Chapter 5: Spotted Hyena

Also the remains of the spotted hyena (Crocuta) they have found in Eurasia, together with the bones of other mammals of the mammoth fauna. When has the spotted hyena lived in Eurasia? How far north? In what kind of a climate and on what kind of a plant-cover has it lived there? What has it eaten?

Vereshchagin and Baryshnikov: "The range of the cave hyena covered all of Europe and the subtropical and middle latitudes of northern Asia, reaching to about 56 degrees N in Siberia. It apparently did not penetrate the arctic zone, nor did it occupy high mountains, since its remains have not been found in alpine sites in the Caucasus (Kurado, Tsona). If the concept of a single African-Eurasian species is correct, then it did not become extinct, but merely suffered a catastrophic shrinkage of the northern Eurasiatic portion of its range. The cave lion presents a potentially analogous case.

"Massive accumulations of cave hyena bones, together with coprolites [dung] and bones of prey animals, have been found in many European caves. For example, Kirkdale Cave in England provided abundant remains of hyenas of all ages, from newborn to old, while Tornewton Cave supplied more than 20,000 cave hyena teeth (Kurtén 1968). In the USSR hyena bones are common in the caves in the Altaij (Hyena Den Cave) and the Far East (Geographic Society Cave) (Ovodov 1980). The species denned in caves and sometimes died there, vanishing into karst wells (Ageeva et al. 1978).

"Like the African spotted hyena, the cave hyena fed primarily on carcasses of large ungulates and thus completed the stage of ‘secondary biomass’ utilization, prior to final destruction by larval insects and bacteria. It is also possible that it hunted rodents, young ungulates, and mammoths. Reduction in the numbers and range of the cave hyena coincided with the disappearance of herds of steppe and forest-steppe ungulates at the boundary between the Pleistocene and the Holocene, when the plains became boggy and the taiga developed."(1984:496, 497).

Map: Late Pleistocene range of the spotted hyena (Croculta crocuta spelaea). From: R.-D. Kahlke (1994) Fig. 9. In Central Siberia the spotted hyena went up to about 65°N. Today, it is living 5400 km further south in latitude. The northern limit of its range in Africa lies now in the Sahel zone, near 15°N, south of the Sahara Desert. It is adapted to the dry steppe and forest-steppe, where large herds of hoofed animals are living. Below map: Reconstruction of glacial-age hyena (Crocuta spelaea). After: W. von Koenigswald (1983:199) Fig. 101.

 

Spotted Hyena and late Pleistocene Climate

The spotted hyena or cave hyena has lived in Eurasia till the end of the late Pleistocene. How far north has it lived there? In what kind of a climate, and on what kind of a plant-cover? - It died out, when the herds of steppe and forest-steppe ungulates disappeared there. How cold is it supposed to have been during the peak of the Last Glacial in Eurasia, where the spotted hyena has lived together with the woolly mammoth and other large hoofed animals? How warm (or cold) is it there now? And in what kind of a climate is zonal steppe and forest-steppe able to grow? How far north the spotted hyena has lived in Eurasia during the late Pleistocene, R.-D. Kahlke (1994) map Fig. 9 shows us.

Central Europe. The spotted hyena has also lived on the Russian Plain, up to 57-58°N, 50°E. Now 18°C July temperature. Then 8°C colder = 10°C July temperature. Now 0°C mean annual air temperature. Then 10°C colder = -10°C mean annual air temperature, with 350 m thick permafrost. Permafrost temperature then –7°C x 1.32 = -9.24°C annual air temperature. Now coldest month –14°C. Then –20°C colder = -34°C coldest month. Now 500 mm annual precipitation. Then 375 mm ppt/yr less = 125 mm ppt/yr in Last Glacial.

West and Central Siberia. In West and Central Siberia the remains of the spotted hyena one has found up to 59°N, 60-90°E. It is supposed to have lived there also during the Last Glacial. – In what kind of a climate? Now 14°C July temperature. Then 7°C colder = 7°C July temperature. Now –3°C mean annual air temperature. Then 9°C colder = -12°C mean annual air temperature in Last Glacial. 400 m thick permafrost. Permafrost temperature then –7°C x 1.32 = -9.24°C mean annual air temperature, with ice wedges. Coldest month there now –22°C. Then 14°C colder = -36°C coldest month. Now 500 mm ppt/yr. Then 375 mm less = 125 mm/ppt in Last Glacial.

