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1999-MAY16-MAY20
2nd International Mammoth Conference

ABSTRACTS (3)

edted by Jelle W.F. REUMER & John DE VOS
DISCLAIMER

The Abstracts of the 2nd International Mammoth Conference,
published in this Volume of Conference Papers, are not published
for permanent scientific record and are therefore not to be considered
as publications in the sense of Article 8 of the International Code of
Zoological Nomenclature. They merely reflect the opinion of the
author(s) at the moment of submission, and they should not be cited
in scientific works without prior written consent of the author(s).

After abstract titles (P) means poster and (L) means lecture (oral contribution)

Y.V. KUZMIN, LA. ORLOVA, I.D. ZOLNIKOV, A.E. IGOLNIKOV - THE DYNAMICS OF MAMMOTH (Mammuthus pnmigenius) POPULATION IN NORTHERN ASIA: RADIOCARBON EVIDENCE (P)

Irina E. KUZMINA, Sofia A. TSYGANOVA - Mammuthus primigenius IN THE URAL (P)

P. LAZAREV - THE MAMMOTH MUSEUM OF THE REPUBLIC SAKHA (YAKUTIA) ACADEMY OF SCIENCES (L)

P. LAZAREV - ANTHROPOGENIC HORSE DEVELOPMENT HISTORY IN NORTHEASTERN SIBERIA (P)

Sergei V. LESCHINSKY - THE CONNECTION OF MAMMOTH MIGRATIONS WITH GEOCHEMICAL LANDSCAPES OF Ca-, Mg-, Na-CLASSES IN THE SOUTHEASTERN PART OF WESTERN SIBERIA (L)

Adnan M. LISTER - EVIDENCE FOR STASIS AND SPECIATION IN THE 'GRADUAL' EVOLUTION OF Mammuthus IN EUROPE (L)

A.M. LISTER - THE DISTRIBUTION OF EURASIAN WOOLLY MAMMOTH COMPARED WITH VEGETATION MAPS FOR THE PLENIGLACIAL (C. 18 KY): EVIDENCE FOR WOOLLY MAMMOTH ECOLOGY AND CAUSES OF EXTINCTION (P)

K. MARKOVA - SMALL MAMMAL COMMUNITIES OF EASTERN EUROPE DURING THE DNIEPER AND VALDAI GLACIATIONS (P)

Evgeny MASCHENKO - MORPHOLOGY OF FIRST GENERATION TEETH IN Mammuthus AND ArchidiskodonAND EVOLUTION OF MAMMOTH AND SOME ASPECTS OF THE BIOLOGY OF M. primigenius (L)

George E. McDANIEL, Jr. & George T. JEFFERSON - Mammuthus mendionalis (Proboscidea.' Elephantidae) FROM THE BORREGO BADLANDS OFANZA-BORREGO DESERT STATE PARK╝, CALIFORNIA:PHYLOGENETICAND BIOCHRONOLOGIC IMPLICATIONS (L)

Dick MOL, Jelle W.F. REUMER, John DE VOS& Piet CLEVERINGA - ON REMAINS OF VERY SMALL (FEMALE) WOOLLY MAMMOTH, Mommuthus primigenius, FROM WESTERN EUROPE (L)

Alexander N. MOTUZKO - TIME OF APPEARANCE OF MAMMOTHS ON THE TERRITORY OF WESTERN SIBERIA (P)

M.R. PALOMBO - Bephas?Mammuthus?Loxodonta? WHO IS THE TRUE ANCESTOR OF THE DWARFED ELEPHANT OF SICILY? (L)

M.R. PALOMBO, A.P. ANZIDEI & A. ARNOLDUS HUYZENDVELD - LA POLLEDRARA Dl CECANIBBIO (ROMA): ONE OF THE RICHEST Elephas (Palaeoloxodon) antiquus SITES IN THE LATE MIDDLE PLEISTOCENE IN ITALY (P)

M. R. PALOMBO & P. VILLA - SEXUAL DIMORPHIC CHARACTERS OF ё (Palaeoloxodon) antjquus FROM 'GROTTE SANTO STEFANO' (VITERBO, CENTRAL ITALY) (P)

S. PEAN & M. PATOU-MATHIS - TAPHONOMY OF THE MAMMOTH SITES (L)

Olga POTAPOVA - SEDENTARY BIRDS IN THE LATE WORM ECOSYSTEMS OF THE NORTH AND MIDDLE URALS (L)

J.L PRADO, M.T. ALBERDI, B. SANCHEZ & B. AZANZA - DIVERSITY OF THE PLEISTOCENE GOMPHOTHERES FROM SOUTH AMERICA (L)

In Poland are known, so far, more than 400 localities of mammoth finds. They are located along the rivers Vistula and Odra. Most of them are located at the above mentioned rivers, as well as around the main scientific centers: Gdansk, Warszawa, and Krakow.

In most of the localities there are finds of teeth or portions of the skeleton. Among them are completely preserved skulls. Two of them are dated by C 14 as to be 36,000 and 14,000 y old. One of them belongs to the latest mammoth in Europe (about 14,000 y BP). It shows abnormally developed tusks and is very in comparison with other mammoth skulls. However, it is a skull of an adult mammoth with dwarfing characteristics. In collections known by the author, there are also finds which show a dwarfing respect. That means that the dwarfing in the mammoth took place and occurred in some periods and under specific conditions in geographic regions, and also in specific environmental and nutrition conditions. European localities of mammoth finds are compared with the Polish finds.

Since the 1960s, a significant amount of radiocarbon (C 14) dates was obtained for mammoth remains in Northern Asia, which covers Siberia and adjacent territories of the Russian Far East, Kazakhstan, northern Mongolia, and northeastern China Using the information available as of 1998, we can now reconstruct main features of spatial temporal dynamics of mammoth population in Northern Asia for the last 40,000-50,000 C14 years. To do this, we applied Geographic Information System (GIS) technology. Using C14 mammoth date lists (Sulerzhitsky 1995, 1997, Sulerzhitsky & Romanenko 1997, Vasil'chuk et a/. 1997), we added information for Kazakhstan, Russian Far East, and northeastern China as well as dates from Siberian Paleolithic sites, and some unpublished data. In total, we collected 3 IOC14 dates from more than 140 localities.

Using the sub-division of the Siberian Late Pleistocene (Kind 1974), we generated maps of spatial distribution of C 14 dated mammoth remains for several time intervals: I ) Early Karginian stages, ca 55,000-33,000 yBP; 2) Konoschelye stage, ca. 33,000-30,000 yBP; 3) Lipovka-Novoselovo stage, ca. 30,000-24,000 yBP; 4) transition to Sartan time, ca. 24,000-20,000 yBP; 5) Sartan Glacial maximum, ca 20,000-18,000 yBP; 6) Late Sartan time, ca. 18,000-13,000 yBP; 7) Kokorevo stage, ca. 13,000-12,000 yBP; 8) Taimyr stage, ca. 12,000-1 1,000 yBP; 9) Norilsk stage, ca. 1 1,000-10,000 yBP; 10) Preboreal period, ca 10,000-9,000 yBP; 1 1) Boreal, Atlantic, and Subboreal periods, ca 8,000-3,700 yBP. The C 14-dated mammoth remains in Northern Asia concentrate into two latitudinal 'belts', the first one in the Arctic, and the second one in the southern part of both Western and Central Siberia. However, this does not mean that the mammoth habitat was divided into two parts, northern and southern ones. In our opinion, the rnammoth habitat in Late Pleistocene covered whole Northern Asia. For the intervals 55,000-33,000 yBP, 33,000-30,000 yBP, 30,000-24,000 yBP, and 24,000-20,000 yBP, 20,000-18,000 yBP, 18,000-13,000 yBP, and 13,000-12,000 yBP, mammoth existed throughout all of Northern Asia. There is no significant difference in the size of mammoth habitat between warm and cold stages; for example, Lipovka-Novoselovo and eariy Sartan Glaciation.

The significant changes in size of the mammoth habitat started in Northern Asia at ca. 12,000 yBP, During this time, ca. 12,000-10,000 yBP, rnammoth settled only lowlands near the Indigirka River mouth, Taimyr Peninsula, and the Sevemaya Zemlya Islands. Around 10,000 yBP mammoth extincted in the Indigirka River basin, and survived only on the Taimyr and Gyidan Peninsulas. The latest C14 dates from these territories are ca. 9,780-9,670 yBP foi-Taimyr and ca. 9,730-9,600 yBP for Gyidan. This supports the earlier conclusion made by L. D. Sulerzhitsky ( 1995, 1997) and A. V. Sher ( 1997) about the sharp decrease of mammoth habitat in Siberia at ca. 12,500-12,000 yBP. In the Holocene, ca 7,700-3,700 yBP, mammoths lived only in Wrangel Island. However, the size ofthose mammoths was rather small compared with typical Mammuthus pnmigenius Blurn. The Holocene rnamrnoth population was described as separate subspecies M. primigenius vrangeliensis (Vartanyan et al. 1993). The C 14 Database allows to estimate the amount of mammoth 14C dates correspond to cold and warm stages within the Karginian, Sartan, and Holocene times, ca 45,000-3,700 yBP. In total, there are 289 C 14 values for this time span. For the warrn stages such as the Holocene, Lipovka-Novoselovo, and Malaya Kheta, the percentage of
dates is 10.4, 15.9, and 22.4, respectively. For the cold stages such as Sartan, Konoschelye, and Early Karginian, the amount of dates is 17.6%, 9.0%, and 4.2%, respectively. 20.5% of the dates correspond to transitional Late Glacial time, ca. 1 8,000-10,000 yBP. Thus, 48.7% of the dates may be correlated with warm stages, and 30.8% referto cold ones. Most probably, the increase amount of dates for the warrn stages may be explained first by favourable taphonomic conditions, because of active solifluction, burial, and consen/ation of mammoth remains as it was noted previously by L. D. Sulerzhitsky ( 1995, 1997). This also means that it is not so evident that warrn environmental conditions were more favourable for the mammoth population than cold ones.

