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Notes on skulls of Pleistocene Saigaof Northern EurasiaG. Baryshnikov a & A. Tikhonov aa Zoological Institute of Russian Academy of Sciences ,Universitetskaya emb.l., St Petersburg, 199034, RussiaPublished online: 10 Jan 2009.

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Historical Biology, 1994, Vol. 8, pp. 209-234 © 1994 Harwood Academic Publishers GmbHReprints available directly from the publisher Printed in MalaysiaPhotocopying available by license only

NOTES ON SKULLS OF PLEISTOCENE SAIGA OFNORTHERN EURASIA

G. BARYSHNIKOV and A. TIKHONOV

Zoological Institute of Russian Academy of Sciences, Universitetskaya emb.l. StPetersburg 199034, Russia

(Received November 2, 1992)

Eighteen fossil skulls of male Saiga from Northern Eurasia and 33 recent skulls from Kalmykiaand Kazakhstan have been studied. Saiga from both the Khazarian Fauna of the Volga andMammoth Fauna of Europe and Siberia are referred to Saiga horealis Tschersky, 1876. Duringthe Pleistocene, 5. borealis distribution extended from England in the west to Alaska in the eastand is characterized by an elongated neurocranium, small frontal angle of the temporal bonefrom the plane of the frontal, and long nasal bones.

S. borealis was a typical representative of the "mammoth biome" in the Pleistocene periglacialsteppes and cryogenic savannahs. Two subspecies are recognized: S. borealis borealis Tschersky(Eastern Sibera and Alaska); and S.b. prisca Nehring, 1891 (Europe, Urals and Western Siberia).At the end of the Pleistocene, when the mammoth disappeared, the range of S. borealis wasreduced. Today they live only in West Mongolia (S. borealis mongolica Bannikov, 1946).S. tatarica tatarica was widely distributed in the other territories of the steppe and semidesertzones of Eurasia. The arid landscapes of Transcaucasia and Kazakhstan were inhabited by Saigawith thinner legs and shorter nasal bones, such as S. tatarica binagadensis Alekperova, 1953,from the middle Pleistocene of Azerbaijan (Bynagady). Fossil skulls from the Ural River thatare large, but with a short neurocranium are identified as Saiga sp. cf. S. tatarica Linnaeus, 1766.

KEY WORDS: Saiga, Pleistocene, Eurasia, Saiga borealis, morphometric characters.

INTRODUCTION

Recent Saiga — Saiga tatarica Linnaeus, 1766 — inhabit steppes of the Lower Volga,Kazakhstan and Mongolia. Herds of this species migrate long distances and individualsare adapted to running in rugged country but not in deep snow cover. These biologicalfeatures of the Saiga make it a good indicator of steppe conditions. Therefore,Quaternary remains of Saiga beyond the recent geographic range of the genusthroughout Northern Eurasia from England in the west up to Chukotka in the east, toAlaska and the Northwest Territory, Canada, are of interest to researchers.

In North America, the Pleistocene Saiga has been described several times underdifferent names: Saiga borealis Tschersky, 1876 (Yakutia); S. prisca Nehring, 1891(Moravia); S. ricei Frick, 1937 (Alaska); and S. binagadensis Alekperova, 1953(Transcaucasus). Specimens refered to these taxa differ from the recent form in thestructure of skull and dentition. The relationships among these extinct groups and theirgeographic distributions during the Pleistocene remain to be clarified. To elucidatethese questions we have studied the available fossil and recent skulls from scientificcollections is Leningrad, Moscow and Krakow.

209

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210 G. BARYSHNIKOV AND A. TIKHONOV

MATERIAL AND METHODS

We studied collections of twelve adult male fossil Saiga reposited in the ZoologicalInstitute, Academy of Sciences, Leningrad, Russia (ZIAS; Volga (1), Ural River (2),Western Siberia (3), Middle and Eastern Siberia (5); others unknown). Two specimenswere also examined from the Paleontological Institute, Academy of Sciences in Moscow,Russia (Volga and Vilui River), and one from the Institute of Systematics andExperimental Zoology, Polish Academy of Sciences, Krakow (Slupyanca Cave). A castof a Saiga skull with horncores from Belgium (Leopold Canal) was also available fromthe Institut Royal des Sciences Naturelles de Belgique (N 16-1021). We also used dataon teeth and the postcranial skeleton from Palaeolithic sites of the Crimea, Urals, andfrom asphalt sands of the Apsheron Peninsula (collection of ZIAS). Modern comparativematerial included 33 recent skulls of adult males from Kalmykia and Kazakhstanreposited in the ZIAS and the Zoological Museum, Moscow University. We also useddata on fossil Saiga from Czersky (1876), Alekperova (1955), Vereshchagin (1959),Sher (1967), Alekseeva (1980), Harrington (1981), Lazarev and Tomskaya (1987), andKahlke (1990).

Since the fossil material is fragmentary and is represented by pieces of skulls withhorncores the measurements listed below were most useful (Figure 1). The T-test wasused to determine the significance of morphometric changes : p < 0.001 (+++); p < 0.01(++); and p < 0.05 (+).

1. Upper neurocranium length: akrokranion - supraorbitale.2. Length behind horns: akrokranion - the hind surface of horncore bases.3. Cranial length: akrokranion - bregma.4. Diameter of the horncore base.5. Greatest breadth across the orbits: ectorbitale — ectorbitale.6. Width between foramen supraorbitale.7. Width between horns.8. Latero-medial diameter of the horncore base.9. Least breadth of the parietal.

