Chapter 2: The Woolly Rhinoceros

Where has the woolly rhinoceros (Coelodonta antiquitatis) lived in Eurasia, together with the woolly mammoth? How far north have they found its remains in Siberia? In what kind of a climate and on what kind of a plant-cover has it grazed up there? Was the woolly rhinoceros adapted to an arctic climate? What have scientists found out about this?

Professor N. K. Vereshchagin and G. F. Barishnikov (1982:271) say about the Woolly Rhinoceros (Coelodonta antiquitatis): "Bone remains, primarily of skull fragments of woolly rhinoceros are known from all of the landscape zones of northern Eurasia excepting the desert. The appearance and body proportions of the woolly rhinoceros were similar to those of the African white rhinoceros (Diceros simus Burchell, 1817), but differed, of course, in having a hairy coat. Short legs and hypertrophy (= very large size) of the horns were characteristic features, especially in the female. The front horn reached a length of 135 cm, and the posterior one, 50 cm. The woolly coat was soft and thick, dingy yellow and reddish brown and 10 or 15 cm long on the trunk. The ears were about three quarters the size of those of the living species. The hooves had a small bearing surface, evidently an adaptation to firm, solid substrates.

"Compressed woody remains of broadleaf species such as willow and alder and also mineral particles have been found in the deep alveoles of the upper teeth of the woolly rhinoceros. Lumps of excrement found in the area of the large intestine within the skeleton of a rhinoceros at Churapcha settlement in central Yakutia contained shoots of grasses, cottongrass, and sedges (Lazarev, 1977b). Pollen found in the faeces consisted of gramineae (89%), Compositae (4.5%), and Artemisia (2.5%). No data are available on their winter feeding habits, but it is quite clear that the woolly rhinoceros was predominantly a grazer. ... The woolly rhinoceros probably inhabited open areas in the cold and snowless steppe-tundra. In valleys and river floodplains it probably kept to shrub thickets." - Vereshchagin N. K. et al. (1982:271).

The woolly rhinoceros from Churapcha in central Yakutia had a shoulder height of about 160 cm. I had a chance, to measure it myself during the Mammoth Exhibition in Darmstadt, West Germany, at the end of November 1994. Most of the adult woolly rhinoceroses, whose remains have been preserved till now, do have a shoulder height of about 160 cm. The rhinoceros, 1.6 m tall at the shoulder, has now a body weight of 1100 to 1500 kg.

The Black rhinoceros (Diceros bicornis) in Tsavo East National Park, in Kenya, East Africa, during the great drought of 1970-71, has starved to death just as soon, as the African elephant. Several hundred of them starved to death with a full stomach, when only about 200 gDM/m² aboveground dry matter had grown there per year. The dry food in their stomach contained then only 2.0% crude protein (dry weight) or more.

Ralf-Dietrich Kahlke (1994:34) says about the woolly rhinoceros (Coelodonta antiquitatis): "During the last glacial complex, Coelodonta was distributed nearly across the whole area of northern and central Eurasia. Finds from Spain and the area of the Pyrenees, from France, England, Belgium, Denmark, the Netherlands and the southern North Sea and from Italy, Yugoslavia and Greece are known. Across central and eastern Europe its range continued to the Caucasus, and to N. and NE-Kazakhstan, and, going around the desert-areas, even till Kirghizia.

"Coelodonta was living in the eastern part of the Russian Federation, and still further east in the Ural and W-Siberia, and in the Baikal area, and northern Outer Mongolia till the Far Eastern Primoski Krai. In the south, it occupied Korea and the northern and northeastern provinces of China. Northwards, the range of this species reached Yakutia and the New Siberian Islands. But the northeastern-most part of the Eurasian landmass Coelodonta seems to have avoided, so that it did not cross over into Alaska.

"During the last Glacial, the most progressive and most robust forms of the woolly rhinoceros were spread out in a broad belt from Spain till NE-Siberia. The idea that C. antiquitatis was a steady companion of Mammuthus primigenius, and that it did have, therefore, similar environmental requirements, as this one had, is true for large parts of Eurasia. Differences do appear mainly north of the 70^th latitude. While finds of mammoths here are not rare at all, the remains of Coelodonta are here sparser or are lacking.