 During the Late Pleistocene mammoths were widely distributed in the Ural Mountains region from the coast of the Arctic Ocean to - on the average - the current of the Ural nver (fig. I ). We investigated more than 4500 fossil mammoth bones. The most numerous congestion of bones is known from the river Usa, the right tributary of the river Pechora.' Mamontovaya Kuria (34,010 ╠ 485 yBP), on the river Pechora the site Byrovaya (25,740 ╠ 500 yBP), and on the river Sos'va near the village ofGari, in the Ekaterinburg area, Paleontological collection in museums in the towns of Ekaterinburg, Ufa, Nizhni Tagil, and others were also investigated. Many ofthese concern collections of skeleton, but also isolated bones and teeth of the mammoth. On the territory of Bashkiria, at the nver Belaya, two caves are located: Kapova cave ( 14,680 ╠ 150 yBP) and Ignatievskaya cave ( 14,240 ╠ 150 yBP). In these caves, images ofMammuthus primigenius and others Late Pleistocene mammals are drawn on the walls. On the eastern slopes of the Urals, at a village called Karelino (Verhoturskiy region, Ekaterinburg area) there is a rock of 50 m length and 15 m height, on which 34 animal figures are represented.

Fossil mammoth teeth from all ages classes are found in the Ural region. The deciduous teeth dP2, dP3, dP4 and the MI,
M2, M3 are known. M3 length is 230-274 mm; m3 length is 245-326 mm. The tusk fragments range from 0.9-2.6 m in
length. Their diameter ranges from 63 mm up to 187 mm, and the circumfence from 220-560 mm. The comparison of skeletal
size extremes shows that animals inhabiting the Ural region were larger than in the centre of the Russian plain.

 The foundation of the specialized Mammoth Museum in Yakutia in 1991 was initiated by M. Nicolaev, the President ofthe Republic Sakha (Yakutia). The foundation of the museum in our northern republic was quite reasonable as a great part ofthe world's famous finds of mammoths, woolly rhinoceroces, Panthera (Leo) spelaeal, and other fossil remnants of Pleistocene mammals were made in permafrost ground within rts territory. The main direction ofthe museum's activities are the following: the work ofthe museum, scientific research, and international relations. There are 4 departments in our museum exibition: I. Research history and Pleistocene; 2. the Mammoth; 3. the Mammoth fauna; 4. Mammoth and man.

The exhibitions tell about the famous Berelekhsky mammoth cemetery, where about 10.000 mammoth fossils were excavated in 1970, including a mammoth leg with hair, etc., and where a srte of Pleistocene hunters was discovered. The museum exhibits skeletons ofthe Diring mammoth and the Churapcha rhinoceros with skin and hair, remnants ofthe hind leg, the mummy ofthe Dukar horse, the inner organs ofthe Shandrin mammoth, a leg with hair from the Bolshoi Lyakhov island, etcetera. Museum research workers study the evolution of Pleistocene mammals, faunal changes on the verge ofthe Pleistocene and the Holocene, the origins ofthe modern Yakutian fauna, climate and vegetational changes of various stages ofthe Quartemary.

Exhibitions of Siberian mammoths held in Japan, South Korea, France and Germany can be mentioned as examples of our international cooperation. Paleontological expeditions were organized jointly with French, German, and Japanese colleagues. Japanese and German cinematographists shot professional films on mammoth sites in Yakutia. The Mammoth Museum became a member of the Museum Council in 1992, promoting its international activities.

Representatives of Equus appeared in Northeastern Siberia in the Eopleistocene period, coming from North America. At that time horses ofthe genus Plesippus were widely spread in North America (W. Matthew 1924). Kretzoi (1938) grouped precaballoid archaic horses into the subgenus Mohippus. Considering that ancestral forms of Asian horses came from North America and moved further to the west, to Europe, and that the subgenus name Plesippus was given by Matthew earlierthan by Kretzoi, this name should be applied to aJI Pleistocene-Eopleistocene precaballoid horses of America and Eurasia.

The rest population of these horses in Northeastern Siberia was represented by ё. (P.) verae (Sher 1971). The evolutionary descendant of this horse was ё. (P.) coHemensis (Lazarev 1980). These Eastern Siberian horses, having common subgeneric features, still differed considerably from contemporaneous European horses. ё. (P.) verae had a large skull and postcranial skeleton, a short subtriangular protocone, and a characteristic hollow on the mesostyle.(P.) coliemensis was smaller than ё. (P.) verae, had a broad forehead and la ong skull with a big axial fracture anda specifically narrow occipital crest. Teeth were short with small folds on the walls of fossae. They lived under the conditions of mildly cold climate, at the time of permafrost formation and the development of 'forest-and-steppe' and forest landscapes. It is worth mentioning that E. Vangengeim (1961) found isolated fossils of ё. (P.) sonmeniensis and ё. (P.) cf. stenonis on the Aldan, but additional research on these horses is necessary.

A phylogenetic line of true horses ofthe subgenus (Lquus is found in the second part ofthe Early Pleistocene in Northeastern Siberia. It consists of Early Pleistocene ё. nordostensis RUSS., Middle Pleistocene ё. onentaiis RUSS. and Late Pleistocene ё. lenensis RUSS.. These horses, like the horses of subgenus Plesippus lived in Eastern Siberia, separated from horses of Western and Southern parts of Eurasia. This separation was caused by climatic changes during the Pleistocene from moderately cold to markedly severe continental, frequent changes of moist glacial and dry interglacial periods, transgressions and regressions, permafrost formation and other extreme living conditions.

The described true Pleistocene horses, beginning with the large Early Pleistocene ё. nordostensis possess charactenstic features: broad foreheads and long muzzles in the skull, an elongated protocone in the upper molars and an assymetrical form of the loop of lower teeth. The horses gradually grew smaller, e.g. ё, nordostensis had a whithers height of 1,7 m, while ё. lenensis was 1,3 m high.

 Investigations concerned the periglacial and extraglacial zones with sharp macroclimatic fluctuations in the Pleistocene. The change of landscapes passed quickly from tundra to steppe and vice versa, practically wrthout a phase of forest distribution. The high density ofherbivore populations (horse, bison, woolly rhinoceros, and others) and the seasonal prevalence in nutntion forced mammoths to migrate large distances. Scientists estimate mammoths' migrations to 650-2500 km (Germonpre 1993). It possibly explains the emergence of arctic populations in the Holocene. Analogies between mammoth and Loxodonta africana are, however, problematic. Movements of African elephants are restricted, although they can move in the Narnib desert for four days without water. In Kenya they cover more than 80 km to places with enough precipftation bringing an abundance of fresh vegetation (Germonpre 1993).

The normal metabolism is made possible by the constant influx of specific chemical elements from the surrounding environment. Deficiency in these elements causes mineral starvation directly connected to lithophagy -the use of rocks, minerals and mineral waters (Panichev 1990). We consider bnefly the mineral starvation on the example of three elements: calcium, magnesium and sodium (Ca, Mg, and Na). Calcium is a major metal in the mammalian organism as it forms the calcic skeleton. Experiments with labelled calcium indicate the continuous exchange of calcium from bones to calcium from food (Perelman 1972). A Ca deficiency causes bone brittleness, rachitis, the loss of breed, and sharply decreasing grov^th. A Mg deficiency causes the heavy disease hypomagnesemia, revealing in convulsions of skeleton muscles and a death that follows quickly. Sodium (Na) losses in the organism heavily affect the nervous system and lead to fatigue and subsequently to emaciation and death (Panichev 1990). Mineral starvation is especially developed in acid gley landscapes of taiga and tundra. The largest Ca, Mg and Na deficiency is observed there, as the elements are easily leached from soils and from the weathering crust. The starvation peak
appears in spnng when there is much potassium (K) in fresh vegetation (c. 1000 times more than in winter). Potassium, being the antagonist of sodium, sharply decreases the level of the latter in the organism, leading to the development of'sodic stress'. Animals eat clay, which has adsorbing properties with respect to potassium, in addition to rocks enriched in Ca, Mg, and Na for the maintenance of the water-salt homeostasis. Elephants can, for example, eat more than 20 kg of rock at atime. Reindeer are known to eat rodents, birds, eggs, fish, and to gnaw bones and drink sea-water as an addition (Perelman 1972).

