distribution area of ​​the northern lion subspecies.

The blue-green circle and the dark yellow-green circle in the upper right corner of the picture are the existing distribution areas of northern lions, and the other color circles are the distribution areas of southern lions. There are two subspecies of lions, and each subspecies has three branches. Image source: universiteitleiden.nl

Note: Lions have been divided into multiple subspecies in traditional taxonomy, but these subspecies are often based too much on morphology (such as mane). The dichotomy mentioned in this article is mainly based on the genome.

The other group is distributed in the savannas, deserts and even the edge of volcanic craters in southern and eastern Africa. This is the southern lion subspecies, which is often seen in documentaries. The average weight of the southern lion is greater than that of the northern lion and there are larger male alliances (when the young male lion grows up and leaves home, he and his brothers form a "gang", a male group, which includes his brothers and sworn brothers).

But overall, their weight is roughly between 100-270 kg, which is smaller than the brutal lion and cave lion, and smaller than their predecessors who belong to the same modern lion but are older.

Middle and Late Pleistocene modern lion fossils from Kenya Source: cambridge.org

From the last tens of thousands of years of the Ice Age to the beginning of the end of the Ice Age (about 12,000 years ago), modern lions expanded from Africa to the Eurasian continent (mainly to southwestern Asia), and even came to Sri Lanka to form the Sri Lankan lion subspecies ( Panthera leo sinhaleyus ).

Modern lions, who have both defended their homeland and expanded slightly, have achieved a distribution across four continents, America, Africa, Europe and Asia, together with their overseas relatives.

Facing a crisis

At the end of the Ice Age, along with the impact of human activities, climate change and prey reduction, lions were pushed back to western Asia, southwestern Asia and the African continent, and the brutal lion and cave lion became extinct, leaving only the modern lion.

Modern lions should be the southern lion subspecies, photographed by the author at Shijiazhuang Zoo

In the era of human civilization, lions in western and southwestern Asia were hunted extensively and eventually became extinct (only more than 500 wild lions remain in India's Gir National Park). In the first half of the last century, lions in northern Africa also disappeared, and the lion population in central and western Africa was even reduced to only a few hundred.

The situation of the southern lion subspecies has improved a lot, but some disease threats (such as tuberculosis), as well as poaching and human-animal conflicts still threaten its survival.

The northern lion subspecies in India is small in number

Image source: www.insideindianjungles.com

Public protests when Zimbabwe's lion star Cecil was killed in 2015

Image source: https://www.cbsnews.com/pictures/cecil-the-lion-killed-by-hunter/

There are only tens of thousands of wild lions left on Earth today, which is still considerable compared to the less than 10,000 wild tigers and snow leopards. However, the threat has not stopped, and how to protect them remains a problem.

What do lions like to eat?

The majestic predator lion has thick canine teeth, short and wide jaws, and relatively flat supraorbital processes on the skull. It also has large, elastic vocal cords and corresponding hyoid bone structures to help it make the "Hedong Lion Roar".

What do these majestic guys eat?

It depends on what delicious food there is where the lion lives.

The average weight of lion prey is the largest among cats. After all, various big guys such as plains zebras, African buffaloes and giraffes will become their staple food (in general, lions like prey between 190-550 kg).

Compared with female lions, the average weight of male lions' prey is larger. It should be said here that "male lions do not hunt" is a rumor. Some "male lions with many wives and concubines" and their brothers will live off their wives, but most male lions with wives and concubines still hunt on their own. They will also help their partners to deal with large prey such as African buffaloes, while gang male lions who don't have a partner have to rely on themselves.

A male lion preys on a warthog. Source: earthtouchnews.com

Attacking wildebeest, source: zoopensefull.com

According to a survey in Hwange National Park, African buffalo accounts for 56% of the diet of male lions, which is higher than the 33% of female lions. In the late dry season, female lions will choose warthogs and zebras as prey, while in the early dry season, they will prey on giraffes, etc. A survey in Kruger National Park found that male lions prefer large African buffaloes, while female lions prefer relatively small plains zebras and wildebeests.

Cooperation in capturing buffalo, source: mediadrumworld.com

Lions in other places are different. In the dry season in Botswana and other places, lions will hunt young African elephants. The main food of lions in Gir National Park in India is the chital deer ( Axis axis ).

If placed in the desert near the sea in Namibia, the local cormorants and penguins, as well as the sea lions on the beach become the main course of the lions - there are not many antelopes and zebras in the area.

When hunting, they will bite off the cervical vertebrae of smaller prey, and suffocate larger prey by biting their throats or mouths and noses. If they are facing large prey such as giraffes, they will attack from behind. When dealing with African buffaloes, they will fight until they are exhausted before preying on them.

The sprint speed of a lion is between 45-60 kilometers per hour. In 2018, there were records of lions running at a speed of 78 kilometers per hour. This speed is enough to shock many prey. Of course, the lion's slender canine teeth, masseter muscles, limbs and claws can tell the prey that death is coming.

Note: Except for snow leopards, other Panthera species can also use the "Hedong Lion Roar" trick, which is to make a deep roar.

Young male lion, photographed by the author at Beijing Zoo

Lion skull model, taken by the author at the Shanghai Natural History Museum

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Zhang Kai, Shen Xueli, Liu Kaichou, Jiang Hao, & Jiang Zuoqigao. (2023). Classification of Felidae: A molecular systematic framework and fossil evidence. Journal of Zoology, 58(1), 1. Author: Daqing Shenlong

Editor: Dong Xiaoxian

Reviewer: Liu Kun and Li Peiyuan