"North of the 45^th latitude, near the Pacific, no sure remains of Coelodonta have been reported. It did not live there. Also the lion Panthera leo spelaea did not live at the Pacific Coast, because it has been then more or less covered with forests, due to the oceanic influences. They did not find there the needed food. The southern limit of the last-glacial range of the woolly rhinoceros in Asia is about the same, as that of the range of the mammoth. Also Coelodonta was not able to move into extremely dry zones. Eastward of Lake Balkhash and across the E-Chinese loess-plateau, though, they were able to walk around them. Coelodonta populations have moved down to 32°N. That is, they have moved about 650 km further S, than the mammoths.

"GUERIN (1980, p. 1005) has compiled about 170 finds from W-Europe with C. antiquitatis. In about 65% of the cases, the rhinoceros was found together with Rangifer. More rarely it was associated with more fastidious kinds, like the wild boar (ca 31%) or even with the hippopotamus (ca. 5.9%). Hence, the woolly rhinocoeros was on the most important parts of its range mainly an animal of the open steppe-tundra." - Kahlke, R.-D. (1994:78).

Whole frozen Woolly Rhinoceros

Also whole frozen bodies of the woolly rhinoceros they have found in northern Siberia. The Baltic-German Baron Erich von Toll reports about a find at the River Khalbui. It was a woolly rhinoceros, not a Merck’s rhinoceros:

"The place, where the body of the Rhinoceros Merkii Jaeg. was found in the year 1877, is at the left bank of the Khalbui, a right-hand tributary of the Bytantai, about 15 verst (16.0 km) above the place, where it flows into the Bytantai. The geographical site should be near 68.2°N. I was lucky that I was led by an eye-witness to the place of discovery, by the son of the discoverer of the rhinoceros carcass, the Yakut Pawel Affanasiewich Gorokhov. He had himself seen the whole rhinoceros in the same position, as his father had seen it lying there. And he had himself helped his father, to chop off the head (which was preserved for science by the trader N. Gorokhov) and of one foot. The trunk of the carcass, the people had left there, and one year later, in spring, it was carried away by the floods!

From what Gorokhov told me, I was able to understand, how the carcass has lain there. At the cutbank of the river, he showed me exactly the height, at which the animal had lain, namely ‘on its belly, with its chin on the edge of the riverbed, while the rest of the body was still sitting at the wall of the cutbank’. ‘The carcass itself’, the eye-witness said, ‘was completely covered with fine sand.’" - Toll, E. (1895:36).

Woolly Rhinoceros at Wilyuy River

Sir Henry H. Howorth has mentioned already the woolly rhinoceros, found at the Wilyuy River in northeastern Siberia. What does Professor J. F. Brandt himself state about this? – Professor J. F. Brandt, Academy of Sciences at St. Petersburg, reports about his findings about a woolly rhinoceros, recovered at the Wilyuy River:

"When thoroughly examining the head of Rhinoceros tich. from the Wilyuy, I noticed that the blood vessels, taken out from its head, were filled up to the capillary vessels with a brown mass (clotted blood), which at some places still had the red color of blood, When seeing that the blood-vessels of the head were filled so much with the remains of small blood-balls, I was unable to suppress the thought that this individual, to which they belonged, has probably died through asphyxia, when it drowned.

"Helmersen and I myself saw on the remains of the Wilyuy rhinoceros two kinds of earth. The one, most common kind of earth, consists of microscopic quartz particles, bedded into a fine, clayey mud, with a few particles of mica. ... The second kind of earth was found only in the shape of spots at a few places on the head. It had a gray-blue color and is easily falling into a powder. It is, as I do assume, blue iron-earth (iron-blue) [vivianit]. Because, when Helmersen took some of it and strew it into glowing coals, it took on first a red-brown color, and then it melted into small gray balls. When treated in a similar way with sodium, small balls of magnetic iron appeared. ... The earths, sitting on the remains of the rhinoceros, must have been laid down by sweet water, which was covering the bodies of the animals, when they had sunken down into the mud." - Brandt, J. F. (1846:223, 224).

The Late Pleistocene woolly rhinoceros (Coelodonta antiquitatis) on the mammoth steppe of Central Europe. After: H.-D. Kahlke (1956) It had a shoulder height of about 1.6 m and weighed 1100 to 1500 kg.

The White Rhinoceros

The woolly rhinoceros is very similar in body build and living habits to the living African white rhinoceros (Dicerus simus). Both of them are mainly grazers. Both of them do have the square upper lip of the grazer. The nutritional needs of the elephant and the white rhinoceros are very similar. The ecological and nutritional requirements of the woolly mammoth and the woolly rhinoceros were also very similar. From this I do conclude that the nutritional needs of the woolly rhinoceros were also very similar to those of the living white rhinoceros.