Mammoth {Wammuthus), the largest representative of the Pleistocene terrestrial fauna in Northern Eurasia has the largest trunk and skeleton compared with other fossil Elephantidae. It therefore felt acute need for mineral nourishment. The wide development of tundra-taiga zones with permanently frozen ground predicts periodical mineral hunger in the animals. Then, accumulative-autonomous and superaqualine landscapes enriched in Ca, Mg and Na played a targe role in the nourishment. The first type is characterised by the removal of weathering products by surface and melting waters to low-lying areas, the latter type by the close occurrence of ground waters. Such landscapes served as the original 'mineral oases' on the migration routes. Rising temperatures or moistening of the climate caused the erosion of rocks enriched in spore elements and the accumulation of talus deposits in depressions. In such humid lowlands the vegetation was impetuously developing, greedily absorbing dissolved forms of mineral compounds (modem analogues are landscapes of the southern tundra on limestones).
Just such places were attractive for mammoths by the abundance of the nourishing forage and the moisture. With falling temperatures migration becomes less important and the main 'mineral oases' are superaqualine landscapes. Groundwaterthen supplies necessary elements to the surface, and the permanently frozen ground is the main geochemical factor. Frozen rocks contain water with negative temperature next to the ice. Such water migrates in the direction of lower temperature: in the winter and autumn to the surface, in the spring and summer in the opposite direction. Necessary elements are supplied from bedrocks into the active layer and then they migrate to the surface by cryogenic intermixing (Perelman 1972). Probably, the claystone was eaten at such periods (bowels of the Kirgilyach mammoth were full of clay). By macroclimate changes (fall and rise of temperature) some landscapes of the Ca-K class were preserved. The indicator of high mineral energy are the plants-calcicoles, widely spread in the palynospectra of deposits (Leschinsky 1998). The development of Ca-, Mg-, and Na-landscapes became possible as a result of the presence of Prequaternary carbonaceous-argillaceous rocks, loess-like loams and sediments of dammed reservoirs enriched by migrating spore elements. Neotectonic movements are not unimportant either, as rock jointing, faults and landslips favour the outcropping of salty waters. In some cases (by the definite relief) it promoted the development ofsolonetzes for beasts (e.g., foothills ofKuznetsky Alatau, crests of Barabinskaya steppe). The global peaks of the mineral starvation occurred, apparently, in cold periods when mass concentrations of animals were observed on solonetzes. At some 'oases' the percentage of mortality and the bunal conditions were sufficient for the formation of thick bone-bearing horizons. The largest famous localities are 'Volchya Griva' in the Barabinsk steppe (more than 5000 bones, Na-Ca-solonetz?) and 'Schestakovo' in the Kemerovo district (more than 3000 bones, Ca-Mg-solonetz?) (Derevianko & Zenin 1998, Leschinsky 1 998).

During the Pleistocene the area occupied by Ca-, Mg-, Na-landscapes changed greatly. In cryochrons it was abruptly reduced because of dammed basins and gleying of soils, and it had received the maximal development in thermochrons as a result of draining of the dammed resei^/oirs and fluvial plains. Mammoth migration paths changed too, apparently. In cold epochs of the Eo-Pleistocene ( 1.8 - 0.8 My), migrations were apparently restricted by the Kuznetsky Alatau, the Kuznetsky trough, the Salairsky ridge and the Near-Obskaya elevated plain. During periods with rising temperatures they had rarely overstepped the limits of present river basins: Ob (~up to 57° N), Tom, Ya/a, Kiya, Chef, Chulim (~up to 56° N). In the Early, and in the main part of the Middle, Neopleistocene (0.8 - 0.2 My) the same situation was maintained, but in warm periods numerous populations of animals had actively passed into the basin of the present Ob (~up to 59°30'N). At the end of the Middle-Late Neopleistocene (0.2 - 0.01 My) the whole of the territory had been covered by migrations, regardless of macroclimate fluctuations but related to the river basins: Ob (~up to 58° N), Tom, Yaya, Kiya, Chulim, as well as the Barabinsky steppe. By calculating migrations it will be allowed to search for new localities with fossil mammals. The most important localities are, in the author's opinion, the places of the confluence and the intersection of migration paths in the junctions of present rivers: Berd - lnya - Ob, Yaya - Kiya - Chef - Chulim, Tom - Ob, Shegarka - Ob, Chaya - Ob, Parabel - Ob and Vasyugan - Ob. Fossil bones have periodically been found at these areas. Taking into account the dependence of ancient people on large mammals (Leschinsky 1998), the distribution ofPaleolfthic localities is also predicted. The main point of such prognosis is the comparison of animal migration routes with the sources of stone material suitable for the instrument manufacture. The most ancient encampments are apparently situated in river basins: Ob (the right bank ≈57 N), Berd, lnya, Tom, Yaya, Kiya, Chef, and Chulim (~up to 56°30' N) rivers.

Derevianko, A.P. & Zenin, V.N., 1998 - On the problem of 'human being and mammoth': geoarchaeological aspect - in: Pleistocene paleoecology and cultures of the Stone age in North Asia and contiguous territories [in Russian] I: 92-99

Germonpre, M., 1993 - Taphonomy of Pleistocene mammal assemblages of the Flemish Valley, Belgium - Bulletin de I'lnstitut Royal des Sciences Naturelles de Belgique, Sciences de laTerre, 63: 271-309

Leschinsky S.V., 1998 - Geology and paleogeography of the late paleolithic encampment 'Schestakovo'. Pleistocene paleoecology and cultures of the Stone age in North Asia and contiguous territories [in Russian] 1: 209-220

PanichevA.M., 1990 - Lithofagy in the life of animals and human being [in Russian], 224 pp.

Perelman A.L, 1972 - Geochemistry of elements in the hypergenesis zone [in Russian], 288 pp.

 The classic evolutionary sequence of mammoth in Europe is based on type samples of/VI. mendionafis from the Upper Valdamo, Italy (c. 1.7 My) and M. trogontherii from Sussenborn, Germany (c. 0.6 My), and Late Pleistocene M. prim/genius (c. 100-10 Ky). The addition of independently dated samples of preceding and intermediate ages is allowing more detailed appraisal of the pattern of change (Lister 1996). In molar morphology, the earliest substantive European samples, from the Red Crag (UK) and Montopoli (Italy), c. 3-2.5 My, are significantly more primitive (9-10 plates in M3) than typical /V\.  eridionalis ( 12-14 plates) (Lister & van Essen, in prep.). There is little further substantive change until late meridionalis, c. 1.0-0.6 My.

The nature of the shift from M. meridionalis to M. trogontherii in Europe is still unclear. Some evidence from sites in Britain (Beeston and West Runton) and Germany (Voigtstedt and Karlich) suggests an overlap in the chronological occurrence of the latest meridionalis and earliest trogontherii, implying a speciation event, as hinted at by Azzaroli (1977) on the basis of skull morphology. The picture is further complicated by the existence of specimens in some respects morphotogically intermediate between these two forms, yet supposedly older that Voigtstedt (Siniaya Balka, Russia: Dubrovo 1964).

The transition from typical early Middle Pleistocene trogontherii to typical Late Pleistocene primigenius has been the source of much debate and confusion. Some authors have suggested late Middle Pleistocene mammoths as a late (advanced) form ofM trogontherii (e.g. M. trogontherii chosaricus DUBROVO, 1966): others see as an alternative (e.g. M. primigenius froosi DIETRICH, 1912) or as a later addition (e.g. Gi-omov & Garutt 1975) an early (primitive) form ofM firimigenius. Detailed study of samples dated to c. 400 Ky (Steinheirn, Germany) and c. 200 Ky (Ilford, UK) indicates that in essential morphology (especially plate number), these mammoths were similar to typical M. trogontherii (e.g. Sussenborn, c. 600 Ky). However, gradual size reduction 600-200 Ky has camouflaged this stasis and given the impression of evolutionary advancement, by its effect of compressing lamellar spacing and thereby elevating lamellar frequency and depressing enamel thickness. When the size factor is removed, these samples are seen to be only slightly advanced overtypical M. trogontherii.

Because of the paucity of well-dated cold-stage samples, the timing of first entry of advanced M. primigenius into Europe is uncertain. At two sites in the UK (Marsworth and Brundon), only slightly post-dating Ilford (c. 200 Ky), two types of mammoth appear to be present in a single honzon. The first is similar to the late trogontherif of Ilford, the second to Late Pleistocene primigenius. Although precise synchroneity of such a co-occurrence can never be proven, the evidence suggests chronological overlap of the two forms and, by implication, a speciation event. The more primitive of the two forms apparently subsequently died out, afthough the possibility that it contributed genes to Late Pleistocene primigenius cannot be discounted.

Azzaroli, A., 1977 - Evolutionary patterns ofVillafranchian elephants in central Italy - Atti Accad. Lincel, Mem., Cl. Sc. Fiz., Mat., Nat. Ser. 8, 14 (lla): 149-168

Dietrich, W.O., 1912 - Bephas primigenius Fraasi, eine schwabische Mammutrasse - Jahresh. Ver. Vateri. Naturk. Wurttemb. 68:42-106

Dubrovo, I.A., 1964 - Elephants of the genus Archidiskodon in the USSR - Pal. Zh. 3: 82-94

Dubrovo, I.A., 1966 - Systematic position of an elephant from the Khozar faunal assemblage - Byull. Kom. lzuch. Chetv. Per. 32:63-74

Grornov, V.I. & Garutt, V.E., 1975 - Mandibel-Reste einer Fruhform des Mammuthus primigenius (Blum.) von Weimar-Ehringsdorf- Quartarpalaontologie 1: 453-464

Lister, A.M., 1996 - Evolution and taxonomy of Eurasian mammoths - in: Shoshani, J. & Tassy, P. (eds.) - The Proboscidea - pp. 203-213. Oxford: OUP

Lister, A.M. & Joysey, K.A., 1992 - Scaling effects in elephant dental evolution: the example of Eurasian Mammuthus - in: Smith, P. & Tchernov, E. (eds.) - Structure, Function and Evolution of Teeth - pp. 185-213. Jerusalem: Freund

 A distribution map for the woolly mammoth in Eurasia has been compiled from the work ofKahIke (1 994), Liu & Li (1984), personal data, and Chang (pers. comrn.). The southern limit of this map is assumed, as aworking hypothesis, to represent approximately the southernmost extent of the species' distribution in the Late Pleistocene, presumed to have occurred during the maximum pleniglaciat (c. 18 Ky). Work by palaeobotanists, based primanly on palynological data, is producing increasingly refined vegetation maps for the Late Pleistocene of Eurasia (Adams & Faure 1997), A simplified vegetation map for 18 Ky has been superimposed on the woolly mammoth distribution. There are some very interesting correspondences between the southern limit of the mammoth's range, and vegetational boundaries. These studies can provide information on the ecological tolerance of the woolly mammoth. They may also bear upon the question ofthe cause of extinction on the Eurasian continent, since they may indicate whether or not habitat types which remained in the postglacial were selected by woolly mammoth even in the Late Pleistocene.