Glacial Maximum in Europe, Plant Production

How much aboveground vegetation (dry matter) has grown on the steppe-tundra or loess-tundra, with its intensive loess production in central and eastern Europe during the coldest part of the Last Glaciation (22,000 B.P.)? – The total aboveground biomass of the Glacial Maximum (22,000 B.P.) was about 121 g/m² (living and dead) dry weight. (A. K. Gliemeroth, 1995:22). The steppe-tundra has grown then about 17 gram aboveground green dry matter per square meter per year (17 g DM/m².yr).

Not even the reindeer could have lived there. The non-lactating reindeer/caribou needs at least 25 g DM/m² yr green vegetation for maintenance. The lactating reindeer needs 35 g DM/m² yr (Bliss, L. C. et al., 1981:440, 441). Nor could the bison have lived on the periglacial loess-steppe during the height of the Last Glaciation. The large 1250-kg bison bull is only able to live, where 189 g DM/m² yr is growing per year (Belovsky, G. R. 1986). And the elephant will starve to death, where only 200 g DM/m² yr has grown per year (Phillipson, J. 1975:171, 176).

Forest-Steppe. The Eurasian forest-steppe is growing now in this climate: 2°C mean ann. air temperature. 60 kcal/cm² solar radiation, 170 days potential vegetation period, 600 mm evaporation. 2125° 10°C.ts., 420 mm ann. precipitation. 1200 g DM/m² yr net primary production.

Zonal Steppe. The Eurasian zonal steppe is growing now in this climate: 5°C mean ann. temperature. 65 kcal/cm² yr solar radiation, 190 days potential vegetation period. 850 mm evaporation. 600-1000 mm/yr potential evapotranspiration. 2275° 10°C.ts. 330 mm ann. precipitation. 1100 g DM/m² yr net primary production. After: Bolin et al., (1979); Walter, H. (1968), Borisov (1965) - The ice-age theory, as now commonly taught, is not based on scientific facts. It is only science fiction.

The spotted hyena (Crocuta spelaea). From: T. Haltenorth and H. Diller, Säugetiere Afrikas (1977) Plate 38.4. The spotted hyena needs about 2 – 3 kg live prey a day. This includes 30% waste. It hunts mainly the wildebeest and the zebra. This animal is adapted to the dry, open savannah, and is able, to live for a long time without drinking any water.

One of the driest areas, where the residential spotted hyena is still able to live in Africa, lies in the Sahel zone, south of the Sahara Desert. A mean annual rainfall of 375 mm (250 – 500 mm) produces there about 2800 kg/km² prey biomass. This means. When the spotted hyena was living on the mammoth steppe of late Pleistocene Eurasia, there had to be then a large prey biomass of at least 2800 kg/km². This amount of prey biomass is only able to grow in a mild, temperate climate, where the growing season of the plants is long.

The arctic tundra (and steppe-tundra) is not able to support the spotted hyena, because there is too little ungulate biomass. The arctic tundra has only an average 22 kg/km² (18 – 26 kg/km²) ungulate biomass (R. J. Hudson and F. L. Bunnel, 1980:21). Because too little fodder is growing up there on the permanently frozen ground. This proves that the spotted hyena is not able to live in an arctic climate, and that it has not lived in late Pleistocene Eurasia in an arctic climate.

 

Recent Spotted Hyena

The spotted hyena (Crocuta) is living now wild only in Africa, south of the Sahara Desert. Where is it able to live there? What does it eat? And which kinds of animals is it mainly hunting?

Hans Kruuk, a Dutch biologist, has studied the spotted hyena in Africa. He reports: "I had four adult hyenas myself which were caught as adults and kept in cages of 3 x 6 m for seventeen days, and then released. These hyenas were fed an average of 3.5 kg per day each (meat including bone and hair) which they always eagerly finished completely. I was able to weigh only three of them before and after the experiment. Francis slimmed from 58.5 kg to 56.0 kg and Graham from 50.8 kg to 50.3 kg, and Hugh gained from 45.4 kg to 48.2 kg, all three on virtually the same amount of food Francis was very nervous and walked a great deal up and down the fence, whereas Hugh was the quietest of the four and lay down most of the time." (1972:75).

