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Home , Forest steppe, Mammoth steppe, Palaeoloxodon, Steppe mammoth

P. V. Putshkov Institute of Zoology,Academia of Sciences, Kiev

The impact of on their : clash of two paradigms

Putshkov, P.V., 2003 - The impact of mammoths on their biome: clash of two paradigms - in: Reumer, J.W.F., de Vos, J. & Mol, D. (eds.) - ADVANCES IN RESEARCH (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 365- 379 [ISSN 0923-9308] Published 24 May 2003

Mammoths, woolly rhinoceroses, musk oxen, primitive , and horses of northern races were stenobiotic cryoxerophiles living only under extreme cryoarid conditions according to the ‘- crash paradigm’. These are considered to be strict grazers on arid steppe-tundra gras- ses; hence, the possibility that mammoths maintain their pastures the way modern do is rejected. Climatists claim that mammoths and their faunal satellites were killed by the warming a result of their inability to feed themselves in any of modern landscapes and to withstand the weather-caused losses. Their survival throughout interglacials is considered as resulting from climatic instability: short warm episodes are said to have alternated with very cold ones. It is belie- ved that a permanent Arctic Ocean ice-shield persisted along the Siberian coast even in summer; its cooling and aridifying influence maintained steppe- and similar throughout Northern . On the contrary, the stable Holocene warming resulted in the disappearance of this ice-shield, in the destruction of the of the mammoths, and in megafaunal extinctions. This conception is based mainly on the properties of Eemian Greenland ice layers. However, these layers do not indicate the real Eemian climate. Non-altered Antarctic ice cores and Atlantic deep sea cores show that the Eemian optimum climate was as stable as the Holocene one. The idea of the persistence of the Arctic ice-shield throughout interglacials also contradicts with this evidence. The real reason why and rhino persisted throughout interglacials (as well as through- out glacials) is in their tolerance to a vast range of climates and their ability to maintain highly pro- ductive pasture ecosystems. Contrary to the opinion of climatists, these pachyderms were polypha- ges that ate various herbaceous and woody plants. They held back and tundra by various direct and indirect influences. Whatever the climate was, pachyderms prevented the appe- arance of closed and they ensured the predominance of grasses and herbs over mosses and shrublets, as well as high mosaicism of the vegetation. The pachyderms trampled on snow and broke the frozen snow crusts, thus facilitating the smaller ungulates to overwinter. Similarly, Paleoloxodon and Dicerorhinus created vast meadows throughout the temperate forest zone. Giant deer, bisons, horses, etcetera, used these meadows. The ecosystem impact of the climate-resistant pachyderms caused remarkable stability of pasture ecosystems throughout the . In addi- tion, the largest carnivores were important prehistoric stabilising agents, due to their pressure on populations of humans, and . The Pleistocene crisis in the Palearctic included the same main processes as in other continental realms: (1) liberation of mankind from carnivore con- trol, (2) human-induced removal of pachyderms, (3) drastic environmental changes harmful to as a result of this removal, (4) secondary extinctions of herbivores as a result of envi- ronmental effects of pachyderms, (5) impoverishment of the large predator guild due to the great prey shortage aggravated by the other reasons, (6) other secondary extinctions, (7) installation of a new equilibrium. Due to the long coevolution of Palearctic megafauna with man, the extinction pro- cess in Palearctic was long-lasting and less catastrophic when compared to the Nearctic situation.

Correspondence: P.V. Putshkov, Institute of Zoology, Academy of Sciences of Ukraine, 15, B. Khmelnitsky street, 252 601, Kiev-30, Ukraine, e-mail: [email protected] or ivanoff@pal- muz.freenet.viaduk.net

Key words: woolly mammoth, , ecological impact, mammoth biome, Pleistocene ecosystems, biotic interactions

365 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

INTRODUCTION giants. As applied to mammoth ecosystems The key point of the Pleistocene extinction the main cause of the ‘mammoth pastures’ debate is (are) the factor(s) that favoured the maintenance were the mammoths themselves. worldwide thriving of diversified sets of large In what follows we define the most important quadrupeds. If this factor was of a climatic divergences of both viewpoints without nature, the climatic explanation of mass giving the detailed argumentation. For such extinctions without replacement of the terres- argumentation in favour of the ‘steppe-tundra trial megafauna would gain credibility. If it crash paradigm’ see Guthrie (1990a, b) and was of a biotic nature, the most probable Sher (1995, 1997a, b); for that in favour of explanation of these extinctions would be the panbiotic explanation see Putshkov (1989a, b; man-caused upset of the Pleistocene ecologi- 1997). cal equilibrium. Some aspects of this problem as applied to mammoth ecosystems are consi- WOOLLY MAMMOTHS AND RHINOS: dered below. The following terms are used: EURY- OR STENOBIOTIC FORMS? Climatists consider these animals as stenobio- ‘kyBP’ - ‘thousands of years before present’ tic cryoxerophiles living only under steppe- ‘large herbivores’ - herbivores weighing more than 50 kg tundra extreme cryoarid conditions. It is sta- ‘giants’ or ‘pachyderms’ - those herbivores weighing ted that the giants could not resist the outcome more than 1000 kg of the Holocene warming, being unable to ‘eurybiotic’ (= euryoecious) - tolerant to a wide feed themselves in the established forests, range of environmental conditions swamps and tundras as well as to withstand ‘stenobiotic’ - species tolerant to a narrow range of the losses caused by deep or/and ice-crusted environmentel conditions snow, by the soaking of their wool, etc. Even ‘climatist’ - proponent of climatic models of Pleistocene modern and forest-steppes are an extinctions inappropriate environment: the summer tem- ‘biotists’ - proponents of biotic models of Pleistocene peratures are too high and the vegetation too extinctions monotonous for efficient feeding ‘Eem’ and ‘Riss/Würm’ are used as synonyms. (Vereshchagin 1979, 1988; Sher 1971, 1995; Velichko 1973; Guthrie 1990a, b; Lister & The dominant climatic concept as applied to Bahn 1994). mammoth ecosystems is the well known However, a comprehensive analysis of the ‘steppe-tundra crash paradigm’. It asserts that available data (Putshkov 1989a, 1991, 1997) woolly mammoth and rhinos existed only shows that both giants were highly polypha- under steppe-tundra extreme cryoarid condi- gous and eurybiotic creatures. Both were well tions and were killed by the outcomes of the adapted for locomotion over swampy, snowy, Holocene warming. The main non-climatic ice-covered and other surfaces as well as for explanation of the Pleistocene extinctions the destruction of the frozen snow crusts was the ‘overkill’ paradigm for a long time during the feeding and locomotion. Size and (Martin 1984). By now it is gradually being strength were potent advantages during all replaced by the ‘key herbivores removal’ seasons. Both pachyderms inhabited a wide (Owen-Smith 1987, 1988; Schule 1990; range of climates (from warm-temperate and Maslov & Antipina 1994; Zimov et al. 1995) mesic to extremely cold and dry) and lands- and panbiotic (Putshkov 1988, 1989a, 1989b, capes (from completely treeless to quite well 1997) models. According to these, only forested; with and without ; with pachyderms were overhunted among the and without considerable swamps). The mainland animals. The other species have giants were invariably accompanied by hor- perished chiefly due to the ecosystem chan- ses, bisons and lions whereas species indica- ges caused by cessation of activity of the ting now exclusively tundra and northern