Kahike, R.-D., 1994 - Die Entstehungs-, Entwicklungs- und Verbreitungsgeschichte des oberpleistozanen iVlammuthus-Coetodonta-Faunencomplexes in Eurasien (GroBsauger) - Abh. Senck. Naturf. Ges. 546: I -1 64

Liu, Tung-Sheng & Li, Xing-Guo, 1984 - Mammoths in China - in: Martin, P.S. & Klein, FLG. (eds.) - Quaternary Extinctions: a Prehistoric Revolution - pp. 517-527, Tucson, Univ. Arizona Press

Adams, J.M. & Faure, H. (eds.), 1997 - R.eview and atlas of palaeovegetation: preliminary land ecosystenn maps of the world since the Last Glacial Maximum. http://www.esd.oml.gov/em/qerl/adams I .html. Oak Ridge National Laboratory, TN, USA

 Small mammal faunas of glacial epochs (Dnieper = Saale; Valdai = Weichsel) have been analysed using the electronic data base SMALLMAM (PARADOX, v.4); the data set compnses information on the Pleistocene small mammal localities over Eastern Europe (Markova 1982). When analysing species composition, distribution and diversity of the small mammal faunas, the ARC/INFO and ARC/VIEW cartographic programs were widely used. The faunas under consideration, dated to the Dnieper and Valdai glacial epochs, differ in their evolutionary level and belong to different assemblages, viz. to the late Khozar and the Mammoth faunal assemblages, respectively.

Dnieper glacial epoch

At least three distinct communities of small mammals have been reconstructed that may be correlated with the Khosar faunal assemblage (Fig. I ): a subarctic small mammal community including Dicrostonyx simplicior, Lemmus sibiricus, Microtus (Stenocranius) gregolis occurred close to the ice sheet; farther south they gave place to a community of periglacial steppe distinguished by the coexistence of subarctic (Dicrostonyx, Lemmus), steppe {Lagurus ex gr. transiens - lagurus, E.olagurus luteus, Allactaga, Spermophitus, Marmota, Ochotono pusillo), and intrazonal species (Arvicola chosancus, Microtus oeconomus). There was a smalt area of periglacial forest-steppe in the west of the Russian Plain (with Lagurus, Eolagurus, Microtus arvolis, Clethnonomys gloreolus and others). In the south a steppe mammal community presumably developed.

Valdai glacial epoch

Not less than four well-defined small mammal communities attributable to the mammoth assemblage by their evolutionary level may be recognised at the time of the last maximum cooling (24 - 15 KyBP). They are as follows (Fig, 2): a subarctic tundra community of solely subarctic mammals (Dicrostonyx gulielmi, Lemmus sibiricus, Microtus gregalis) inhabited a narrow zone adjacent to the ice margin. South of it occurred mammalian species characteristic of the subarctic zone, together with typical steppic animals and a number of forest eurybiont species, forming a periglacial forest-tundra community with Dicrostonyx gulielmi, Lemmus sibincus, Clethnonomys gloreolus, Microtus gregalis, Microtus agrestis, Lagurus lagurus, Spermophilus, Mormota, and others ). South of that zone zoocoenoses were dominated by steppic species, though there were also subarctic small mammals present in small proportion and some woodland and meadow species. Such was the periglacial forest-steppe community including Dicrostonys gulielmi, Microtus gregalis, Allactaga major, Arvicola terrestris, Cricetus cricetus, Cricetuius migratorius, Spolax, Lagurus lagurus, E.olagurus luteus, Clethnonomys glareolus, Microtus agresVs, Microtus orvalis, Microtus oeconomus. The southern limit of this community ran along 48°N in the west of the Russian Plain and retreated northward in the east. It roughly coincides with the southern permafrost limit (Nechaev 1986). Finally, in the south of the Russian Plain small mammals show no effect of the ice sheet: a steppe small mammal community with Ochotona pusilla, Spermophilus, Marmota bobac, Allactaga, Alactagulus, Pygerethmus, Spalax, Cricetuius migratorius, Cricetus cricetus, Lagurus lagurus, ELolagurus luteus, and others was developed everywhere south of 48°N.

The structure of periglacial, forest-tundra and forest-steppe communities resembles that of communities within large ecotones. Characteristically, the cold intervals of the Pleistocene featured a specific 'indistinct' zonalitythat was essentially different from that of interglacials. Only subarctic species are found within a narrow belt ( 100-1 50 km) along the ice margin. This community was similar in composition to modem tundra zoocoenoses. It may be called harmonious in structure, though its area was shifted southward due to the glacier advance. These harmonious faunas are exempted from the rule. In the south they bordered on zoocoenoses that have no analogue at present: those were communities of periglacial tundra and forest-steppe resulted from subarctic species penetration from the north and steppe mammals from the south; they could be formed after the forest
biome had disintegrated. The faunas underconsideration are known as 'intermingled', 'disharmonious', or 'periglacial' (Vangengeim 1977, Sernken 1983, Graham 1976, Baryshnikov& Markova 1992). Southernmost regions of the Russian Plain still featured steppe zoocoenoses, the latter differed from their modem analogues chiefly in the large mammal composition. The ice sheet impact was negligible there. Therefore, the periglacial faunas dated to the Dnieper and Valdai glacial epochs were distinguished by the joint habitation of animals which belonged to different ecologic groups and different natural zones. A majority ofpenglacial communities have no modern analogues and could exist in mosaic periglacial environments. Those specific communities resulted from various mammals migrating in opposite directions (mammals of the subarctic zone moved from the north, while steppe endemic species advanced from the east and south). It was also essential that the continuous areas of woodland mammals had disintegrated, and many species typical of broad-leaved forests retreated southward into the
mountains.

 The evolution of mammoth-like elephants (genera Mornmuthus and Archidiskodon) is connected with their adaptations to cold climatic conditions of Eurasia during the Pleistocene and was accompanied with morphological changes of teeth and postcraniaJ skeleton (Lister 1996). The following dental morphlogical features are significant: growing hypsodonty, growing number of plates constituting the crown, diminishing enamel thickness, etc. (Maglio 1973). The systematics of the mammoth-like elephants is still under discussion because the changes in dental and skeletal morphology are interpreted differently (Garutt 1986, Baigusheva& Garutt 1987, Dubrovo 1994, Todd & Roth 1996). To clarify these problems, the morphology of stages of the first teeth generation (dp2-dp4) in Mammuthus and Archidiskodon were studied.

It was found that a comparative study of the morphology of first generation teeth in Mammuthus and Archidiskodon revealed many similar features. In both genera all parts of the dp2 crown show simultaneous development. Beginning in dp3, the arrows show a developmental sequence from the anterior part of the tooth to the postenor. The difference in plate number and enamel thickness of dp3 between Marnmuthus and Archidiskodon is also manifest in the dp4 through N3 generation. The structure of first generation tusks (di) in /V\ammuthus and Archidiskodon is also similar and reminds of the non-specialised incisors of other mammals. The second generation tusks (1) are highly specialised incisors typical of representatives of the family Etephantidae. The position of di in its alveola, the structure of the latter and the presence ofatveola of a rudimentary tusk in M primigenius suggest homologies of the rudimentary tusk with the incisor di I, of the first generation tusk (di) with dl2 and of the
permanent tusk with 13. Minor differences in the structures of first generation teeth and tusks between Mammuthus and Archidiskodon suggest their late differentiation into genera within the group of mammoth-like elephants.

Analysis ofgrov/th dynamics in M. primigenius calfs from the Late Pleistocene Sevsk locality population and similarities in some morphological features of/VI. pnmigenius and modem elephants, as well as some indirect evidence from elephant group structure, provide an opportunity to reconstruct peculiarities of the biology and ethology of the woolly mammoth (Agenbroad 1990, Haynes 1991, Maschenko 1993). Shoulder height of a newborn mammoth calf varies from 700 - 800 mm. It is less than in ё. maximus (780 - 950 mm) and much less than L africana (900 - 1050 mm) (Stenley 1943, Sikes 1971). Changes in dimensions of the humerus and femur during the first years of life show a different speed of growth in M. primigenius calfs, corresponding to different stages of physiological development. Judging by absolute grov^th of (dimensions of) humerus and femur one may discriminate between stages of (1) immediately proceeding and following birth, (2) maturation and (3) maturity.
The speed of growth is the greatest during the prenatal stage, whereas dunng maturation (8-10 years) it significantly diminishes. In M. primigenius females, growth stops completely at the age of 17-20, whereas in males it goes on, although at a practically reduced speed, untill 35-40 years. Changes in dimensions of growing bones of calves show that dunng the first year of life the speed of growth in mammoth was higher than that in L africona calves. The height of the body changed from 75 cm to 1 15 cm (L africana 90 cm and 120 cm, respectively). Faster growth of mammoth during the first year of life is connected with seasonal climatic changes and cold winter. Born in spring, mammoth calves had to grow fast in orderto survive in winter. Earlier wearing of dp2 and dp3 in mammoth calves in companson to elephant calves (Roth & Shoshani 1988) is indicative of their earlier feeding on vegetation, because of reduced lactation in females as a result of lack of green fodder in the winter.

