Chapter 4: Mammoth Fauna on Arctic Tundra

Could the woolly mammoth and its companions have lived on the arctic tundra, growing now in northern Alaska? Would the woolly mammoth, the bison, the wild horse, the moose, the reindeer, and the other large mammals find up there now enough to eat? What have some of the world’s leading experts found out now about this?

L. C. Bliss is Professor of Botany at the University of Washington, in the northwestern USA. And James H. Richards is at the Department of Range Science, Utah State University, at Logan, Utah. They have investigated the "Present-day arctic vegetation and ecosystems as a predictive tool for the arctic-steppe mammoth-biome". They published it in the book Paleoecology of Beringia (1982).

L. C. Bliss and J. H. Richards state: "The concept of an arctic steppe-mammoth biome with a diversity of small and large herbivores (plant-eaters) and carnivores (flesh-eaters) is difficult to imagine, in light of the modern arctic and northern boreal forest biomes and their rather limited faunas. ... It is not enough, to ask, what kinds of plant communities might have been present; rather, we must ask, what was the net primary production of a diversity of plant communities and what was their potential carrying capacity for a diversity of herbivores and a human population." (1982:241).

"The concept of an arctic steppe-mammoth biome is based upon: (1) a drier and colder climate, but with warmer summers; (2) a tundra vegetation, dominated by species of grass, sedge, and Artemisia, and (3) a diverse herbivore fauna. ... For this modeling effort, the time period from 25,000 to 11,000 BP was chosen on the grounds, that Beringia (= and the Last Ice Age) was then near its maximum extent." Bliss and Richards (1982:252).

"Reconstruction of vegetation in Beringia from lake-sediment cores, colluvial and alluvial cores, exposed sediment sections, and patterns of modern low-arctic plant communities, and floristics of modern low-arctic plant communities leads to the assumption, that four major habitat types occupied regional landscapes in Beringia.

Tall-shrub communities of Salix alaxensis, S. pulchra, and S. arbusculoides occupied floodplains and low river terraces. A rich understory of herbs and grasses in open areas no doubt occurred, as with present shrub communities.

"In poorly drained lowlands, sedge-moss meadows prevailed with a species composition and plant structure, similar to those found in similar habitats today.

"The rolling uplands, that were much better drained, contained a mosaic of upland sedge (not Eriophorum tussock tundra), grass, and Artemisia. Based upon the evidence, that the Pleistocene ungulates had relatively small hoof sizes, compared to modern species, wetlands or soft substrates are presumed minor and winter snows probably not deep nor compacted. Modern moose, caribou and musk ox are adapted, to pawing through snow (about) 50 cm deep, but horses are not. These assumed low rolling uplands, with their tundra-graminoid (grass) vegetation, must have been very important herbivore habitat during summer and winter.

"At higher elevations with well-drained soils, little snow cover, and considerable influence of wind, cushion plants such as Dryas, Saxifraga, Draba, and dry-site sedges, rushes, and lichens predominated. These communities would have been similar to wind-exposed mountain slopes in the Low Arctic and the polar semideserts of the High Arctic today.

"Based upon modern Alaskan landscapes and their dominant plant communities, it is assumed, that in a landscape unit of 1000 kmē, 5% would have been tall shrublands, 10% wet sedgelands, 65% upland sedge, grass and Artemisia, and 20% upland cushion plant-lichen vegetation. This landscape is assumed to occupy a valley system, extending 10 km either side of a river to a ridge crest along each side of the valley and 50 km downstream.

"It is assumed, that this late Pleistocene ecosystem had six dominant large herbivores. Mammoth (Mammuthus primigenius) and horse (Equus) were monogastrics; ruminants included caribou (Rangifer, musk ox (Ovibos), bison (Bison) and moose (Alces)." Bliss, L. C. et al. (1982:252, 253)

Figure 1: Theoretical ecosystem in Beringia 25,000 BP showing the four dominant communities and six dominant herbivore species. The mammoth takes 40% of the food, which it needs, from the tall willow area in the river valley. In the wet sedge area, the mammoth takes also 40% of its needed food. And in the upland- sedge-grass area 65%.