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(, arctic fox, lemmings) or dry case of the more general phenomenon that steppe (saiga, etc.) conditions are present in has already been noted by Darwin and called some but absent in other mammoth sites. the ‘paradox of prehistoric pastures’ There are also many sites where both pachy- (Putshkov 1989a, 1992a). The cited features derms are accompanied mainly by animals of were inherent not only to mammoth ecosys- temperate forests and forest-steppes. tems, but to other dominant Pleistocene main- land ecosystems too (Schule 1992, May REGIONAL PARADOX OR WORLD- 1993). The main prehistoric plant communi- WIDE GIANTS PASTURES? ties were often similar not to modern zonal It is impossible to put the variety of mam- formations, but to the places of their contact moth environments to the Procrustean bed of (ecotones) or to non-climax communities of the ‘crioxerotic treeless steppe-tundra’ sensu diverse stages of the plant succession. Due to Velichko (1973) or Sher (1971). Refusing to this there were an increased local biotic admit this fact openly, climatists did recogni- diversity, including the unusually (in compa- ze it silently by proposing an excessively rison with most Holocene ecosystems) high broad understanding of ‘steppe-tundra’. Now large density and diversity. Even they include into ‘steppe-tundra’ (Sher 1995, more, the ecosystems that were more favou- 1997a) or ‘’ (Guthrie 1990a, rable for large mammals than the Holocene 1990b) even the mesic parklands of scattered ones had been globally distributed during northern taiga trees (, , , etc.) most of Cenozoic era. with rich herbaceous cover of meadow plants. It has been shown that quasi-global pre- However, even such interpretation could not Holocene distribution of fertile pasture eco- be applied to certain well-forested and forest- systems could not be explained by mere cli- ecosystems that indicated matic reasons; it was a natural and inevitable warm-temperate and/or mesic climate condi- consequence of the feeding of giant herbivo- tions (see Putshkov 1989a, 1997 for a res and of other activity as shown by ecosys- review). tem impacts of modern pachyderms (see On the other hand, all mammoth ecosys- Putshkov 1989a, 1992a, 1997 for a review). tems indeed possessed qualities very impor- tant for the large herbivores (Yurtsev 1976; CRYOARID CLIMATE KEY REASON Sher 1982; Vereshchagin & Baryshnikov FOR MAMMOTHS’ WELLBEING? 1983; Musil 1985; Verkhovskaya 1988; The dominant taiga and tundra vegetation Guthrie 1990a, b): (1) highly mosaic vegeta- (mosses, lichens, shrublets, ) is unfa- tion cover, (2) the prevalence or, at least, vourable for the large grazers feeding. Plants abundance of plants and plant associations grow slowly, their nutritive value is low; precious for large grazers feeding, (3) in case spruce and many other plants are even toxic of tree presence, closed forests never appe- (Guthrie 1990a, 1990b). Such vegetation is ared: open , parklands or cryophy- not merely a climatic product: dominant tic with rich herbaceous cover grew plants maintain their communities themsel- instead, (4) the rich megafaunal set of mam- ves. So, moss and/or fallen conifer needles moth ecosystems strikingly resembled such carpets resist to evaporation thus cooling and sets of African savannas in spite of certain excessively moistening the soils. Hence, mos- differences. These features are inconsistent ses and shrublets obtain advantages over the with low productivity and certain other meadow grasses even in the scattered larch modern taiga and tundra vegetation particula- stands of NE (Sher 1997a: 25), rities. Hence the catchword ‘steppe-tundra though lime-rich fallen larch needles improve paradox’ has appeared (Vereshchagin 1988). the soil quality for meadow plants However, this paradox is only a particular (Shennikov 1964). In the spruce forests the

