Chapter 6: The Lion and the Mammoth-Steppe

Alan Turner has written the book The Big Cats. He is a paleontologist and Senior Lecturer in the School of Biological and Earth Sciences at Liverpool John Moors University, in England. The fine drawings and paintings in this book, Mauricio Antón has done. He is an artist and scientific illustrator based in Madrid. On Plate 7 of this book, there is a painting, with lions and mammoths and caribou in Alaska during the upper Pleistocene. It is autumn. The dry brown grass is perhaps up to 10 cm tall. There are a few patches of low shrubs: Dwarf birch (buckbrush, as we call it in the Yukon). Its leaves have turned red. This tundra-steppe, further back, on ground rising up a little higher, is covered at a few spots with thin snow. There are no trees and tall shrubs whatsoever.

The woolly mammoth and the lion have certainly lived in Yukon-Alaska and northern Siberia. But in what kind of a climate? And on what kind of a plant-cover? Could the mammoth have found enough to eat up there in an arctic climate? And could it have found up there in an arctic climate enough to drink? - Could the lion have lived up there in an arctic climate? Is an arctic climate able to produced enough hoofed animals, to feed a residential lion, raising cubs? Could the "tundra-steppe" have supported the mammoth fauna? What have other workers found out about this?

American lion (Panthera leo atrox) attacking Equus lambei, a small Yukon horse. Illustration by Bonnie Dalzell. From: C. R. Harington, Neotoma (1990) National Museum of Natural Resources, National Museums of Canada, Ottawa.

C. R. Harington, in Ottawa, is Canada's leading Pleistocene paleontologist. He says on his homepage Animals of Beringia (1996) about the American lion: "American lions (Panthera leo atrox) were among the largest flesh-eating land animals that lived during the Ice Age ... in America. They ranged from Alaska and Yukon as far south as Peru. Because so many well-preserved specimens (over 80 individuals) have been found in tar pits at Rancho La Brea (Los Angeles, California), we have an excellent idea of their body structure. And unique, vivid glimpses of their past, about 40,000 to 10,000 years ago, have been recorded in European caves by our Paleolithic ancestors. Intriguing details in these artistic works suggest that the lions of Eurasia and America differed in some features from the living African lion.

"American lions were characterized by their enormous size and relatively long, slender limbs. Males were nearly 25 percent larger than male African lions. According to calculation of body weight based on femoral (thigh bone) size, male American lions would have averaged about 235 kg, females about 175 kg. The lion was adapted to steppe-like terrain. From about 300,000 to 10,000 years ago, cave lions (Panthera leo spelaea) lived in steppelike and parkland regions in the north and semi-desert areas in the south of Eurasia. Evidently they were not adapted to dense forests or deep snow. Fossils have been found as far west as England and as far east as the Alazeya River in Siberia some 1,600 km west of the nearest known specimens of the American lion (Kaolak River, northern Alaska). That distance is of little significance considering the wide-ranging habits of those lions, their adaptability to cool climate and the fact that they were able to pursue bison, horse and mammoth herds spreading across the grassy Bering Istmus which existed during late Pleistocene glaciations.

"Therefore, the cave lion stock that gave rise to American lions probably entered Alaska from Siberia during the second-last (Illinoian) glaciation. Lions had penetrated the North American plains by late Illinoian or early last (Sangamonian) interglacial time. As ice of the last (Wisconsinan) glaciation (about 80,000 to 10,000 years ago) spread, American lions were isolated in unglaciated parts of the northwest (Eastern Beringia) and south. Toward the end of the last glaciation, lions ranged southeast to Florida and as far south as Mexico and Peru. Perhaps rather dense forests prevented their entering eastern Canada and the northeastern United States.

"In Canada, most American lion remains have been found in Yukon deposits of last glacial age. The best of the fossils is a virtually complete skull from the Dawson City area. Other Yukon fossils are from Old Crow Basin and Bluefish Caves (north of the Arctic Circle), Sixtymile, Dublin Gulch and Big Creek. Lions seem to have been more common and widespread in the Yukon than other large predators of the time, such as the short-faced bear and the scimitar cat (Hometherium serum a relative of the sabretooth). Other Canadian fossils are from Edmonton, Blindloss and Medicine Hat in Alberta. ... According to their anatomical structure, American lions were at least as fleet as African lions, which are able to reach speeds of 48 km per hour in bursts when hunting. They may have been best adapted to feeding on bison. ... Probably they also hunted small horses." Harington, C. R. (1996).