The analyses of the Sevsk Late Pleistocene mammoth population (absolute dating 14,000 y) is indicative of earlier maturation in mammoths, causing earlier slowing down and cessation of growth. That might be one ofthe causes of the smaller size of Late Pleistocene representatives ofMammuthus pnmigenius, in comparison to earlier mammoth. The phenomenon is thus connected with species survival strategy. This biological strategy, together with morphological peculiarities provides on additional reason to segregate the genera. Mammuthus and Archidiskodon.

Agenbroad, L.D└ 1990 - The mammoth population of the Hot Springs Site and associated megafauna - in:

Agenbroad, L.D└ Mead, J.D. & and Nelson, L. (eds.) - Megafauna and Man: The Discovery of America's Heartland - pp. 5-32. The Mammoth Site of Hot Springs, Hot Springs, South Dakota

Baigusheva, V.S. & Garutt, V.E., 1987 - The skeleton ofArchidiskodon trogontherii (Pohlig, 1885) from the north-eastern Azov sea coast. Transactions Zoological Institute of USSR 186: 21 -37 (in Russian)

Dubrovo, I.A., 1994 - Fossil elephants from the Commonwealth of Independendent States - in: Agenbroad, Larry & Mead, jim (eds.) - The Hot Springs Mammoth S'rte - pp. 426-451. The Mammoth Site of Hot Springs, South Dakota. Freske Printing, Inc., Rapid City

Garutt, V.E└ 1986 - The origin and phitogenetic relationships ofElephantidae. Transactions Zoological Institute of USSR 149: 15-32 (in Russian)

Haynes, G., 1991 - Mammoths, mastodonts and elephants, Cambridge University Press, 413 pp.

Lister, A.M., 1996 - Evolution and taxonomy of Eurasian mammoths - in: Shoshani, J. & Tassy, P. (eds.) - Proboscidea. Evolution and palaeoecology of elephants and their relatives - pp. 203-213. Oxford University Press

Maglio, V.J└ 1973 - Ongin and Evolutin of the Elephantidae. Transactions American Philosophical Society, N.S. 53 (3), 149 pp.

Maschenko, E.N., 1993 - Mammoth heard structure from the Late Pleistocene locality ofSevsk. Transactions Zoological Institute of USSR 246: 41-59 (in Russian)

Roth, LV, & Shoshani, J└ 1988 - Dental identification and age determination in Elephas maximus. Journal Zoology 214:567-588

Sikes, S.K., 1971 - The Natural History of African Elephant - Weidenfeld and Nicolson, London, 1 68 pp.

Stenley, S.F., 1943 - Notes on age and sexual maturity, gestation period and growth of Indian Elephant, Llephos maximus - Proceedings Zoological Soc.iety London 1 13: 21 -27

Todd, N.E. & Roth, L.V., 1996 - Origin and radiation of the Elephantidae. - in: Shoshani, J. & Tassy, P. (eds.) - Proboscidea. Evolution and palaeoecology of elephants and their relatives - pp. 193-202. Oxford University Press

 In December of 1986, a nearly complete skeleton of a mature female southern mammoth, Mommuthus mendioncilis (Nesti, 1825), was discovered in the Borrego Badlands of northern Anza-Borrego Desert State Park (ABDSP) by G. Miller of Impenal Valley College Museum, El Centre California (Miller et oL, 1991 ). Magnetostratigraphy and tephrochronology date the specimen at approximately I. I MyBP, and place it in the lower Matuyama chron (Rerneika & Beske-Diehl 1996, McDaniel & Jefferson 1997). The morphology of various skeletal elements indicates that this specimen represents a very late evolutionary stage in the Mommuthus mendionolis chronocline. The site lies in the fluvial floodplain deposits of the Ocotillo Conglomerate (Dibblee 1954, Remeika & Pettinga 1991, Remeika 1992) in a geologically complex area south of the east end of the San Jacinto Fault Zone and is transected from southwest to northeast by the Valle Escondido anticline (Remeika pers. comm.1996).

The skeleton was deposited within a I m deep, ephemeral braided stream channel system typical of a distal bajada. Distal alluvial fan sheet flood deposits overtie the specimen. The positions of various skeletal elements in the bone bed indicate that the carcass had largely decomposed and the remains had been moved prior to bunal. A short duration transport within a fluvial system is inferred. The long axis orientation of various skeletal elements indicates a northeast to southwest direction of flow. The carcass had been heavily damaged by carnivores and scavengers. Osteopathologic evidence of long-tenn, progressively debilitating arthroses is present in the degenerative state of the tempero-mandibular articulation, making chewing painfully difficult. The specimen was about 55-60 AEY (Haynes 1991) at the time of death. Predation is a distinct possibility in an aging infirm adult. The remains of this late stage Mammuthus meridionalis co-occur with that of the imperial mammoth, M. imperator (LEIDY, 1858) from the Ocotillo Conglomerate. Significant differences in the morphology of the dentition, tusk, skull, mandible and scapula of the two taxa suggest that the steppe mammoth, M. armeniacus (FALCONER, 1857) rather than M. meridionalis was immediately ancestral to M. imperator. The skull and mandible ofM meridionalis, represented by
ABDSP(IVCM) V5126, is shorter antero-posteriorly and wider than that ofM imperator, represented by contemporaneous ABDSP(IVCM) V4214 and V4260. The index of curvature of the incisor of the former taxon is significantly less than that of the latter. It follows that M. mendionalis represents the end of one lineage, and that M. imperatorthe beginning of another lineage. TheMarnmuthus meridionalis-M. imperotor-M. columbi lineage ofMaglio ( 1973), now is seen as two separate dispersals from Eurasia to North America, the earliest occurring approximately 1.8 MyBP with the appearance ofM. meridionolis, and the second, M imperator at 1.2 NyBP. A third dispersal, that of the woolly mammoth, M. primigenius (BLUMENBACH, 1799), occurred in the late Pleistocene.

Dibblee, T.W., Jr., 1954 - Geology of the Impenal Valley region, California - California Division of Mines Bulletin 170 (2): 21-28

Haynes, G., 1991 - Mammoths, mastodons, and elephants - Cambridge University Press, England

McDaniel, G.E., Jr. & Jefferson, G.T., 1997 - A nearly complete skeleton of Mammuthus meridionalis from the Borrego Badlands, Anza-Borrego Desert State Park, California - in: Mojave Desert Quaternary Research Symposium, Abstracts of Proceedings - San Bernardino County Museum Association Quarterly 44 (1): 29-30

Miller, G.J., Remeika P., Parks, J.D., Stout, B└ & Waters, V.E., 1991 - A preliminary report on a half-a-million year old cut marks on mammoth bones from the Anza-Borrego Desert ln/ingtonian - Imperial Valley College Museum Society Occasional Paper 8: 1 -47

Remeika, P., 1992 - Preliminary report on the stratigraphy and vertebrate fauna of the middle Pleistocene Ocotillo formation, Borrego Badlands, Anza-Borrego Desert State Park, California - in: Mojave Desert Quaternary Research Symposium, Abstracts of Proceedings - San Bemardino County Museum Association Quarterly 39 (2): 25-26

Remeika, P., & Beske-Dlehl, S└ 1996 - Magnetostratigraphy of the Western Borrego Badlands, Anza-Borrego Desert State Park, California: implications for stratigraphic age control - in: Abbott, P.L└ & Seymour, D.C., (eds.) - Sturzstroms, and Detachment Faults, Anza-Borrego Desert State Park, California - pp. 209-220, South Coast Geological Society, Field Trip Guidebook

Remeika, P., & Pettinga, J.R└ 1991 - Stratigraphic revision and depositionai environments of the middle to late Pleistocene Ocotillo Conglomerate, Borrego Badlands, Anza-Borrego Desert State Park, California - Abstracts Symposium on the Value of the Desert - p. 13. Anza-Borrego Desert Foundation, Borrego Springs, California

 Remains of woolly mammoth Mammuthus primigenius (BLUMENBACH, 1799) from the remote Wrangel Island (East Siberian Sea) hit the news is 1993. Their geological age was Holocene (Vartanyan et al. 1993). The youngest C 14 datings reached ages of about 3700 yBP! The remains, mostly third molars (Mm3), were ascribed to dwarf rnamnnoths. Garutt et o\. ( 1993) considered these relatively small molars to be of a new subspecies, N\. primigenius vrangeliensis. The estimates of the withers height of these dwarf mammoths from Wrangel Island vary considerably. According to some researchers it was 1.5-2 m, others mentioned 1.8m, still others 2.0m (e.g. Lister & Bahn 1994, Agenbroad 1998). All estimates of the withers height are based on the size of the molars.