367 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

meadow vegetation could not grow, not only Siberian area seems to be well founded. This because it needs drier and richer soils but view corresponds with the direct data on also due to overshadowing and increased soil these animals feeding too (see below). acidity (Guthrie 1990a, 1990b; Rabotnov Second, mammoths with satellites became 1996). Stands of other full-grown boreal and extinct even there where deep snow, wet win- temperate trees also oppress herbs and gras- ters and a continuous moss carpet over ses to a various extent. Due to all this, spruce swampy soils have never appeared due to the forest and -to a lesser extent- other northern constant domination of the dry and markedly plant communities ‘exclude’ many cold-adap- continental climate throughout the ted herbivores from ecosystems. The density Quaternary. Such is the case of Trans- and ranges of cervids, bisons, bighorn sheep baikalian and Mongolian steppes and forest- largely depend on the resources of the restric- steppes as well as one of the Central Yakutian ted azonal such as riverside or alpine scattered larch stands with rich mesic meadows, willow thickets, glades, post-fire meadow cover over dry soil. Horses and sites, steppe-like vegetation of the southern cattle feed here in the wild all year round; in slopes, etc. For ‘strict grazers’ such as horses, winter they dig for food through thin crumbly musk-oxen, woolly mammoths and rhinos snow. Even these ecosystems are less there is no place at all (Guthrie 1990a, favourable for large grazers than mammoth 1990b). Thus, climatists argue that only meadows of Kazantsevo (Eem) and Kargin climate driven aridization could convert taiga (Mid-Würm warming) optimum times were and tundra to fertile mammoth step- (Verkhovskaya 1988). It shows once again pes (Sher 1995, 1997a). However, such a sta- that climate driven aridisation was not the tement contradicts other, no less important key factor of such phenomenon: during these evidence. warmings the East Siberian climate was war- First, climatists often consider the so-called mer and wetter than it is now (Putshkov steppoids, the azonal cryoarid steppe-like 1989b, 1997). plant associations of North-Eastern Siberia Third, the experimental enriching of north- and Alaska, as relics of the notorious steppe- ern soils with fertilizers, as well as grazing tundras. However, due to soil infertility, by horses and cattle, lead to the ousting of warmth and humidity deficiency, mighty mosses, sedges and shrublets by meadow winds and insufficient snow cover in winter grasses and forbs (Guthrie 1990a: 200-205; the steppoid plants are low, scattered and Zimov et al. 1995). This allows us to think slow growing. Their productivity, at least that intensive grazing by mammoths and their under Chukotka conditions, is ‘insufficient satellites could drive taiga and tundra com- even for a hare feeding’ (Kozhevnikov 1977: munities to a zonal condition even 445). It is meso- and hygrophilous meadow without any general climate-caused increase vegetation that has the high productivity and in aridity (Putshkov 1989a, b; 1993b; 1997; nutritive value for herbivores in Arctic and Zimov et al. 1995). Climatists neglect or dis- Subarctic (Yurtsev 1976; Kozhevnikov 1977, card such a possibility. How well justified is 1986; Verkhovskaya 1988). Such vegetation their position? and not the xerophilous one has dominated in NE Siberia during global warmings. The role NORTHERN GIANTS,ARID GRASSES of meso- and hygrophilous plants remained AND NORTHERN TREES essential even during coolings that Guthrie (1990b: 50, 51) claims that there caused expansion of steppoids. Therefore the could be no valid analogy between the ‘unu- opinion that mesic and hygrophilous plants sual ecological effect’ of mixed grazers/ were the staple fodder of mammoths -and browsers such as African elephants, and that their satellites- in the most of the Euro- of ‘strict grazers’ such as woolly mammoths

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and rhinos. The woolly giants ‘did not rely on ‘eternalized’ or early successional browse for the dry winter season’, never stages of the forest restitution in various cli- ‘...made significant use of woody plants, matic zones (see Putshkov 1989a, 1997 for especially the birch, , spruce, larch and review). that were taking over the northern step- Thus, the grazing adaptations of mam- pes as mammoths became extinct ...’. As their moths, woolly rhinos, horses, hemiones, predominant diet were ‘arid grasses’ the bisons, yaks, musk-oxen, bighorn sheep, sai- giants ‘... could not have held back the mesic gas and reindeer could not be considered as forests’ either by direct or by indirect effects. signs of their harmlessness to woody vegeta- Eating exclusively grasses they could only tion. In line with this are direct data on mam- accelerate the demise of their grassland habi- moths and their satellites’ food. Contrary to tats, when climate became favourable for Guthrie’s claim, grasses and sedges from fro- woods or moist tundra. Such statements are zen Siberian mammoths, horse and car- largely based on clear-cut grazing adaptations casses are meso- and hygrophilous and not of mammoths and their satellites, such as ‘arid’ ones (Ukraintseva 1985, 1993). Next to hypsodont teeth with a complicate masticato- these, brackens, mosses and forbs were usual ry surface, the bifid tip of the mammoth trunk and, at times, important food ingredients. allowing to pluck even very low plants, cra- These data correlate well with the fact that it nial and upper lip morphology of woolly is meso- and hygrophilous meadow vegeta- rhino, wide muzzles of horses, musk-oxen tion and not a xerophilous one that has and and bisons. The other proof is the predomina- had a high productivity and nutritive value tion of grasses and sedges in the digestive for herbivores in the Arctic and Subarctic tract of frozen carcasses of the mentioned (see above). The xerophilous steppe vegeta- animals (Formozov 1990; Guthrie 1990a, tion should be the staple of mammoths and 1990b). However, these assertions are rather their satellites’ diet only in more southern misleading and biased. regions (Putshkov 1997). First, observations on the modern ungulates Woody plants are also invariably present in and the isotopic evidence from dental sub- the frozen mammoths´ carcasses: the macro- stances of extinct herbivores have demonstra- rests of currant, chosenia, willow, larch, tree ted that no grazing specialisation makes the birch and alder were found (Ukraintseva browsing impossible. Hypsodont teeth do 1985, 1993). Hence, Guthrie’s statement that enable ungulates to feed efficiently on gras- mammoths did not eat the three latter species ses but they by no means prevent the use of is incorrect. The feeding on pine, spruce and the less abrasive food - bark, young branches poplar is not proven though not excluded and leaves (Schule 1990, 1992; Haynes 1991; (ibid.). Though the browsing fraction is low Putshkov 1997). The hypsodont northern gra- as opposed to the grazing fraction it should zers such as bisons, wild sheep and goats, be stressed that we do not know the winter chamois and ghorals switch in winter to food of mammoths, when browsing could browsing (Formozov 1990; Heptner et al. play a far more important role: carcasses 1961). Both modern elephants and many tro- from permafrost belong to animals, that pical herbivores also have a mixed diet or a perished during warm seasons (ibid.). diet switching seasonally from grazing to Opinion that mammoths always ate the non- browsing (Kingdon 1979, 1982; Penny 1987; graminoid plants only in small quantities for Owen-Smith 1988). It should be particularly the sake of the metabolism correction emphasised that specialised grazers destroy (Guthrie 1990a, 1990b) contradicts to the iso- tree seedlings and saplings far more efficient- topic evidence in the hydroxyapatite from ly than brachyodont browsers. For this reason tusks of Great Lakes mammoths that inhabi- they often impeded the reforestation and ted the spruce-dominated woodlands. These