The horse finds 20% of its food in the wet sedge area, and 80% in the upland sedge-grassland. The bison gets 30% of its food in the wet sedge area, and 50% in the upland sedge-grass area. Bliss and Richards (1982:253).

Table 5: Estimated above ground phytomass, net annual plant production (NPP), and percent of NPP, harvested by large herbivores in Beringia in the Late Pleistocene. Total area in this model, 1000 kmē.

Average body weight of mammoth 2230 kg, of horse 150 kg, of bison 450 kg, of caribou 100 kg, of musk ox 180 kg, and of moose 300 kg.

Net primary production: Tall shrub area 400 t kmē yr (400 g DM/mē yr), wet sedge meadow 75 t kmē yr (75 g DM/mē yr), upland grass, sedge, low shrub 100 t kmē yr (100 g DM/mē yr), and upland cushion plant area 25 t kmē yr (25 g DM/mē yr).


Total net primary production in 1000 kmē area:

Tall shrub 20,000 t yr

Wet sedge meadow 7500 t yr

Upland grass, sedge, low shrub 65,000 t yr

Upland cushion plant 5000 t yr – Bliss and Richards (1982:253)

That is 975,000 tons above ground plant production (NPP) dry weight on 1000 kmē per year.

That is 975 gram above ground dry plant matter per square meter per year (975 g DM/mē yr). In other words: During the height of the Last Glaciation, in central Beringia, the arctic tundra is supposed to have produced 975 g DM/mē per year.

Total herbivore biomass would be 800-1450 t on 1000 kmē of Beringian landscape. (1982:255). That is 800-1450 kg/kmē.

Table 7: Total herbivore biomass 799-1445 t kmē (mammoth, horse, bison, caribou, musk ox, moose) on 1000 kmē. (1982:255). That is an average 1122 kg/kmē herbivore biomass, during the Last Glacial Maximum, 25,000 to 11,000 BP, at the peak of the Last Glaciation.

"Herbivore biomass for this Beringian ecosystem (800-1450 kg kmē) is significantly higher than estimates for modern arctic ecosystems (300-400 kg kmē) (Guthrie, 1968a; Hubert 1977), systems that are low in herbivore species diversity." Bliss and Richards (1982:255, 256).

1125 (800-1450) kg kmē : 350 (300-400) kg kmē = 3.214 times larger.



L. C. Bliss and James H. Richards conclude in their ecological model:

·         During the height of the Last Glaciation, while Beringia had reached its largest size, the woolly mammoth, the wild horse, bison, and other hoofed animals have lived in north-eastern Siberia, Alaska and the Yukon.

·         They were living on an arctic plant-cover, on arctic tundra. This arctic tundra in Beringia was growing during the height of the Last Glaciation in a drier and colder climate than now, but with warmer summers.

·         It grew so much fodder, that the woolly mammoth, the horse, the bison and the other ungulates were able, to live there all year round, not just during the short arctic summer.

·         25,000-11,000 years ago, the arctic tundra in NE Siberia, Alaska and the Yukon produced 975,000 tons of dry above ground plant matter on 1000 kmē. That is 975 g DM/mē yr.

·         The total herbivore biomass (total weight of hoofed animals), living then on this 1000 kmē arctic tundra, was then 799-1445 tons kmē, or an average 1122 kg kmē. Today, only 350 (300-400) kg kmē hoofed animal biomass is able, to live now in Central Alaska. That is 3.2 times less.

Professor L. C. Bliss and James H. Richards have made in their ecological model about annual aboveground plant production and hoofed animal biomass in the Alaskan arctic tundra, growing at the height of the Last Glaciation, certain assumptions. And from this, they have drawn then certain conclusions. Are these basic assumptions and conclusions correct? Are they reasonable? Are they scientific? Are they true?