"They can go without food for days – for instance, the hyena with the radio collar in Ngorongoro at intervals of at least 2, 2, 1, 5 and 2 days. When they do eat, they consume large quantities in one session. Once I presented the remains of two female Thomson’s gazelle (all except intestines, stomach, liver, lungs, etc.) to a male hyena I met wandering on the plain, which I estimated to weigh between 45 and 50 kg. This animal looked healthy and did not have a full stomach, but I did not know how long ago it had its last meal. When presented with the meat, the hyena took off and within 45 min, ate14.5 kg of it.

"... the average daily killing rate in the dry season in the Ngorongoro is 2.0 kg per day. This includes meat scavenged by lions, jackals, and vultures, but only the amount of meat scavenged by lions is substantial in this wastage... Actual food consumption at the time in the crater must have been 1.5-1.8 kg per day.

"Ngorongoro, prey killed by spotted hyenas: Average per hyena per day: 2.0 kg per hyena per day for calculating the role of adult wildebeest, zebra, and so on, in the hyena’s diet. ... I estimate the daily killing rate in the Serengeti to be approximately 3 kg. I use the words ‘killing rate’ here in the sense of kilograms of meat appropriated, either killed or scavenged; the 3 kg per day also includes waste.

"In Ngorongoro, some 430 hyenas kill or scavenge 2 kg per day each; so total population disposes of 430 x 2 x 365 = 313,900 kg per year. Similarly, in the Serengeti, 3,000 hyenas disposing of 3 kg per day each would account for 3,000 x 3 x 365 = 3,285,000 kg per year." - Kruuk, H. (1972:76-81).

Condition of Prey Animals

Which kinds of animals does the spotted hyena hunt mainly for food? How does it find in a large herd those hoofed animals, which it could most easily capture?

Hans Kruuk: "I found it difficult to obtain information about the condition of the animals caught by the hyenas; carcasses are often consumed entirely and the vital organs disappear very rapidly after the prey has been killed. It is clear that hyenas will select a physically inferior animal from a group if they happen to see it, and they probably search for such prey. But most likely the hyena’s criteria are far more sensitive than mine, because I can rarely see anything different in the behavior of an animal that has been selected by hyenas. In a number of such instances, there was indeed something wrong with such selected quarry.

"The hyena’s selection and hunting methods cause them to select the least physically fit from the population, at least in wildebeest and gazelle; in zebra, the interaction is complicated by the active defense of the family group by the stallion. This by no means signifies that hyenas eat only very young, old, or diseased animals, but if these are available, they will be selected. In the Ngorongoro the latter categories are hardly present at all, and the hyenas have to be content there with what are probably perfectly healthy prey in the prime of their life." (1972:99, 101).

"In the range of available prey size, hyenas appear to take the medium-sized ungulates (wildebeest, zebra) rather than the larger (buffalo) or smaller ones (Thomson’s gazelle). Hyenas have not shown any obvious preference between wildebeest and zebra, and the presence of these species in the animal diet corresponds closely with the relative abundance.

"Hyenas would indeed be eminently suitable for a role in which they would adjust the ungulate’s number to the range conditions without large fluctuations in numbers. Any animal that, because of a slight undernourishment, was in worse condition than its neighbor, would be cropped immediately, and a rise in numbers of undernourished animals could be dealt with without delay. ... Similarly, a sudden improvement of the grassland might cause hyenas to catch fewer prey of one kind so they would have to switch to another." - Kruuk, H. (1972:102, 104).

Competition

How does the spotted hyena get along with its fellow-hyenas, for instance, when a large pack of them is devouring a carcass at the same time?

Hans Kruuk: "Competition between feeding hyenas expresses itself in speed of eating rather than in actual fighting, and the time in which a prey animal can vanish is truly amazing. To mention some examples: in Ngorongoro, a yearling wildebeest (estimated weight approximately 100 kg) was chased for 5 min by a hyena and died 2 min after being caught by 2 hyenas; 7 min after its death, one of the twenty-one hyenas eating from it ran off with a leg; another 3 min later only the head and spine were left; and 3 min more from the time of death there was only a dark spot on the grass to show what had happened.

"On another occasion, in the crater, hyenas killed, simultaneously, a female zebra and her two-year-old foal (estimated weights approximately 220 and 150 kg). Altogether, thirty-five hyenas ate from the two carcasses, clearing the spot of all remains of the two-year-old zebra in 24 min, and of the adult female in 36 min after their deaths..

"In the Serengeti, I have seen a gazelle fawn (approximate weight 2.5 kg) swallowed up by a single hyena in just under 2 min, and the largest amount of food I ever gave to a wild hyena (two disemboweled gazelle carcasses, of which the hyena ate 14.5 kg) disappeared in 45 min. Hyenas can swallow large pieces of meat or bone in one gulp, which must help them in consuming their food quickly." (1972:126).