369 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

data show ‘a much greater proportion of the course of succession, birch or aspen browse in their diet than had previously been woods with juvenile spruce or pine in the thought’ (Haynes 1991: 88). In line with this undergrowth precede the climax stages of is the pattern of the variability of Old World mature spruce or cedar pine forest. The rapid- woolly mammoths’ dentition. The so-called ly growing or aspens protect young boreal mammoths had teeth adapted to forage conifers from excessive sunlight and belated with greater amount of forbs and woody frosts (Rabotnov 1996). Such protection is plants, the periglacial ones possessed teeth necessary for young conifers during many more specialised to coarser grasses. The for- years: spruce and cedar pine needles that mer (boreal) type always occupied most of resist to winter frosts of - 40oC or more, are ; during warming it replaced the perig- damaged by night frosts of - 7oC in warm lacial type more or less completely in seasons (Walter 1982). The repeated damage European , and in Siberia too, as the of the broadleaved protective cover by mam- role of arboreal vegetation in ecosystems moths and their satellites should inevitably increased (Foronova & Zudin 1986). A simi- lead to the destruction of many spruce and lar phenomenon has been observed in the pine seedlings and saplings (Putshkov 1997). woolly rhinos skull structure that changed However, even dense birch or aspen grove from ‘strictly grazing’ to ‘grazing/browsing’ stage often could not be attained due to mam- type in regions and/or epochs with more or moth’ -with satellites- pasture activities. The less well developed woody vegetation (N.E. repeated action of hooves of various sizes Garutt, pers. comm. 1995). It means that and forms destroyed the moss cover and pre- strictly grazing white rhino is not so good vented its reappearance. It caused the deep model of woolly rhino adaptability. soil thawing and drying in summer. Eating up Even if woolly mammoths and rhinos enormous quantity of plant biomass, animals indeed recurred to feeding on large trees abundantly fertilized the soil with the dung relatively rarely, one must remember that and urine; dung beetles reinforced the fertili- pachyderms destroy young and full-grown zing effect. Under such conditions grasses trees not only by feeding but also as a result and herbs, rapidly growing on fertile soils of other aspects of their behaviour. and resisting herbivores more efficiently by Mammoths, much as the modern elephants rapid compensatory growth than by deterrent (Kingdon 1979; Owen-Smith 1988), would substances (Guthrie 1990a, b), had an advan- destroy many trees during their sexual beha- tage. They ousted slowly growing plants, viour, simply out of pleasure, to clear paths, including spruce seedlings that could realise for the sake of communication, etc. (ibid., their odds (protection with toxic compounds) Putshkov 1989a, 1997). Rhinos, wisents and only on poor, unfertilised soils. Due to all stags also break young trees simply for the this steppe-like plant communities established sake of physical exercise (ibid.). under drier conditions, and the meadow-like It is plausible that spruce has hardly been communities under wetter climatic conditions. eaten by extinct large herbivores as it is Thus, mammoths prevented the closed unwillingly eaten by the living ones (Heptner forests appearance even in the regions clima- et al. 1961; Walter 1982). But, on the other tically fit for the dark-needled taiga. The fer- hand, the mammoth fauna did coexist for tile meadows maintenance in the present hundreds of thousands of years with spruce larch taiga zone was by far an easier task for and pine woods, though not so closed as them. The Pleistocene predomination here of Holocene forests (Vangengeim 1977; Musil birch over larch (Sher 1995, 1997a) was the 1985; Stuart 1991). Most likely mammoths consequence of intense pasture activity too. prevented the appearance of closed spruce Birch is excluded now from vast monodomi- forest without heavy browsing on . In nant larch taiga regions due to the thick con-

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tinuous moss carpet presence causing the pieces by hooves or eaten up in spite of its high position of upper limit of permafrost and low digestibility (food masses from some fro- water logging of the thawed soil layer. The zen mammoths are rich in mosses; moss carpet elimination and other large gra- Ukraintseva 1993). Thus, repeated pasture zers effects (see above) caused the lowering exploitation ‘eternalised’ the competitive of the permafrost upper limit as well as soil advantages of meadow grasses and forbs over warming, drying and fertilizing effects pro- mosses and shrublets. The Pleistocene Arctic fitable to grasses, forbs and birches. The lat- and Subarctic meadows were at the same ter preceded in the succession. Rapid time sustained by the large herbivores herds growth gave to birch an additional advantage: and nourished them (Putshkov 1989a, 1989b, birches coped easier than larches with the 1997; Zimov et al. 1995). Due to impacts of harm caused by mammoths’ cold season the herds on soils and vegetation, the inverte- browsing. brate communities differed from those of modern taiga and tundra. Among insects, the MAMMOTHS AND TUNDRA ZONE role of species connected with steppe and VEGETATION meadow vegetation remained considerable, Acatchword ‘tundra is taiga lacking trees’ even during the warm mesic phases (Kiselev (Chernov 1980: 7) means that presence of a 1980; Sher 1995). Among soil oribatid mites, thick moss carpet is equally characteristic for there were even species now extinct both biomes. This carpet’s effects in tundra (Krivolutsky & Druk 1982). The impact of are as baneful for meadow grasses and herbs large herbivores has favoured the northward as they are in taiga. It is noteworthy that in expansion of cold resistant trees that obtained tundras small meadow patches appear not possibilities for germination, sprouting and only due to local abiotic conditions, but also subsequent growth over moss carpet-depri- when soil receives fertilisers such as garbage ved, well dried, deeply thawed and fertilized and excrements. These patches occur on the soils. Consequently, birch and other trees monotonous moss-covered flats in places penetrated during Kazantsevo (Eem), Kargin with high lemming density, around Arctic (Mid-Würm) and Early Holocene warmings foxes burrows, on tops of hillocks serving as far more northwards than now. dining places for birds of prey and near the human settlements. Dense, highly productive ZOOGENIC VEGETATION MOSAICISM and rapidly growing grasses and forbs deve- AND STEPPOIDS PRODUCTIVITY lop there, oppressing mosses and lichens up Mammoths were the principal forces that to the moment when soil looses its fertility created and perpetuated the zoogenic (ibid.). meadows. Thanks to them horses, bisons, In the Pleistocene such meadows inevitably musk oxen, other ungulates had high popula- became the dominant vegetation in all areas, tion densities and could freely migrate climatically fit for present-day typical and through and brushy spaces by shrub tundras, since everywhere soils got meadow or cryophitic ‘’ corridors enough fertilisers in the form of urine, excre- (Putshkov 1989b, 1993b, 1997). These herbi- ments and large animals carcasses (Putshkov vores in their turn considerably reinforced 1989a, 1989b, 1997; Zimov et al. 1995). The and modified the ecosystem impact of pachy- reappearance of a moss carpet was hindered derms, increasing the mosaicism of plant because grass and forbs cover did not prevent communities as do now various herbivores in the deep soil thawing during the warm season elephants dominated ecosystems (ibid., and has dried up the soil considerably by the Kingdon 1979, 1982). Not only overgrazing, high evaporation rates. If a moss carpet but also undergrazing leads to degradation of nonetheless reappeared, it were soon torn to meadows in forest and tundra zone regions