·         Neither the Alaskan arctic tundra, nor any other type of arctic tundra is producing now 975 g DM/mē yr. Nor is it able it support 1122 kg/kmē of ungulate biomass. That is just an unfounded assertion, without any valid observational or experimental proof. This does not agree at all, with how much ungulate biomass the forest-tundra, the arctic tundra, and the polar desert in the Northern Hemisphere actually are able to support now.

·         R. Dale Guthrie (1968a) reports for the area near Fairbanks, central Alaska, a large mammal biomass of 300-400 kg/kmē, on discontinuous permafrost. But that is in the forested zone (boreal forest, taiga, forest tundra), not in the arctic tundra, on continuous permafrost, where it was so cold in summer, that trees were not able to grow. Certainly not during the height of the Last Glaciation! So this proof is not valid.

·         Ben Hubert (1977) estimated a biomass of muskoxen on Devon Island of 300-400 kg/kmē. But that is only on the wet sedge meadows of Truelove Lowland. This is only a tiny part of Devon Island. Truelove Lowland is an oasis, a nutritional hot spot, within a large polar desert.

·         "When considering those areas occupied by muskox, this is 23% of the above-ground forage, available in the preferred habitat on the entire lowland. Summer grazing of the Truelove Lowland is highly sporadic and muskox are virtually absent during periods of melt-off (May-June), (= because the lowland is then flooded) and during the period August to February." White, R.G. (1981:429).

·         "Previous studies with grazing herbivores in temperate grassland systems show, that the availability of forage limits intake of digestible organic matter below a total biomass (dry matter) of 200-220 g mē, or an available green biomass of 0.25 x 200 or 50 g mē.

·         "Predictions of the amount of available green biomass, which would support maintenance of body weights in lactating and non-lactating reindeer, are respectively 25 and 35 g mē, which is in good agreement with field results of 30 to 40 g mē for sheep and 30 to 37 g mē for lambs. Hence, although the total aboveground biomass of vascular plants may frequently be in excess of 100 g mē, for most for the year the amount of green or preferred material may be less than 30% of this figure and a decline in body weight of animals would be expected. At biomass levels below 20 g mē, feeding time must be maximised." White, R. G. (1981:439-441).

·         If the muskox is able to live there, this does not mean yet, that also the elephant, bison, and other large mammals are able to live on the arctic tundra and polar desert.

·         Not only the amount, also the quality of the food is important. The elephant in Africa will starve to death with a full stomach in a dry grassland, where the grass shoots are 2-3 m high. Because this dry, brown grass contains too little protein and too many fibers. The elephant cannot eat it, digest it. It is not food for the elephant anymore.

·         The plant-cover on Seward Peninsula, westernmost Alaska, in the middle of former Beringia, was preserved beneath a thick layer of volcanic ash, like the City of Pompeii beneath the volcanic ash of the Vesuv. According to the carbon-14 dates, this plant-cover has grown at the height of the Last Glaciation.

·         It completely disproves the ecological model of the mammoth fauna on the Alaskan arctic tundra. The summer was not longer and warmer, but shorter and cooler than now. Its plant-cover resembles more that of a very dry northern arctic tundra, changing over into semi-polar desert, as we find it now on the southern parts of the Canadian High Arctic Islands. Neither the mammoth, nor the bison, nor the horse, nor the moose could have lived there. They would have pitifully starved there to death, due to lack of energy and digestible crude protein. Their model is based on unfounded speculation, not serious scientific fact.


Aboveground annual plant production (dry weight)

How much aboveground vegetation do the different zonal plant-covers of the northern part of the Northern Hemisphere now produce? How much of this vegetation are the animals able to eat? And how much hoofed mammal biomass do these different zonal plant-zones now support?