Postcranial skeletal elements of Holocene age have been found on Wrangel Island in addition to the mentioned molars. These postcranial elements show that we are dealing with characteristically Late Pleistocene woolly mammoths: an average withers height of between 2.5 and 3.0 m. Por example, a femur is known from Wrangel Island with a length of 0.98 m. This can be compared to the famous Berezowska mammoth, which was found at the beginning of the 20th century and that had a withers height of 2.65 m and a femur of 1.03 m. Other postcranial elements from Wrangel Island show that the mammoths had withers heights such as we also know from other localities in Eurasia. In general a strong decrease in body size can be seen in Pleistocene mammoths: /V\. meridionoiis from the Early Pleistocene is the largest (withers height up till 4.2 m), the Middle Pleistocene M. trogontherii reached heights of 3.5 -4.0m, and Late Pleistocene M. pnmigenius is the smallest with heights between 2.5-3.0m. A good example of relatively small animals are the mammoths from Sevsk (Russia). At this locality at least 33 individual animals were excavated of about 13,950 y old. Seven nearly complete skeletons belonged to animals ranging in age from new-born to very old. The largest skeleton, with awithers height of 2.4 m belonged to an adult bull (Lister & Bahn 1994).

The Southern Bight of the North Sea between England and the European continent is a rich locality for finding larger mammal remains. As early as 1986, Van Essen (1986) mentioned a remarkably small M3 ofM {)nmigenius (see Fig. I, roughly natural size!) Since, many relatively small molars, characterized by very thin lamellae are found in the North Sea. Similar finds are known from the Dutch continental area. All are of a Late Pleistocene age. The geologically youngest specimens from The Netherlands are from the IJssel Valley (province of Gelderland). A well-preserved skull of an adult female with mandible and complete dentition (second and third molars in function) was dated at Utrecht University to 22,1 60 +/- 260 yBP (UtC-4550). The oldest C14 dated specimens are skeletal remains from the bottom of the North Sea, belonging to adult animals with a withers height of less than 2.2 m (>45,000 yBP, University ofGroningen GrA-I 1640). Also in other European (e.g., England, Germany) and North Amencan (Fairbanks, Alaska) museum collections we have found small third molars ofA4 primigenius from Late Pleistocene deposits. We ascribe such remains of small adult Late Pleistocene mammoths to female animals. The post-cranial material of West-European Early Pleistocene M. meridionalis also contains both very large and very small specimens. There is apparently aconsiderable size range in mammoths. We prefer to consider this remarkable difference in sizes a result of sexual dimorphism. Adult female mammoths were considerably smaller than adult males (bulls).

The only true dwarf mammoth of Late-Pleistocene age isM exilis, known from the Channel Islands off the coast of California, U.S.A. They descend from the Columbus mammoth M. columbi, an abundant species in the Late Pleistocene of continental North America. This latter mammoth species reached awithers height of between 3.5 and 4.0 m. Only one fairly complete skeleton ofM exilis is known from Santa Rosa Island (Agenbroad 1998, Agenbroad et a!. 1999): its withers height was 1.6 m. M. exUis, with a withers height between 1.5 and 1 .8 m, is notably smaller than all supposedly dwarfed Late Pleistocene mammoths mentioned in the literature.

Agenbroad, L.D., 1998 - Pygmy (Dwarf) Mammoths of the Channel Islands of California - Mammoth Site of Hot Springs, SD Inc., Hot Springs, South Dakota, 27 pp.

Agenbroad, L.D., Morns, D. & Roth. L└ 1999 - Pygmy mammoths Mammuthus exilis from Channel Islands National Park, California (USA) - in: Haynes, G., Klimowicz, J. & Reumer, J.W.F. (eds.) - Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem - Deinsea 6: 89-102

Garutt, V.E., Avenanov, A.O. & Vartanyan, S.L, 1993 - On the systematic position of Holocene Dwarf Mammoths, Mommuthus {)nmigenius (Blumenbach, 1799) from Wrangel Island (North East Siberia) (in Russian) - Doklady Akademii Nauk332:799-801

Lister, A. & Bahn, P., 1994 - Mammoths - Macmillan, USA: 1 68 pp.

Van Essen, H., 1986 - Signalementvan een diminutieve M3 sup. van een wolharige mammoet - Cranium 3 (1): 6-7

Vartanyan, S.L, Garutt, V.E. & Sher, A.V., 1993 - Holocene dwarf mammoths from Wi-angel Island in the Siberian Arrtir - Nature 362: 337-340

 Mammuthus trogontheni rests were found in the localities of Srorodum, Abalack, and Tchemae on the territory of Western Siberia. The localities settle down on the right bank ofthe river Irtysh river upstream from the town of Tobolsk (58° N latitude). Remains of the following animals were discovered alongside with Mammuthus trogontheni: Desmano sp., Sorex sp., Citeilus sp., Ochotono sp└ Lepus sp., Sicista sp., Ciethnonomys ex. gr. glareolus, 0. sp.C ? ex. gr. rutilus), Proiagurus pannonicus, Lemmus cf. sibincus, Dicrostonyx mendionalis, Mimomys ex. gr. sovini, M. ex. gr. pusUius, M. ex. gr, intermed/us, Allophajomys pHocaenicus, Trogontherium cf. cuvieri, E.quus ex. gr. sonmeniensis, Bison sp└ Alces lotifrons, Carrielidae (Paracamelus ?) gen. This species structure of the fossil fauna is typical for the final stages of theEarly Pleistocene (approximately Cromerian I, I I ).

Real mammoths {Marnmuthus primigenius) appeared in central localities of Western Siberia at the beginning of the Middle Pleistocene (= Cromerian I I I, IY). In the localities ofVoronovo and Urtam on the left bank of the river Ob (56° N) the following taxa were found alongside with Mammuthus pnmigenius-. Ochotona sp., Lepus sp., Citeilus sp└ Cncetuius sp., Myospolax sp., A/tooicetus sp└ Lagurus sp., Eofagurus sp., Ciethnonomys sp., Microtus oeconomus, Stenocranius gregalis, Mimomys ex. gr. intermedius, Mimomys (Arvicola ?) sp└ {Lquus ex. gr. sanmeniensis-mosbachensis, Bison priscus, Alces cf iatifrons, Elasmoterium sp└ CoeiodontQ antiquit.avs, and Cervus elophus. Dunng Tobolsk Interglacial (Holsteinian ) the mammoths moved southward to 54° N (locality Tatarka). They inhabited the entire territory of Western Siberia. The assemblage with Mammuthus pnmigenius contains species such as Bison priscus cf iongicomis, Coeiodonta antiquit.atis, Cervus cf elaphus, Saiga cf rice), Ursus cf deningeri, Megoloceros sp. But the structure of mammoth faunas in southern regions of Western Sibena usually included supplementary steppe and desert species, and the fauna in the northern regions included tundra species. A morphological structure of fossil molars of the mammoth line is shown in Table I.

 Dwarfed elephants occurred in the Middle Pleistocene of the Mediterranean island of Sicily. First described by Busk (1867) on remains from Malta, CLiephas falconeri was considered for more than fifty years as the last step of a progressive size reduction process. This started with Elephos antiquus and continued through ?Bef>has (Palaeoloxodon) antiquus leonardii, ё. mnaidrensis and ?&ephos melitensis (according to different opinions by various authors) (Caloi et al. 1996 and references therein). Stratigraphic and geochemical data have now demonstrated that the smallest forms were older than the medium-sized elephants of the ё. rnnoidrensis group (Esu et al. 1986, Burgio & Cam 1988, Bada et 01.1991). The derivation of the so-called 'ё.' falconen from ё. antjquus is not sure. There is no conclusive osteological evidence forthis, and the very simplified molar morphology does not exclude a phylogenetic relationship with the Polaeoloxodon line. However, the morphology of the skull seems to be closer to the Loxodonto or Mammuthus lines. With respect to the postcranial bones, it is difficult to estimate to what degree the size reduction and the locomotion have influenced the morphological modifications ofthe limb bones compared to the ancestral species (Caloi & Palombo 1994, Palombo 1996). In spite of this, our preliminary investigations have shown that some morphological features of 'ё.' falconen from Spinagallo Cave in SE Sicily (Ambrosetti 1968) are closer to the Mammuthus line than to the Poloeloxodonto line. The analysis of carpal and metacarpal bones and ofthe tusk structure is still in progress and we need more data to resolve the problem of the origin ofthe Sicilian dwarf elephants.

Ambrosetti, P., 1968 - The Pleistocene dwarf elephants of Spinagallo (Siracusa, South-Eastern Sicily) - Geologica omana 7:277-398

Bada, J.L, Belluornini, G., Bonfiglio, L, Branca, M., Burgio, E. & Delitala, L.M., 1991 - Isoleucine epimerization ages of Quaternary mammals from Sicily - II Quaternario 4 (la): 49-54

Burgio, E. & Cam, M., 1988 - Sul ritrovamento di Elefanti fossili ad Alcarno (Trapani, Sicilia) - It NaturalistaSiciliano 12:87-97

Busk, G., 1 867 - Descnption ofthe remains of three extinct species of elephant, collected by Capt. Spratt, C.B.R.N., in the ossiferous cavern ofZebug, in the island of Malta - Transactions ofthe Zoological Society of London 6: 227- 306

Caloi, L & Palombo, M.R└ 1994 - Functional aspects and ecological implications in Pleistocene endemic herbivores of Mediterranean islands - Historical Biology 8: 151-172

Caloi, L, Kotsakis, T, Palombo, M.R. & Petronio, C., 1996 - The Pleistocene dwarf elephants of Mediterranean islands - in: Shoshani, J. & Tassy, P. (eds.) - The Proboscidea - pp. 234-239. Oxford University Press

Esu, D└ Kotsakis, T. & Birgio, E└ 1986-1 vertebrati e i molluschi continentali pleistocenici di Poggio Schinaldo (Palermo, Sicilia). Bolletino Societa Geologica Italiana 105: 233-241

Palombo, MK, 1996 - Lar-ge Pleistocene mammals ofthe Mediterranean islands - Vie et Milieu 46 (3/4): 365-374

 The La Polledrara de Cecanibbio site, together with the Castel di Guido site (Sala & Barbi 1996), is one of the richest deposits with Elephos (Paloeoloxodon) ontiquus in the Italian late Middle Pleistocene (- Early Aurelian Land Mammal Age sensu Gliozzi et al. 1997). The deposit, located on the highest terrace of the Northwestern area around Rome, at an altitude of about 83 m above sea level, was discovered in 1984 (Anzidei & Amoldus Huyzendveld 1992, Anzidei et of. 1989, in press). During numerous excavation campaigns, about 750 m2 of a paleosurface belonging to an ancient stream bed were uncovered. It contained a high concentration of large mammal bones (over 8,000) associated with a lithic and bone industry. The site is included in the terminal series of the pyroclastic deposits of the 'Sabatino' volcanic complex, up to now correlated with the Auretia Formation (Conato et at. 1980, De Rita et at. 1992) and correlated with OIS 9. Recent stratigraphical research seeims to indicate an erosive contact between the layers including the La Polledrara di Cecanibbio site and the deposits of
the Aurelia Formation. Therefore, La Polledrara might be older and might be coirelated with aterminal phase of OIS 10 (Anzidei et a!, in press).