371 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

(Shennikov 1964, Zimov et al. 1995). Seeds the herds. A patchwork of feeding places con- of many meadow and steppe plants are distri- nected by wider and narrower paths trampled buted by large animals (ibid., Janzen 1984). by animals appeared. This patchwork was Ungulates are the best agents for the sprea- renewed after snowfalls and took a new pat- ding of seeds of herbs and grasses of distur- tern when the foraging places changed. bed places. Plants, covered with excrements, Feeding and moving mammoths, rhinos, large mostly die, giving place to young plants, herds of bisons and horses broke the frozen rapidly growing from the seeds that have snow crust over large areas. Other herbivores been deposited with the dung and then buried easily grazed over the fields where the snow- by dung-beetles. Moving over considerable crust was smashed by the mammoths. They distances, large herbivores ‘sowed’ the plants also browsed upon branches and trees broken far from the places where the seeds were by the pachyderms. Animals used mammoth- eaten. Thus they locally restore plants disap- made paths and roads much in the same man- pearing for various reasons. Different herbi- ner as nowadays Far Eastern deer, boars and vores were efficient as feeders on different tigers use the roads trodden by timber-car- woody and herbaceous plants, and more or rying lorries (ibid.). Thus, they spared forces less efficient propagators of their seeds. evading snow crust and deep snow. Thanks to These phenomena have largely contributed to the spared forces a cave lion caught prey in the maintenance of the locally diverse vegeta- winter as easy as in summer while ungulates tion, mosaicism and productivity (ibid., efficiently raked the snow in places untou- Putshkov 1989a). ched by mammoths, when the touched areas The opinion that cryoxerotic vegetation became unusable as a result of overgrazing or was the principal food source for North-East to secondary condensation of snow by winds, Siberian mammoth does not correspond with sun or animals. In summary: many other the evidence. However, it is probable that the large mammals had better chances to over- opposite view (that completely negates the winter thanks to the mammoths (ibid.). alimentary significance of the cryoxerotic vegetation for the North-East Siberian MAMMOTHS AND THE PLEISTOCENE Pleistocene ungulates) is biased too: the PASTURE ECOSYSTEMS STABILITY Pleistocene Arctic and Subarctic steppoids So, woolly mammoths (or for the earlier might be more fertile than Holocene ones due times their ancestors) ‘eternalised’ the highly to the activity of mammoths and their satelli- productive and mosaic pasture ecosystems. tes (see Putshkov 1997 for the details). The high tolerance mammoths have to envi- ronmental changes assured the stability of MAMMOTH IMPACT ON SNOW such ecosystems in spite of frequent climatic COVER oscillations. Coolings and warmings, aridifi- Mammoths (etc.) increased not only the cations and humidifications caused shifts in mosaicism of the vegetation but the mosai- the composition of plant species. Yet, the cism of the snow cover as well (Putshkov eurybiotic giants, together with other large 1989a, 1997). Partly the latter was a conse- herbivores, always provided advantages for quence of the former: snow was deeper and the rapidly growing grasses and forbs over ‘crumblier’ in forests, while snow was more other plants, thus maintaining highly produc- condensed and shallower in open areas. tive pastures (Putshkov 1989a, 1992a, 1997). Macromosaic animals caused subsequent While the pre-Holocene major climatic war- variations of this snow cover. Mammoths mings favoured closed forests, moss/shrub tusks, rhino horns, and hooves of various tundras existed approximately in the same ungulates hooves raked and shoveled the regions as today. The giants prevented such snow over large areas during the feeding of biomes from installing. They maintained