Hunting Zebra

How is the spotted hyena hunting the zebra? And how fast is it able to eat it up?

Hans Kruuk: "By 19.07 hr seven more hyenas had been attracted by the commotion and there were fifteen hyenas following the zebras, but otherwise the picture was changed. Suddenly one hyena managed to grab a young zebra while the stallion was chasing another member of the pack. This young zebra, which was between nine and twelve months old, fell back a little, and within seconds twelve hyenas converged on it; in 30 sec they had pulled it down while the rest of the family ran slowly on. More hyenas arrived and the little zebra was completely covered by them. At. 19.17, 10 min after the victim had been caught, the last hyena carried off the head and nothing remained on the spot but a dark patch on the grass and some stomach contents. Twenty-five hyenas were involved, and the whole process of dismembering took exactly 7 min." (1972:177, 178).

"Zebras are hunted by packs of up to twenty-five hyenas, whereas wildebeest are chased by solitary hyenas or hyenas in groups of two or three, which are later joined by others. ... Before the hunt, hyenas have different ways of ‘testing’ wildebeest and zebra; wildebeest are often made to run by a hyena’s dashing into the herd; the hyena then stands watching the running prey. With zebra, a pack of hyenas usually merely walks close to a group and looks at them from nearby before the zebra start slowly drifting away or the zebra stallion charges the hyenas." (1972:201).

 

Spotted hyenas  hunting down a zebra, which they have first separated from its herd. They usually kill their prey by eating it. From: A. Turner and M. Antón, The Big Cats (1996:167) Fig. 5.13.

Prey Biomass

The spotted hyena is hunting mainly ungulates of medium size, like the wildebeest and the zebra. One spotted hyena, or groups of two or three chase the wildebeest. When they have caught the prey, other hyenas join them. The zebra, they are hunting in packs of up to 25 hyenas, as Hans Kruuk (1972:201) found out. – How much prey biomass does such a pack of 25 spotted hyenas need per day and per year? And how large must be then the ungulate biomass on their home range?

We shall assume here that one spotted hyena needs 2.5 kg prey biomass (with bones and waste) per day. 2.5 x 25 = 62.5 kg prey biomass per day x 365 = 22 812 kg prey biomass per year. If the ratio of prey to predator is balanced, the hyena is taking only the surplus, added each year through new growth of 10%. The total hoofed mammal prey biomass must be then 228,120 kg (of wildebeest and zebra). If the average zebra weighs 200 kg, the pack of 25 spotted hyenas must kill then 1141 zebras in one year.

In Eurasia, when the spotted hyena and the woolly mammoth were living there, the wild horse and the wild ass were taking the place of the zebra. Eurasia’s mammoth fauna was very similar to the fauna, living now on East Africa’s savannas.

In Africa, the spotted hyena is living now just as far north as the lion: up to the savanna-grasslands, south of the Sahara Desert. The residential lion, raising cubs, needs at least 1,000 kg/km² prey biomass. If there is less food, most the cubs will starve to death. Also the spotted hyena, raising cubs, is only able to live permanently, where there is a prey biomass of at least 1,000 kg/km².

The Kruger National Park, in South Africa, for example, has a prey biomass of 1,034 kg/km². The Etosha National Park, in Namibia, has a prey biomass of 2,873 kg/km². The Nairobi National Park, in Kenya, has 3,052 kg/km². The Serengeti, Tanzania, has 4,222 kg/km². Manyara, Tanzania, has 7,785 kg/km². And the Ngorongoro Crater in Tanzania has a prey biomass of 10,363 kg/km², as reported by G. B. Schaller (1972). The figure for Etosha is from H. B. Berry (1981). In all of these African parks, the spotted hyena is living together with the lion and other large carnivores.

The lowest possible prey biomass for the large predator fauna in these African parks (with its spotted hyena, lion, leopard, cheetah, and wild dog) is 100 kg/km² to 1 kg/km² of predator biomass. Only at this ratio of 100 kg/km² to 1 kg/km² is the prey/predator mass balanced. Spotted hyena, lion, leopard, cheetah, and wild dog are taking then not more prey biomass, than is being added each year through new growth in the herds of the hoofed animals: through the young calves, which are born each year.