Result: The male American lion averaged about 235 kg, and the female about 175 kg. That is the average body weight of the lion, not its largest weight. This cat was living in North America up to the shores of the Arctic Sea. Its remains one has also found in northern Alaska and the northern Yukon Territory. Could this large cat really have lived up there during the height of the Last Glaciation, as C. R. Harington believes? Would it have found there enough to eat? How much ungulate prey biomass does the American lion need, when raising cubs? Could arctic tundra or tundra-steppe have fed those large herds of hoofed animals, needed to support the lion? Let us start here with today's African lion.

African Lion

The American wildlife biologist George B. Schaller states in his book The Serengeti Lion (1972): The average adult lioness weighs about 120 kg, and the male lion about 170 kg. The adult lioness needs 5 kg food per day, and the male 7 kg. 33% of the average carcass, devoured by the lion, it does not eat, like large bones, stomach contents. A moderately active lioness needs about 5 kg/day, a male 7 kg. The average African adult male lion, weighing 170 kg, needs 2,555 kg meat/year. 33% of the prey is inedible. So he must kill 3,813 kg ungulate prey/year. - The average adult lioness, weighing 120 kg, needs 5 kg meat/day, and 1,825 kg meat/year. She must kill then 7.463 kg prey/day and 2,723 kg/year.

The residential lion is living in a pride (or group). This pride is made up of the ruling male lion, of several adult lionesses, and their cubs. They are living on a home range, controlled and defended by the ruling male lion. The young male lions, when mature, must leave their pride and their home range. They become then nomads. The nomadic lion will only become a residential lion, raising cubs, when he has found a place, where enough hoofed animals are living: where the ungulate prey biomass is large enough. That is: where it is also large enough during the lean time of the year. Any surplus during the rest of the year is irrelevant. How large must this ungulate prey biomass be?

We shall be able to answer this, if we do treat the residential lion like the residential tiger. Each residential tigress has her own home range. And two, three or more female home ranges are controlled by the ruling male tiger. That is: His home range overlaps with those of his females. Thus, I do assume here, that also the residential lion is not living in a pride, together with several adult lionesses and their cubs on his home range. I do assume now, instead, that also the residential lion controls the separate home ranges of 3 lionesses. From each one of these 3 female home ranges, he is taking about 1/3 of his food. How much ungulate prey biomass will the residential lioness then need, when raising 2 cubs, when these cubs are 6 months old (in their first year), and 1.5 years (in their second year of life)? Her home range is here 18 km². "kg BW/day" = kilogram body weight a day. "% BW/day" = percent body weight per day.

African Lioness with 2 cubs, 6 months old

Male lion, 170 kg BW, free-ranging. He needs 7 kg meat/day = 4.118% BW/day. That is 2,555 kg ungulate biomass/year (wet wt). He is eating 67% of his prey. He must kill then 6.145% BW/day ungulate prey = 10.446 kg/day, and 3,813 kg/year live prey. One third of this prey, he takes from the home range of this lioness: 1,271 kg/year.

Lioness, 120 kg BW, non-reproducing, free-ranging. She must eat 5 kg meat, organs/day (wet wt), 4.167% BW/day. That is 1,825 kg/year. She must kill 7.463 kg ungulate prey/day: 6.219% BW/day. That is 2,724 kg/year (with 33% waste).

One 6-month-old male cub, 20 kg BW. It needs 5.312% BW/day meat (wet wt): 1.062 kg/day. It needs then 7.928% BW/day live prey: 1.586 kg/day, and 579 kg/year live prey.

One 6-month-old female cub, 20 kg BW. It needs 4.780% BW/day meat: 0.956 kg/day, and 7.134% BW/day live prey: 1.427 kg/day, and 521 kg/year.