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open woodlands or parklands with rich impacts superposition. In regions with clima- meadow or steppe-like herbaceous vegetation tically driven thick snow cover and humid instead of closed forests, and meadows or soils, mammoth's activities created satisfacto- meadow dominated parklands instead of tun- ry conditions only for herbivores of conside- dras. During the coolings zoogenic and cli- rable mild soil and snow cover tolerance. matic agents of deforestation often acted in Hence, the interglacial range of saiga and, to concert, creating steppe-tundras sensu Sher a lesser extent, that of musk-ox, dwindled (as (1982). However, these treeless formations compared to glacial range) to a greater extent have never covered Northern Eurasia comple- than did the ranges of mammoths, woolly rhi- tely, because there were regions wet enough nos, horses and bisons. But in regions of cli- for closed forests to persist in certain parts of matically driven thin snow cover, so wide- Europe and Eastern Asia. Here mammoths spread even now in Eastern Siberia, snow- and/or straight-tusked elephants constantly scraping and frozen snow crust smashing maintained open woodlands with meadow activities of mammoths created satisfactory patches (Putshkov 1989a). Predators modified conditions even for saigas and hemiones effects of herbivores on vegetation and their (Putshkov 1997). interactions between each other. The largest carnivores pressure on human and popu- THE HOLOCENE:A NORMAL OR AN lations was not a less important element of ABNORMAL INTERGLACIAL? the ‘prehistoric equilibrium’ than their control Climatists claim that the Holocene climate is over populations (Putshkov 1988; sharply different from the interglacial clima- 1989a,b; 1992a; 1993a,b; 1997). The combi- tes, though they disagree on the kind of the ned action of various megafaunal species difference. Ideas that the Holocene climate is explains the notorious mosaicism and the sta- less equable, more severe, or more mild than bility of ‘mammoth ecosystems’ and other the interglacial climates, as well as the opi- Pleistocene ecosystems much more convin- nion that the Weichselian/Holocene transition cingly (ibid.) than an appeal (Guthrie occurred faster than did previous climatic 1990a,b) to climatic causes. shifts have failed to be tested by the evidence (see Putshkov 1989b, 1997 for a review). CO-OCCURRENCE OF MAMMOTH, Instead, an interglacial ‘flickering’ climate SAIGA AND REINDEER idea has been proposed (Sher 1995, 1997a, Climatists pay particular attention to the co- 1997b; Lister & Sher 1995). It states that occurrence of saiga and reindeer in numerous during interglacials short warm episodes ‘mammoth sites’. For them it means that: (1) alternated with very cold ones. Due to this Pleistocene climate has no modern analogue, the continuous Arctic Ocean ice-shield persis- for the mentioned ungulates nowhere occur ted even in summers. Its cooling and drying together now; (2) like saiga, woolly mam- influence has maintained steppe-tundras in moth and woolly rhino were did not tolerate North-East Siberia and similar pasture eco- woodland, boggy ground, deep snow and fro- systems westwards and southwards. Holocene zen snow crust conditions (Formozov 1990; warming, unlike the previous ones, was sta- Sher 1971; Velichko 1973; Guthrie 1990b). ble. As a result, it caused the disappearance Such a position neglects many other ‘mam- of the mentioned ice-shield along the moth’ and ‘woolly rhino sites’ where saiga, Siberian coast and, as a consequence, that of reindeer or both are absent (Putshkov 1989a, ‘mammoth ecosystems’. 1991, 1997). Biotists see the answer in the The basic arguments of the mentioned idea eurybiocity of the pachyderms and in the are the Eemian (125-115 kyBP) Greenland variability of ‘mammoth landscapes’ that ice-core properties (GRIP project results) that were created by climate and pachyderm were interpreted as reflecting the surprisingly

373 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

fast alternation of short-lived (from 20 to ones. It may reflect an intervening succes- 1700 years) warm and cold phases. The latter sional stage of a mixed mesic forest of cool- are said to be almost as cold as the full gla- temperate type that was fixed by large herbi- cial cooling. However, it has been conclusi- vores pasture activity. Furthermore, there are vely shown that these cores do not reflect the doubts whether the sedimentation period ind- true Eemian climate, as they are being altered eed corresponds to Eemian (Riss-Würm) by folding and other processes caused by ice sensu stricto: only the duration of this period flow and bedrock influences (Alley & Bender (9,7 kyr) is established varvometrically but 1998). The non-distorted Eemian and not the age of this vanished lake (ibid.). Holsteinian Interglacial layers of Antarctic The doubts seem even more justified when ice-cores (Vostok project results) do not show looking at uninterrupted sediment sequences any abrupt climatic flickering either. in deep-sea cores from various parts of the Other arguments are changes in Eemian North Atlantic. A number of the independent pollen spectra recorded in certain European indices show that during the Eemian sensu lacustrine and sea shelf sediments. However, stricto (125-115 kyBP: sea sediments stage it is not clear to what extent these changes 5e) there were no sharp changes of the mean correspond to the real intensity of the clima- paleotemperatures of deep-sea and surface tic oscillations, for they might reflect local water, as well as those of iceberg production succession events caused by the interactions rates. The same is true for the Holocene. The of biotic and abiotic factors. It is beyond Eemian border between polar and boreal question that, due to the presence of rhinos waters was constantly placed between and elephants, the particularities of the Greenland and Iceland, in the very region, interglacial vegetation successions could not where it is placed now (McManus et al. be the same as those of Holocene succes- 1994; Adkins et al. 1997). On the other hand, sions. So, the increase of herbaceous plants within the predominantly warm Early as revealed by the Eemian sediments of two Weichselian time (115-70 kyBP: stages 5d, lakes in the Massif Central in France 5c, 5b and 5a) there was indeed a series of (Thouveny et al. 1994) might be provoked strong cooling when the polar water border even by a weak decrease in humidity. approached Ireland, though not to Northern Superimposing upon the damage caused by as it did during the Middle and Late large herbivores, it could markedly oppress Weichselian stadial coolings (ibid.). trees in favour of grasses and herbs. Such Sher (1971, 1995, 1997b: 4-6) insists that interpretation seems plausible considering the the Arctic Ocean ice-shield has been destroy- concomittant changes in tree composition: an ed to a lesser degree during interglacials than oak/hazel-dominated community has been during the Holocene; notably the direct con- replaced by spruce/fir-dominated, and subse- tact of the mainland and pack-ices remained quently by pine-dominated ones; oak and intact even during summers in the East spruce, however, did not vanish (Thouveny et Siberian Arctic. He interprets the modest al. 1994). These changes by no means con- interglacial transgression rates in most of the firm the idea of an intra-Eemian ‘quasi-sta- East Siberian Arctic, compared to other dial’ cooling: spruce, pine and oak grow now Arctic regions, as sufficient proofs of this in the Massif Central mountains. Similarly, idea. However, even in East Siberia the traces the birch domination and spruce presence of these transgressions are found óver the episode recorded in ancient lake sediments present day sea-level. It means that some of near Bispingen, North-Western Germany the Interglacial transgressions and, therefore, (Field et al. 1994) does not necessarily repre- the extent of the Arctic Ocean ice melting sent the ‘Central Siberian’ rates of climatic were greater than the Holocene ones. severity; both taxa are not exclusively boreal Notably, the lowlands of Yamal, Gydan and