We shall find out more about this in, "Grazing in Tundra and Northern Boreal Environments" by R. J. Hudson and F. L. Bunnel in: Grazing Animals, F.H.W. Morley (Editor) (1980):

Climate. "The high albedo [reflection] of snow and water reduces absorption and keeps net radiation income small, about 15 kly (15 kcal/cmē) y on the tundra, 15-20 kly (15-20 kcal/cmē) y at the forest-tundra ecotone (transition zone) and 20-35 kly (20-35 kcal/cmē) y in the boreal forest. ... The tundra experiences only about 125 snow-free days annually, while the boreal forest experiences about 185." Hudson, R. J. and F. L. Bunnel (1980:205)

Pleistocene fauna. "Tundra and boreal biomes supported a much more diverse and abundant fauna in the Pleistocene, during times, when environments may have been less harsh and vegetation more productive. ... Reasons for the sudden impoverishment of this megafauna are disputed." (1980:206). Emphasized by me.


Typical values for standing crop,

net annual primary productivity and sustained forage production


Standing crop (g/mē)

Net annual primary productivity (g/mē y)

Sustained forage production (g/mē y)

Polar desert




Tundra (proper)




Forest tundra




Open boreal woodland




Boreal forest (proper)




Boreal mixed wood




After Hudson, R. J. and F. L. Bunnel (1980:209) Table 11.1.

"The standing crop of the forest-zone includes also the wood of the trees, which the animals are not able to eat. Sustained forage production includes non-woody vascular plants, lichens and current growth of browse. Assumes 50% of non-woody vascular plants, 10% of lichens and current annual growth of shrubs can be grazed each year. No correction is made for availability under snow cover." R. J. Hudson and F. L. Bunnel (1980:209)


Secondary productivity of northern grazing systems


Animal species

Biomass (kg/kmē y)

Productivity (kg/kmē y)

Polar desert

Muskoxen, Peary caribou




Caribou, muskoxen



Forest tundra




Open boreal woodland

Caribou, moose



Boreal forest

Moose, reed deer, reindeer, musk deer



Boreal mixed wood (Europe)

Moose, red deer, roe deer, wild boar



Boreal mixed wood (North America)

Bison, wapiti, moose



After Hudson, R. J. and F. L. Bunnel (1980:210) Figure 11.2. Changed from kg/ha to kg/kmē.

"In general, biomass and productivity of large herbivores parallel latitude and gradients of primary productivity. Polar deserts support ungulate biomasses of only 0.1 kg ha (0.0-10 kg/kmē) (Kevan, 1974). Tundra ranges, on average, support about 0.18 kg ha (18 kg/kmē) (Klein, 1970). Biomass increases in the boreal forest to a maximum of over 50 kg ha under optimal conditions at the boreal forest/prairie transition (Telfer and Scotter, 1975).

"These latter values compare favourably with maximal values of several hundred kg ha observed in the most rich African savannahs. However, the high densities, observed in Elk Island National Park, may be influenced by fences, which prevent animal dispersal and therefore normal regulation of populations. More typical values, represented in European forest, are 4-10 kg ha (Grodzinski, 1975, citing others)."

"In comparing biomass densities of individual species, it is important, to distinguish those, which attain high local densities, but which are not widely distributed, from those, which are more widely dispersed and exploit a wider range of habitats. Bison, muskoxen and wild sheep are examples of the former, while moose and reindeer are examples of the latter. (1980:211).

"In the high arctic, muskoxen often achieve higher local biomass densities, than do caribou, and in the boreal forest, bison usually attain a higher biomass, than the smaller species, with which they are associated. A similar pattern has been observed in grazing systems in southern latitudes.

"In polar deserts, muskoxen are most abundant in wet sedge meadows, whereas Peary caribou are common only in uplands, supporting plant communities, dominated by Dryas, sedges, grasses and lichens." Hudson, R. J. and F. L. Bunnel (1980:211)



Number of species

Census Area (kmē)

Biomass Density kg/kmē







Yukon, Canada




Geist (1978)

N.W.T., Canada




Parker (1975)

After: Redmann, R.E. in, Paleoecology of Beringia (1982:230) Table 230. Kg ha changed into kg/kmē. These ungulate biomass values are mostly from the forest tundra, not from the arctic tundra.