The whole African lion family, with two cubs in their first year of life, must kill at least 5,095 kg ungulate biomass in one year. That is about 9% of the total prey biomass. This means: There must be then a prey biomass of hoofed animals of 56,611 kg on their 18 km² home range. That is a lean season prey biomass of hoofed animals of 3,145 kg/km². This lion family must kill at least 47 wildebeest (weighing 108 kg each) in one year, or 31 zebras (164 kg each), or 12 buffaloes (420 kg each). The individual body weights are 3/4 of the average female, according to G. B. Schaller (1972).

African Lioness with 2 cubs, 1.5 years old

Male lion, 170 kg BW. He is taking 1/3 of his food from the female's 18 km² home range: 1,271 kg/year live prey.

Lioness, 120 kg BW. She needs at least 2,724 kg/year live prey, when non-reproducing.

One 1.5-year-old male cub, 124 kg BW. He needs 4.476% BW/day meat: 5.550 kg/day. And he needs 6.680% BW/day live prey (with 33% waste); 8.283 kg/day, and 3,023 kg/year live prey.

One 1-5-year-old female cub, 95 kg BW. It must eat 4.270% BW/day meat, 4.056 kg/day. And it needs 6.373% BW/day live prey: 6.054 kg/day, and 2,210 kg/year live ungulate prey.

The whole African lion family, with two cubs in their second year of life, must kill then at least 9,228 kg hoofed animals in one year. That is about 9 percent of the total prey biomass on their home range: a lean season prey-biomass of 102,533 kg/18 km², or 5,696 kg/km². This lion family must kill then in one year at least 85 wildebeest (108 kg each), or 56 zebras (164 kg each), or 22 buffaloes (420 kg each).

This means: The residential lion will only be able to raise its two 1.5-year-old cubs on its 18 km² home range, if there is a prey biomass of hoofed animals of at least 5,696 kg/km². That is, why the nomadic lion will only set up its home range, where there are many hoofed animals. Important to the residential lion is here only the ungulate prey biomass during the lean season of the year. Any surplus during the rest of the year is irrelevant.

Lions (Panthera leo atrox) in Central Alaska, during the time of the woolly mammoth. They have killed a steppe-bison bull and are feeding now on it. Drawing from: R. D. Guthrie, Frozen Fauna of the Mammoth Steppe (1990) Fig. 3.16.

The Lion of the Mammoth Steppe

Could the Eurasian cave lion, ranging up to the shores of the Arctic Sea, really have lived up there during the height of the Last Glaciation? And would the American lion, hunting up to the northern coast of Alaska and the Yukon, really have lived up there during the height of the Last Glaciation? Would the arctic-steppe, mammoth steppe, or periglacial steppe have supported those herds of hoofed animals? How large must the ungulate prey biomass in Alaska/Yukon have been, so that the American lion could have lived up there is a resident, raising its cubs?.

The average body weight of the adult American male lion was about 235 kg, and that of the female about 175 kg (C. R. Harington, 1996). We shall assume here now: The residential male American lion controls the home ranges of three females. He is taking 1/3 of his food from the home range of each one of his females. The home range of the American lioness in this model is also 18 km². She raises two cubs. How much food must the mammoth steppe of Alaska and the Yukon Territory then have produced? In what kind of a climate could those herds of hoofed animals have lived up there? And in what kind of a climate could the zonal mammoth steppe or savanna-like forest-steppe have grown up there?

American Lioness with 2 cubs, 6 months old

Male lion, 235 kg BW, free-ranging. He needs 9.677 kg meat/day = 4.118% BW/day. That is 3,535 kg year (wet weight). He is eating 67% of his prey. He must kill then 6.145% BW/day ungulate prey = 14.441 kg/day, and 5,271 kg/year live prey. One third of this prey, he takes from the home range of this lioness: 1757 kg/year.

Lioness, 175 kg BW, non-reproducing, free-ranging. She must eat 7.292 kg meat, organs/day (wet wt), 4.167% BW/day. That is 2,662 kg/year. She must kill 10.883 kg ungulate prey/day: 6.219% BW/day. That is 3,972 kg/year (with 33% waste).

One 6-month-old male cub, 28 kg BW. It needs 5.312% BW/day meat (wet wt): 1.487 kg/day. It needs then 7.928% BW/day live prey: 2.220 kg/day, and 810 kg/year live prey.