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Taymyr peninsulas were inundated during the mes, horses inhabited the rich open wood- Eemian, whereas their highlands (such as the lands indicating climatic conditions warmer Rranga mountains) became islands (Gus’kov and wetter than the Holocene ones (May 1986). Sher’s idea also contradicts other 1993; Kolfschoten 1995). The contradiction important evidence (Putshkov 1991, 1997). is eliminated when the inevitable impact of Sher (1995: 320) considers the eastwards mammoths and/or straight-tusked elephants is increase of steppic and/or tundra elements in considered: proboscideans with accompa- the Eemian paleofaunas, or the "eastward nying fauna converted forests to woodland/ ‘steppisation’ of the interglacial environment" parkland/grassland mosaics under any clima- as proof of the interglacial steppe-tundra per- tic conditions. Throughout the forest zone on sistence. He reckons all faunas including intensively grazed and sufficiently wet places mammoth, woolly rhino, bison and horse as zoogenic meadows originated, while the drier "mere insignificant variations of the steppe- places overgrew with a steppe-like vegeta- tundra faunistic complex" (Sher 1997a: 24). tion. The latter became more important Such point of view neglects the vast evidence towards the east and south where the climate proving the high eurybiocity of the mentionec became drier. But in the deep parts of the animals (see Putshkov 1989a, 1991, 1997 for forest zone the dense forest patches (though a review). The paleofaunas including these not as closed as now due to presence of species accompanied by steppe and/or forest pachyderms paths) also were alternated with dwellers could not be considered as steppe- considerable parkland, open bushland and tundra faunas if the predominantly tundra grassland areas of zoogenic origin. Not only species such as Lemnus, Dicrostonyx, ungulates, but even stenotopic animals living Alopex, Rangifer are absent. Apart from the now almost exclusively in steppes (Lagurus, other examples (ibid.), we mention the Citellus, Cricetus, Ochotona, etc.) or in tun- Eemian faunas with that inha- dras (Lemnus, Dicrostonyx, etc.) penetrated bited the warm forest-steppe of Eastern deeply into the forest zone and maintained Europe (Voronezh, Kuban’, Moldavia) there viable populations. Most likely such is (Alekseyeva 1980) and the cool-temperate the main reason of coexistence of forest, forest-steppe near Novosibirsk (Vassiliev tundra and/or steppic rodents in the intergla- 1995). cial faunas of Europe and Siberia mentioned Another fault of this position is the disre- by Sher (1995; 1997a). In some cases, howe- gard of the abundant evidence proving that ver, these mixed faunas do not belong to Eemian and some earlier interglacial opti- Eemian optimum stages, but to cooler earlier mum climates were milder than the present or later ones as, e.g., the Bielorussian faunas Holocene one (Putshkov 1991, 1997; rich in lemmings (Motuzko 1989). Kolfschoten 1995). It is noteworthy that The dissimilarities of interglacial ‘mammoth interglacial ‘steppisation’ or, to be more pre- biome’ vegetation with that of the modern cise, ‘prairisation’ has touched even England, taiga and tundra, and the persistence of pasture though its Eemian optimum climate surely ecosystems in Siberia throughout interglacials was milder than the present one (Stuart are also claimed as proofs of the interglacial 1991). The enormous aurochses and bisons, steppe-tundra persistence (ibid.). However, giant deer, steppe rhinos and hippos grazed these phenomena find their explanation in the here over vast meadows bordering with the impact of the pachyderms on the ecosystem. rich warm-temperate woods nourishing the The climatic interpretation would gain credibi- straight-tusked elephants and fallow-deer lity only if there were reliable and independent (ibid.). The same is true for the interglacial megafaunal indices showing sharp differences optimum faunas of continental Europe where between interglacial and Holocene climates. the mentioned large herbivores and, someti- But the situation is just the reverse.

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Consequently, the climatic paradigm is still (c) predation from wolves that multiplied stuck upon the question: why were numerous with the decline of lions, leopards and hye- pre-Holocene glacial/interglacial transitions nas, (d) local catastrophic events, epizootics, by no means as destructive for large mam- degeneration harmful to depressive popula- mals as the Weichselian/Holocene transition tions. is believed to be? (5) Southward retreat of lions, hyenas, leo- pards and dholes caused by the impoverish- ment of large prey aggravated by competition EXTINCTIONS IN PALEARCTIC with man (lion, hyena and leopard) or with There are detailed interpretations of these wolf that was previously suppressed by larger events from climatic (Vereshchagin & predators (dhole). Baryshnikov 1985; Guthrie 1990a, 1990b; (6) Other secondary extinctions among Sher 1995, 1997a, 1997b), half-and-half mammalian parasites and soil mites depen- (Stuart 1991) and ecological (Putshkov ding on the extinct herbivores activity. 1989a, 1989b, 1992b, 1993 a, 1993b, 1994) (7) New equilibrium installation. The points of view. According to the latter one the Holocene ‘wild’ ecosystems differ from the crisis in Palearctic included the same proces- Pleistocene ones first and foremost by the ses as in other realms: lack of giant herbivores and their ecological influences. (1) Liberation of mankind from being con- trolled by carnivores, resulting in the increase Due to the long co-evolution with man the of human populations and in certain carnivo- three first processes in the Old World were res extinctions. long-lasting and gradual. Therefore, conside- (2) Removal of giant herbivores by man rably more forms than in America could took place because pachyderms were vulnera- adapt themselves to environmental changes. ble to human predation due to their low reproductive rates. CONCLUSION (3) Drastic environmental changes caused The key reason of the maintenance of ‘mam- by this removal. Instead of pachyderms moth ecosystems’ were the mammoths them- sustaining mosaic woodlands, parklands, selves. Being highly eurybiotic, woolly mam- meadow and other associations, closed moths and rhinos could not be driven to forests, moss/shrublet tundras, and new step- extinction by the Early Holocene climatic pe type originated. Due to the lack of pachy- changes even if these changes, contrary to the derms, vegetation became everywhere less evidence, were unique events throughout the mosaic, fires more frequent and/or intensive. Quaternary. The ecological impact of mam- Animals could no longer use ‘mammoth moths prevented the extinctions of other paths’ through thickets and snow. megafaunal species. The climate-driven envi- (4) Secondary extinctions of herbivores ronmental changes became harmful for smal- (giant deer, musk-oxen, primeval bisons, two ler megafauna only when the pachyderms Central Asian antelopes, European asses, were gone or became too rare for buffering ‘forest’ races of horses and cave bears), the climatic influences. Human activity is the depending on environmental effects of the sole factor that could cause woolly mam- pachyderms took place chiefly due to the moths and rhinos to decline. mentioned changes added with the following factors: (a) competition with extant herbivo- ACKNOWLEDGEMENTS res that were preadapted for the pachyderm- The author very much appreciates the help less environment, (b) human hunting and bur- and financial support obtained from the ning activities and their ecological outcomes, Organising Committee of the 2nd Inter-