Among the most common species (Bosprimigenius, Bephos antiquus, Cervus elaphus modem form, Equus cabolius, Conis aff. C iupus, Stephanorhinus sp.), Bephos bones are the most interesting, especially for the presence of two well preserved skulls. They offer a broader knowledge on the morphology of the Italian subspecies of Llephas antiquus, up till now often studied on incomplete or deformed skulls. All skeletal elements are represented: numerous tusks (over 30), mandibles, isolated molarteeth and postcranial bones (some ofthem in anatomical connection), belonging to at least fifteen individuals. The studies, presently in progress, may contribute to the knowledge of the morphological and biometrical variability of the Bephas antiquus populations of the late Middle Pleistocene, and test the variability of some characters that are considered useful for gender determination.

Anzidei, A.P., Angelelli, F└ Amoldus Huyzendveld, A., Catoi, L, Palombo, M.R. & Segre, A.G└ 1989 - Le gisement pleistocenique de La Polledrara di Cecanibbio (Rome, Italic): la faune - L'Anthropologie 93 (3): 749-782

Anzidei, A.P. & Amoldus Huyzendveld, A., 1992 - The Middle Pleistocene site of La Poltedrara de Cecanibbio (Rome, Italy) - in: Papers Fourth Conference Italian Archaeology - New developments in Italian Archaeology I: 141-153

Anzidei, A.P., Amoldus Huyzendveld, A., Caloi, L, Palombo, M.R & Lemorini, C., in press - Two Middle Pleistocene sites near Rome (Italy): La Polledrara di Cecanibbio and Rebibbia-Casal De'Pazzi - Atti Coil. 'The Palaeolithic occupation of Europe. The role of Early Humans in the accumulation of European Lower and Middle Palaeolithic bone assemblages, Neuwied, 18-21 May, 1995

Conato, V., Esu, D., Malatesta, A. & Zarlenga, F└ 1980 - New data on the Pleistocene of Rome - Quatemana 22: 131-176

De Rita, D., Milli, S., Rosa, C. & Zarlenga, F└ 1992 - Un'ipotesi di correlazionetra la sedimentazione lungo la costa tirrenica della campagna romana e l'attivitavulcanica dei Colli Albani - St. Geol. Camerti, vol. spec. CROP I I (199 I/I 992): 343-349

Gliozzi, E└ Abazzi, L└ Ambrosetti, P., Argenti, P., Azzaroli, A., Caloi, L, Capasso Barbato, L, Di Stefano, G., Esu, D., Ficarelli, G., Girotti, 0└ Kotsakis, T., Masini, F└ Mazza, P., Mezzabotta, C└ Palombo, M.R, Petronio, C└ Rook, L., Sala, B└ Zanalda, E. & Torre, D└ 1997 - Biochronology of selected mammals, moltusks, ostracods from the Middle Pliocene to the Late Pleistocene in Italy. The state of the art - F^iv. Italiana Paleont. Stratigr. 103 (3): 369-388

Sala, B. & Barbi, G└ 1996 - Descrizione deila fauna - in: Radmill, A.M. & Boshian, G. (eds.) - Gli scavi a Castel di Guido - Istituto Italiano Preistoria Protostoria Firenze: 55-90

 The sexual detemnination of fossil elephant skeletons is generally considered an important contribution to understanding their social structure and it can be also useful in taphonomical studies. There are different main categories of potential information about the sexual dimorphism in the proboscidean skeleton: (1) preserved genitalia, (2) skeleton size and robusticity, (3) skull and tusk morphology, (4) pelvic morphology. Many studies were undertaken to determine the efficient dimorphic characters from a biometrical and morphological point of view. The greater part of study was been done on hA. pnmigenius (Averianov 1996) or on other remains of the genus Mammuthus (Lister 1996), Contrary to this, sexually dimorphic characters of ё. (Polaeoxodon) antiquus were not much known yet (Dubrovo & Jakuboswski 1988└Palombo 1986). To test these sexually dimorphic characters, an almost complete skeleton ofё. antiquus, which was found in the lacustrine deposit 'Grotte Santo Stefano' near Viterbo (Central Italy), and which is late Middle Pleistocene in age, was examinated. In addition to dimorphic
characters shown by the skull, the tusk and by pelvic morphology, there is some information about the differences between male and female ofё. (Polaeoloxodon) antiquus to be found in morphology and biometry of the molars, atlas, epistropheus and carpal and metacarpal bones.

Averianov, A.O., 1996 - Sexual dimorphism in the mammoth skull, teeth, and long bones - in: Shoshani, J. & Tassy, P. (eds.) - The Proboscidea - pp. 260-267. Oxford University Press

Dubrovo, I.A. & Jakubowski, G., 1988 - The carpus morphology of the forest elephant (Pataetoxodon) and its significance fortaxonomy - Prace Muzeum Ziemi 40: 65-95

Lister, A.M., 1996 - Sexual dimorphism in the mammoth pelvis: an aid to gender determination, - in: Shoshani, J. & Tassy, P. (eds.) - The Proboscidea - pp. 254-259. Oxford University Press

Palombo, M.K, 1986- Observations sur Qephos antiquus Falconer & Cautley du Pleistocene moyen d'ltalie: essai devaluation des caracteres dentaires. Geologica Romana 23 ( 1984): 99-1 10

 In order to answer the questions about the setting of the faunal assemblages with mammoth remains, pertinent taphonomic criteria are looked for and proposed here in a methodological way. All factors, which can lead to an accumulation of mammoth bones, should be taken into account from the beginning of the study. First, the biological or non-biological hypotheses should be tested. Among the predator agents, the anthropic factors should be distinguished from the carnivore ones. If man is responsible for the accumulation, differences should appear between hunting and scavenging activities. The collecting strategy might have been done quickly (fast access to the carcass), or slowly, depending on the people's needs (food or other utilization), To help to organize this methodology, references are taken throughout European Upper Paleolithic settlements, East by West, and also in Middle and Lower Paleolithic, and even Neogene, Proboscidean open air sites.

 Sedentary birds are a group of species that can be important in reconstructing the Pleistocene epoch environmental conditions, based on fossil remains in the Pleistocene epoch. Numerous remains of birds (about 10,000 bones) were collected in two archeological sites - Medvezhya Cave (approx. 62° N 58.5° E) and Grotto Bolshoi Glukhoi (approx. 58.5° N 58° E) in the western foothills of the Northern and Middle Urals. Both caves are located within the boreal zone about 400 krn apart. Late Pleistocene mammal bones from Medvezhya Cave (MC) were radiocarbon dated from 18,700 ╠ 180 yBP to 1 6,130 ╠ 150 yBP (brown clay, honzon 5B), 13,260 ╠ 230 yBP to 12,230 ╠ 100 yBP (brown clay, horizon 5A), and 12,670 ╠ 90 yBP (gray clay, horizon 3) (Sinitsin & Praslov 1997). The upper part of honzon IX or cultural layer 6 of Grotto Bolshoi Glukhoi (GBG) had a carbon date of > 33,900 yBP (Guslitser & Pavlov 1993). Bones from the same horizon (IX) were radiocarbon dated at 38,200 ╠ 900 yBP (GIN-8404). The horizons of gray (IV), humus (V), and brown (VI and VII) clays of GBG have not been radiocarbon dated at this time. Taking into account the deposits sequences of lithology and fauna) compositions I believe that mentioned horizons of GBG may correspond to the gray, humus and brown clays of MC.

Recovered megafauna consisted of reindeer (Rangifer tarandus L), arctic fox (Atopex logopus L.), Pleistocene hare (Lepus tonaitJCUS GUR.EEV, 1964), cave bear (Spefaearctos speleus ROSEN. AND HEINR. 1794), Ural's horse (E.quus uralensis KUZMINA, 1975), musk-ox {Ovibos pationtiS SMITH, 1827), bison (Bison priscus BOJANUS, 1827), saiga (Saiga borealis TSCHERSKY.1876) and mammoth (Mornmuthus primigenius BLUMENBACH, 1799) in horizons 5 and 3 of MC, and horizons VII and VI in GBG (Kuzmina 1971, Kuzmina & Sablin, 1991 ). The remains of sedentary birds from MC and GBG include 31 species, the majority of which are grouse and birds of prey. These and passerine bird species are ecologically tied to each other as 'predator and prey' (L togopus + L mutus≈N. scondiaca + F. g/rfalcon + F. peregrinus + A. gentilis', Capercaitlie≈F. cherrug + A. gentilis', L. tetrix + 6. bonasia + P. perdix≈A. gentilis', Passeriformes≈Strix oluco). The eagle owl could also have hunted owls, grouse and passerine species. Study of the complete microfossil assemblages from the caves showed that avian and rodent faunal remains accumulated from regurgitated pellets of eagle owl, Bubo bubo (L), tawny owl, Strix aiuco L, and in smaller part by middle-sized carnivorous predators.