One 6-month-old female cub, 28 kg BW. It needs 4.780% BW/day meat: 1.338 kg/day, and 7.134% BW/day live prey: 1.997 kg/day, and 729 kg/year.

The whole American lion family, with two cubs in their first year of life, must kill at least 7,268 kg ungulate biomass in one year. That is about 9% of the total prey biomass. That is, when raising two cubs in their first year of life. This means: There has to be then a lean season prey biomass of hoofed animals of 80,755 kg on their 18 km² home range. That is a prey biomass of hoofed animals of 4,486 kg/km². If living in Africa, the American lion family must kill in one year at least 67 wildebeest (weighing 108 kg each), or 44 zebras (164 kg each), or 17 buffaloes (420 kg each). The individual body weights are 3/4 of the average female, according to G. B. Schaller (1972).

American Pleistocene lion, Panthera atrox. From A. Turner and M. Antón, The Big Cats (1997:76) Fig. 3.30.

American Lioness with 2 cubs, 1.5 years old

Male lion, 235 kg BW, free-ranging. He is taking 1/3 of his prey from the female's 18 km² home range: 1,757 kg/year live prey.

Lioness, 175 kg BW, free-ranging. She needs at least 3,972 kg/year live prey, when not reproducing.

One 1.5-year-old male cub, 170 kg BW. He needs 4.476% BW/day meat: 7.610 kg/day. And he needs 6.680% BW/day live prey (with 33% waste): 11.356 kg/day, and 4,144 kg/year live prey.

One 1.5-year-old female cub, 120 kg BW. It must eat 4.270% BW/day meat: 5.124 kg/day. And it needs 6.373% BW/day live prey: 7.648 kg/day, and 2,791 kg/year live ungulate prey.

The whole American lion family, with two cubs in their second year of life, must kill then at least 12,664 kg hoofed animals in one year. That is about 9 percent of the total prey biomass on their home range: 140,711 kg/18 km², or 7,817 kg/km². The American lion family must kill then in one year at least 117 wildebeest (108 kg each), or 77 zebras (164 kg each), or 30 buffaloes (420 kg each). This means: The residential lion will only be able to raise its two cubs in their second year of life on its 18 km² home range, if there is a lean season prey biomass of hoofed animals of at least 7,817 kg/km². In an arctic climate this is not possible.

European cave lion, Panthera leo, from the time of the woolly mammoth. After A. Turner and M. Antón, The Big Cats (1997:78) Fig. 3.31. The male lion of the mammoth fauna in Eurasia and North America does not seem to have had a mane like the African lion of today.

Ungulate biomass in Northern Hemisphere today

The Canadian Professors of Biology R. J. Hudson and F. L. Bunnel (1980) Table 11.2, have found out, how much ungulate biomass the different ecosystems of northern North America and northern Eurasia are able to support. They also show us, how much the ungulate biomass increases there per year. This is important, because the lion or any other large predator, can harvest only as much, as the herds of hoofed animals do increase in one year. If the predator took more, it would soon starve to death.

·         The open boreal woodland (taiga) in the northern hemisphere (in northern Europe, Siberia, Alaska, and northern Canada), where caribou/reindeer and moose/European elk are living, has a large ungulate biomass of 200-500 kg/km², or an average 350 kg/km². The ungulate biomass increases by 100 kg/km² a year.

·         The forest-tundra, with its caribou/reindeer, has a ungulate biomass of 50-200 kg/km², or an average 125 kg/km². Ungulate biomass increase per year, 10 kg/km².

·         The arctic tundra (Barren Grounds), with its caribou and musk-oxen, has a ungulate biomass of only 18-26 kg/km², or an average 22 kg/km². Ungulate biomass increase per year, 2 kg/km².

·         The polar desert, with its musk-oxen and Peary caribou, has an ungulate biomass of only 0.0-10 kg/km², or an average 5 kg/km². Ungulate biomass increase a year, 0.0-0.5 kg/km².

The cave lion of northern Siberia was not able to live up there during the Last Glaciation. And the American lion was not able to live in Yukon/Alaska during the Last Glaciation. It would have starved to death up there within a few weeks, if not days. The assumed adaptation of the cave lion and American lion to an arctic climate is not science, only science fiction. One should give it up now and look for a better explanation.