Thus, when the spotted hyena, lion and mammoth were living in Eurasia, there had to be then also a prey biomass of at least 100 kg/km² per 1 kg/km² of predator. We must ask ourselves now: Could this prey biomass have grown in an arctic or subarctic climate, in arctic tundra, steppe-tundra or forest-tundra?

Prey Biomass and Annual Rainfall

How much large prey biomass is growing now in Africa at which amount of annual rainfall? How much is it raining per year at the spotted hyena’s northern range-limit, south of the Sahara Desert?

At the northern limit of the present range of the spotted hyena, south of the Sahara Desert, the following amount of rain is falling per year: In Senegal, NW Africa, 500 mm. In Mali 250-500 mm. West of Lake Chad 250-500 mm. At Lake Chad 250 mm. East of Lake Chad 500 mm. And in the Sudan, NE Africa, 250-500 mm.

R. Norman Owen-Smith, University of Witwatersrand, South Africa (1988:273) Fig. 14.3, has published the annual rainfall and total biomass of large herbivores in different African parks. It shows us, how much large prey biomass may grow at which annual amount of rainfall. It goes up to about 800 mm rainfall per year.

I have drawn the amount of rain and of large prey biomass of these different African parks on millimeter paper, to find out, how much large prey biomass is able to grow there per millimeter of rain. – At 250 mm rain per year in Africa, there will be up to 700 kg/km² of large prey biomass. At 300 mm ppt/yr 1,600 kg/km²; at 350 mm ppt/yr 2,400 kg/km²; at 400 mm ppt/yr 3,200 kg/km²; at 450 mm ppt/yr 4,000 kg/km², and at 500 mm ppt/yr 4,900 kg/km² large prey biomass (as an upper limit).

South of the Sahara, where the spotted hyena (and the lion) are still able to live (at the northern limit of their range), 250-500 mm of rain is falling per year. It produces 700 to 4,900 kg/km² of large prey biomass. The average rainfall of 357 mm (250-500) per year produces a large prey biomass of 2,800 kg/km². The nomadic spotted hyena, not raising any cubs, is able to live, where there is less large prey biomass than this. 

Arctic and Subarctic Prey Biomass

The advocates of the ice age hypothesis, as now commonly taught, do claim in all earnestness that the spotted hyena, the lion and the woolly mammoth have lived in Europe and Asia during the height of the Last Ice Age. These animals are supposed to have been adapted to a severe arctic climate. It was colder there, than it is now in northeastern Siberia, they say. These animals are supposed to have lived in Eurasia on arctic tundra, tundra-steppe or loess-steppe. So we must ask ourselves again: How much large prey biomass does the arctic and subarctic climate now produce? Would the spotted hyena have found there enough to eat? We have learned about this already, when studying the lion.

The Canadian Professors of Biology, R. J. Hudson and F. L. Bunnel (1980:21) Table 11.2, have found out: The open boreal woodland 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 northern forest-tundra, with its caribou/reindeer, has a large ungulate biomass of 50-200 kg/km², or an average 125 kg/km². The arctic tundra has an ungulate biomass of only 18-26 kg/km², or an average 22 kg/km². Some leading advocates of the modern ice age theory even claim that the mammoth fauna has lived in central Europe also during the peak of the Last Glaciation on a polar desert! The polar desert, with its muskoxen and Peary caribou, has now a large mammal prey biomass of only 0.0-10 kg/km², or an average 5 kg/km².

The Canadian biologist R. E. Redmann (1982:230) Table 4, states about the large herbivore biomass in the arctic tundra and forest-tundra in the Northern Hemisphere: Finland 430 kg/km²; Alaska, USA, 298 kg/km²; Norway 233 kg/km²; Siberia, Russia, 81 kg/km²; Yukon Territory, NW Canada, 80 kg/km²; Northwest Territories, central northern Canada, 36 kg/km² large herbivore biomass.

All this clearly shows me: The spotted hyena, the striped hyena and the cave lion were not able to live in an arctic or subarctic climate, in ice and snow. They were not able to live on arctic tundra, tundra-steppe, loess-steppe, forest-tundra, nor on any other kind of arctic plant-cover: They are not able to live in an arctic climate, because they would not find there enough to eat. They would have pitifully starved there to death within a few weeks or days! I have disproved this now quantitatively. This alone would be enough already, to disprove the whole ice-age theory, as now commonly taught, once for all times, and with it, the asserted adaptation of the woolly mammoth to severe arctic cold.