Avifauna) assemblages from the Wurm deposits of the caves indicate predominantly open landscapes (tundra-steppe) (65-92% in MC and 34-81 % in GBG), with the presence of forest vegetation in river valleys on the western slope of the north and middle Ural Mountains. Forest formations could have comprised 8-35% of the landscape around MC and 19-66% surrounding GBG. The assemblages include many species of forest dwellers (hazel grouse, Ural's owl, hawk owl, Tengmairn's owl, three-toed woodpecker, crossbill and two-barred crossbill, nutcracker) that prefer spruce, fir or larch, with mature or thin forest stands. These data agree more with that of the rodent fauna (Guslitser et ol. 1990, Guslitser & Pavlov 1988), than from pollen data (Guslitser & Kanivets 1965). Most of the skeletal remains of the genus Lagopus species belong to the willow grouse, and one subspecies of tundra ptarmigan. These accounts for about 95% of the species composition in the Late Wurm deposits ofMC and GBG. Tundra subspecies of ptarmigan (Lagopus mutus ssp.) differs from modem tundra ptarmigan subspecies by a short and thick-set tarsometatarsus. It comprises approximately 50% of the bone assemblage of genus
Lagopus in MC and GBG. The relatively large amount of bones of isolated subspecies of mountain ptarmigan (which today inhabits alpine arctic tundra, at elevations of 1200 -1 600 m above sea level) shows that, in the Late Pleniglacial, its range comprised the eastern belt of the western North Urals, with an elevation of about 300 - 400 m above sea level, about 50 krn west of the current range there and 150 km south of the current range in the Middle Urals. Predominantly open landscapes, including tundra, tundra-steppe and steppe, allowed willow grouse and the tundra ptarmigan subspecies to penetrate far to the south, especially during winter. The extreme rarity of young individuals of both species of the genus Lagopus (less than 0.5% of the bones) in the fossil assemblages provides evidence that these birds were eaten during winter or winter/spring periods. The modern seasonal foraging pattern of the willow grouse and ptarmigan, making seasonal movements with reindeer herds, suggests that these birds and reindeer probably made seasonal movements together in the late Pleniglacia). In the Late
Pleniglacial the tundra species of ptarmigans probably followed the seasonal north-south movements of reindeer, and possibly the Ural's horse and bison, which made foraging easier for the birds in snow-covered conditions. The presence of the migrating herds may explain the abundance of these birds on the western slope of the middle Urals during the Late Pleistocene. Predators, such as snowy owl and gyrfalcon followed the seasonal movements of grouse, as -they do today. The presence of tundra species of ptarmigan in GBG is the most southern record for that bird in the Pleistocene in European Russia.

Guslitser, B.N└ 1965 - Caves of Pechora's Urals - Nauka, Moscow-Leningrad, 134 pp. (in Russian)

Guslitser, B.N. & Pavlov, P.Yu., 1988 - Upper Paleolithic Site in Medvezhya cave (new data) - Materialy po arkheologii Evropeiskogo Severo-Vostoka. Vyp. 1 1: 5-18 (in Russian)

Guslitser, B,l└ Pavlov, P.Yu. & Panukova, N.N., 1990 - Application ofpaleomicrotheriological methods in study of Paleolithic site in Medvezhya cave - Kratkie soobshcheniya Institute Arkheologii 202: 1 10-1 14 (in Russian)

Kuzmina, I.E., 1971 - Formation ofTheriofauna of the North Urals in the late Anthropogene - Trudy Zoologicheskogo institutaAN SSSR49: 44-122 (in Russian)

Kuzmina, I.E. & Sablin, M.V., 1991 - The Remains of Mammals from Grotto Bolshoi Glukhoi in the Middle Urals - in: Abstracts of International Symposium 'Problems of the Historic and Cultural Environment of the Arctic'. Syktyvkar,
pp. 77-78

 The species of the family Gomphotheriidae from South America range from the Middle Pleistocene (Ensenadan Land-mammal Age) to the latest Pleistocene, Lujanian Land-mammal Age (Alberdi & Prado 1995). These groups were descendants of a gomphothere stock from North America and arrived to South America during the 'Great Amencan Biotic Interchange' (Webb 1985), Based on vegetational distribution maps of South America during the glacial and interglacial phases, and the distribution of South American Gomphotheres, it appears that Cuvieronius inhabited an arid landscape, whereas Stegomastodon seems to have predominated in areas identified as savannah (Webb 1991 ). Cuvieronius hyodon seems to have been adapted to atemperate-cold climate, since in the intertropical zones it has been found only at the highest levels. On the contrary, in Chile it expanded to the littoral zone, that surely offered more similar living conditions, in terms of temperature, than the Andes corridor. This species would feed mainly on hard grasses, leaves, and seeds from bushy plants. Species of Stegomastodon would be better adapted to warm or temperate climatic conditions, because their most austral distribution does not surpass the 37° S parallel in the Buenos Aires province (Tonni 1987). The frequency of Stegomastodon platensis diminishes in the Pampean Region by the latest Pleistocene, when environmental conditions became colder and drier. In lower latitudes, where Stegornostodon is more frequent, it would occupy savannahs orxerophytic pastures.

Simpson & Paula Couto (1957) considered that all South American forms must be included in one subfamily: the Anancinae, as there are only few and slight differences among them. We agree with this opinion and recognise in South America only two genera: Cuvieronius with only one species: Cuvieronius hyodon, and Stegornostodon with two species: Stegomastodon waringi and S. plotensis (Alberdi & Prado 1995). C. hyodon is geographically restricted to the Andean Region in Ecuador, Peru, Bolivia, Chile, and Northwest Argentina (Hoffstetteri 952, Casamiquela et ol. 1996). S. waringi was recorded in the Santa Elena peninsula in Ecuador (Hoffstetter 1952, Ficcarelli et 0/., 1996), and in the tropical zone of Colombia (Coireal Urrego 1981 ) and Brazil. S. platensis was recorded in the middle to latest Pleistocene in Argentina, especially the Pampean Region, and also dunngthe late Pleistocene of Uruguay (Mones & Francis 1973) and Paraguay (Cabrera 1929, Simpson & Paula Couto 1957). During the Pleistocene in South America two savannah corridors would have developed, one called 'the Andes route', or 'the height corridor' and another one the so-called 'East route' or 'plain corridor' (Webb 1978, 1985). Both corridors have special characteristics and very probably they constituted the main dispersion route for different mammal groups.
The Andes route was a more direct North-South connection and provided greater environmental homogeneity of non-woodland habitats in comparison with the East route. On the other hand, the low temperatures associated with the Andes route could have constituted a barrier for certain groups. These two corridors have conditioned the paleobiogeographic history of most North American mammals in South America. In fact, different models can be postulated for different groups depending on their capacity to produce distinct adaptive types along the duration of their dispersion process. In the case of the South American gomphotheres, the small Cuvieronius utilized the Andes corridor, whereas the large Stegomostodon dispersed through the East route (Figure I ).

We reconstruct the diets of Cuvieronius h/odon, Stegomastodon ptotensis and S. wanngi through isotopic analysis. Cuvieronius from Tarija indicates that they were almost exclusively mixed feeders. S. piatensis from the Middle Pleistocene of Argentina shows mixed feeding to browsing adaptations, and the same species from the Late Pleistocene in Argentina indicates atrend from mixed feeding to browsing adaptations. S. wanngi from the Peninsula of Santa Helena shows atrend of mixed feeding to grazing adaptations (Sanchez et o\. in press).

Cabrera, A., 1929 - Revista del Museo de La Plata 31: 61 -144

Casamiquela, R.M. et al., 1996 - in: Shoshani, J. & Tassy, P. (eds.) - The Proboscidea - Oxford Universrty Press

Correal Urrego, G., 1981 -Fund. Inv. Arqueol. Nac., Bogota 12: I -148

Ficcarelli, G. et aL, 1993- Geobios 12 (2): 231 -240

Hoffstetter, R└ 1952 - Memoires Societe Geologique de France 66: 1 -391

Mones, A. & Francis, J.C└ 1973 - Comunicaciones Paleontologicas del Museo de Historia Natural de Montevideo (4): 39-97

Sanchez, B. et a!., in press - International Congress, 19-22 Mayo 1999, La Paz, Bolivia

Simpson, G.G. & Paula Couto, C└ 1957 - Bulletin of the American Museum of Natural History 1 12 (2): 1 25-190

Tonni, E└ 1987 - Ameghiniana 24 (3-4): 323-324

Webb, S.D., 1978 - Annales Review Ecology Systematic 9: 393-426

Webb, S.D., 1985 - in: Stehii, F.G. & Webb, S.D. (eds.) - The Great American Biotic Interchange - pp. 357-386 - Plenum Press, New York and London

Webb, S.D└ 1991 - Paleobiology 17(3): 266-280