Where in Africa is the white rhinoceros now living? To what kind of a climate and to what kind of a plant-cover is it adapted? How does it graze? How tall are the grasses, the white rhinoceros is grazing during the course of the year? How much crude protein (dry weight) does this grass contain? And at how much rain per year is this grass growing there?

R. Norman Owen-Smith, University of Witwatersrand, South Africa has studied this animal. He writes: "Favored habitats seem to have been semi-arid savanna, although in Zimbabwe animals are commonly associated with drainage line grasslands. ... The mean annual rainfall at Mpila Camp in Umfolozi (South Africa) is 700 mm (1959-1980), 70% of which falls during the six summer months of October to March. Rainfall increases northwards to reach a mean of 985 mm (1932-1980) at Hilltop Camp in Hluhluve. The vegetation in Umfolozi is dominated by small trees and shrubs of the genus Acacia in the woody layer, and the grass Themedia triandra in the herb layer, although extensive areas of mixed grasses occur." (1988:28, 29).

"Short grasses were the most important food source during the wet season while the grass remained green or mainly green. Shade grasses, in particular Panicum maximum, were sought out especially during the early dry season, when they tended to remain green longer than other grasses. During the dry season rhinos transferred their attention increasingly to medium-tall climax grassland dominated by Themeda triandra. T. triandra was also favored during the wet season when kept short. This species provided the greatest fraction (about 30%) of the food intake on a year-round basis. Forbs made up only 1% of the annual diet, and seemed mostly to be ingested accidentally along with grass. No browsing was observed, apart from occasional instances of chewing on woody stems.

"During the dry season months they concentrated their grazing on short grass grasslands. As the dry season advanced, they shifted their grazing to areas of medium-tall Themeda grassland, though initially seeking out patches of short grass. The fringe of short grass associated with termite mounds was especially favored. By the end of the dry season most of the Themeda grassland on gentle terrain had been grazed down, and animals then moved up onto hillslopes to graze remaining reserves of taller grassland.

"While grazing, white rhinos swung the head in an arc to crop the grass that came within reach with each forward step. The mean height of the grass grazed increased from about 100 mm during the wet season to about 200 mm during the dry season. This was cropped down to a height of 25-60 mm except when tall dry grass was being eaten. On short grass the feeding rate averaged 72 bites per minute.

"The mean crude protein content in whole plant samples of Themeda triandra growing on sandy soils in Umfolozi was 5.9%, compared with 7.8% for plants of the same species collected from bottomland sites with clayey soils. Panicum maximum showed a similar difference (11.7% versus 14.9%).

"White rhinos of the northern species were recorded in Uganda feeding mainly on medium-height grasses. ... The general height of the grass cropped was 250-300 mm, this being reduced to a level of about 50 mm." Owen-Smith, R. N. (1988:41-44).

The African White rhinoceros (Ceratotherium simum), with the square upper lip of the grazer. After: T. Haltenorth and H. Diller (1977) Plate 22. Its horns are round, while those of the woolly rhinoceros were flat, like a knife.

Fat White Rhinoceros

How fat does the white rhinoceros get, when living in tropical and subtropical Africa? Why does it get fat?

R. N. Owen-Smith: "Notably, white rhinos may build up deposits of subcutaneous fat to aid their survival through the dry season. Selous (1899) commented that ‘towards the end of the rainy season, in February and March, white rhinos used to become excessively fat, and would keep in good condition until late in the dry season. I have seen them so fat that between the skin and the flesh over the greater part of the body there was a layer of fat over one inch in thickness, whilst the whole belly was covered in fat to two inches thick.’

"The existence of similar fat deposits in the northern subspecies of white rhino was confirmed by Cave and Allbrook (1958). Hippos were renowned among early hunters for their fat, and Ledger’s (1968) data show carcass fat contents for hippos of 7-11% (= of their body weight), more than twice the mean value for African wild ruminants." - Owen-Smith, R. N. (1988:87).

"White rhinos and hippo transform stands of tussock grass to lawnlike expanses of low-growing or creeping species. Decumbent (= lying down on ground) grasses have a higher leaf to stem ratio, and commonly also higher protein and lower fiber content in their leaves than taller grasses. Thus expanses of short grass offer a higher concentration of leaf material and a high food quality than tall grass stands (McNughton 1985). However, white rhinos are dependent upon reserves of taller grass for subsistence through the dry season when short grass has been grazed down to stubble. If all areas of tall grass were converted to short grass, white rhinos would become more vulnerable to starvation-induced mortality during drought periods." – Owen-Smith, R. N. (1988:259)