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national Mammoth Conference that made Janzen, D.H., 1984 - Dispersal of small seeds by big possible his participation in the conference herbivores: foliage is the fruit - American Naturalist and the publication of this article. He thanks 123 (3): 338-353 W. Schule (Freiburg), O.P. Zhuravlev, D.V. Kingdon, J., 1979 - East African Mammals. Vol. III, Ivanoff, A.G. Kotenko, V.M. Tytar, L.V. part B - London Puchkova and N.B. Puchkova (Kiev) for the Kingdon, J., 1982 - East African Mammals. Vol. III, useful discussions. parts C & D - London Kiselev, S.V., 1981 - Late Cenozoic Coleopterans of REFERENCES North-East Siberia - Moscow (in Russian) Adkins, J., Boyle, E., Keigwin, L. & Cortijo, E., 1997 - Kolfschoten, T. van, 1995 - On the application of fossil Variability of the North Atlantic thermohaline circula- mammals to the reconstruction of the palaeoenviron tion during the last interglacial period - Nature 390: ment of northwestern Europe - Acta zoologica 154-156 cracoviensia 38 (1): 73-84 Alekseyeva, L., 1980 - Peculiarities of the theriocomplex Kozhevnikov, Yu.P., 1977 - A critical review of the data of Russian Plain at the Last Interglacial - in: concerning the history of the Chukotsk flora - Anthropogene mammals of East Europe, Leningrad: Botanichesky Zhurnal N 3: 445-460 (in Russian) 68-74 (in Russian) Kozhevnikov, Yu.P., 1986 - On the question of the Alley, R.B. & Bender, M.L., 1998 - Greenland Ice-Cores: Beringean steppe-tundras existence - in: Frozen in Time - Scientific American 1998 (2): 66- of the Beringean sector of the 71 Subarctic, Vladivostok: 45-51 (in Russian) Chernov, Yu.I., 1980 - Life of the tundra - Mysl’, Krivolutsky, D.A. & Druk A.Ja., 1982 - Oribatid mites Moscow (in Russian) from the Siberian Quaternary Sediments - in: XI Field, M.H., Huntley, B. & Muller, H., 1994 - Eemian Congress INCVA (Abstracts) , Moscow, V. 1: 183- climate fluctuations observed in European pollen 184 record - Nature 371: 779-783 Lister, A. & Bahn, P., 1994 - Encyclopédie complète des Formozov, A.N., 1990 - Snow cover in the life of mammouths - Lausanne/Paris mammoths and birds - Moscow (in Russian) Lister, A. & Sher, A., 1995 - Ice cores and mammoth Foronova, I.V. & Zudin A.N., 1986 - New approach to extinctions - Nature 378: 23-24 research of fossil elephants of Archidiscodon- Martin, P.S., 1984 - Prehistoric overkill: the global Mammuthus lineage in North Eurasia - in: model - in: Quaternary extinctions. A prehistoric Biostratigraphy and paleoclimates of Pleistocene of revolution, Tucson, Arizona: 354-403 Siberia, Novosibirsk: 6-31 (in Russian) Maslov, S.V. & Antipina E.E., 1994 - On the role of the Gus’kov, C.A., 1986 - An essay of the construction of man in the changes of the Earth Biosphere at the the paleooceanic maps for the Late Pleistocene of the Pleistocene/Holocene border - in: All-Russian collo North Siberia on Foraminifera - in: Biostratigraphy quium for the study of the Quaternary Period, and paleoclimates of Pleistocene of Siberia, Moscow: 157 (in Russian) Novosibirsk: 111-115 (in Russian) May, T., 1993 - Beeinflussten Grossäuger die Guthrie, R.D., 1990a - Frozen fauna of the mammoth Waldvegetation der pleistozänen Warmzeiten steppe - Chicago Mitteleuropas? - Natur und Museum 123 (6): 157-170 Guthrie R.D., 1990b - Late Pleistocene faunal revolution McManus, J.F., Bond, G., Broecker, W.S., Johnsen, S., - a new perspective - in: Megafauna and man: disco Labeyrie L. & Higgins, S., 1994 - High-resolution very of America’s heartland: 42-53 climate records from the North Atlantic during the Haynes, G., 1991 - Mammoths, mastodonts and last interglacial - Nature 371: 326-329 elephants - New York Motuzko, A.N., 1989 - On the possibilities of the small Heptner, V.G., Nassimovitch, A.A. & Bannikov A.G., mammals fauna utilisation for the stratigraphy of the 1961 - Mammals of Soviet Union. V. 1. Upper Pleistocene deposits - in: Quaternary Period. Peryssodactyles and Artiodactyles - Moscow (in Paleontology and Archeology, Shtiinza, Kishinev : Russian) 44-52 (in Russian)

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