10. Greatest mastoid breadth: otion - otion.11. Greatest breadth of the occipital condyles.12. Height of occiput: akrokranion - basion.13. Frontal angle (in degrees).

MORPHOMETRIC CHARACTERISTICS OF THE SKULL OF SAIGA BOREALIS

Czersky (1876, p. 150) described from the Vilui River, Yakutia, an extinct species ofAntilope (Saiga) borealis and assumed that the structural features of its skull indicatedsharp differences in climate between the Pleistocene distribution and the recent.. Theholotype of this species has been lost. To this new species, Czersky included theseparate horncore from the Nizneudinskava Cave, Sayani (Czersky, 1876, p. 149). Later,Frick (1937, p. 546) described Saiga ricei from Alaska (RAM 30495 from Lilian Creek,in the region of Fairbanks). Examining the fossil material from Eurasia, Vereshchagin(1959, p. 444) noted morphological differences between the East Siberian Saiga andthe recent species, in particular the somewhat elongated neurocranium of the skull, theweak spread of the horns, and straightness of the horncores.

Sher (1967), who studied serial fossil material, concluded that Siberian and AlaskanSaiga were very similar and placed them in the same species - S. ricei. He thought

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PLEISTOCENE SAIGA IN EURASIA 211

Figure 1 Diagram of Saiga's skull measurements.Designation of measurements in text.

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212 G. BARYSHNIKOV AND A. TKHONOV

that S. ricei differed from the holotype of 5. borealis by a smaller orbital width(128 mm and 133.8 mm, respectively). However, this character however is variable,since the margins of the orbits frequently are broken. Orbital width is thus not diagnosticfor the specimens we studied and does not separate S. borealis from 5. tatarica.Proceeding from the assumption that all skulls of the Pleistocene Saiga from the basinof the Lena River (Vilui River included) are similar, we can retain the name S. borealisfor this form as the species having priority (Kahlke, 1975 p. 138) based on the followinginformation.

Studying the fossil Saiga from Siberia, Sher (1967: 106) noted the followingcharacteristic features: skull relatively narrow across orbits, horncores strongly bentbackwards, and neurocranium elongated. The latter two characters confirm thedifferences between S. borealis and S. tatarica. The minimal values characterizing thelength of the hind part of skull in the extinct species (measurments 1-3) on specimensfrom Siberia exceeds the mean indices in the recent S. tatarica (Table 1). These formsdiffer in the index of relative skull width at the base of the horncores in relation toneurocranial length, which is much smaller in S. borealis. The angle of the frontal bonesfrom the plane of temporal in S. borealis is also significantly smaller. As a rule thisangle does not exceed 32 degrees. In S. tatarica it is 33 degrees on the average.

The following parameters should be added to the above ones: the occiput in the fossilSaiga is broader (in relation to its height) and the nasal bones are longer. The nasalsare preserved completely on only one specimen (Minusinsk, ZIAS N 15097, Figure 2)and are 67.4 mm in length. In recent S. tatarica, the nasals range from 50.8 mm to63.2 mm (x = 57.6, n = 32), and in 5. mongolica 48.3 mm to 56.8 mm (x = 52.1, n = 3).

A \W4

=1 cm.

Figure 2 Saiga borealis, from Aidorach River near Minusinsk ZIAS N 15097. A - front view, B - sideview.

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Table 1 The size of skulls of Saiga borealis from Siberia and Alaska

Measurements

(mm)

1."2.3.4.5.6.7.8.9.10.11.12.13.

River TuraZIASb> N°3383

32.8,32.4137.253.2

110.129.7,31.6

Western Siberia

River Uschaikanear TomskZIAS N° 32727

94.560.5

120.130.3,29.2

13353.7

103.226.3,28.2

77.591.752.345.628

KuzbassZIASN°32750

97.865.3

129.131.2,29.6

13648.0

102.330.8,29.7

74.592.852.348.932

River Aidorachnear MinusinskZIAS N° 15097

96.065.9

123.430.0,29.9

13351.3

105.632.1,31.4

72.891.253.048.134

Middle Siberia

The Yeniseinear KrasnoyarskZIAS N°14585

97.466.1

121.331.7

12951

102.429.2708751.545.0

River Vilui(holotype)(Chersky, 1876)

96.363.0

33.0133.8

97.829.0

Yakutia

River ViluiPIASC)

N°750-146451

99.365.9

115.526.5

119.7 •49.497.826.765.183.146.449.7

rw

oowin5Q>

fri

s

a) - description of measurements on page 000b) - Z I A S - Zoological Institute Academy of Sciences, Leningradc) — PIAS — Palaeontological Institute Academy of Sciences, Moscow

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Table 1 (cont.)

Measurements(mm)

1.2.3.4.5.6.7.8.9.10.11.12.13.

River Vilui(Sher, 1967)

99.066.0

33.6. 122.6

102.932.2

22

River OlenekZIAS N°7076

121.729.2,28.8

-13152.899.7

28.9,27.573.9

Yakutia

River OlenekZIAS N°7077

96.767.2

117.230.2,28.7

-13149.698.2

27.9,26.772.286.545.945.224

The delta ofLena ZIASN° 13636

89.164.2

28.8,28.7-128

40.590.6

26.7,25.669.885.645.748.522

River Kolyma(Sher, 1967)

94.765.0

28.3113 def.

94.825.8

24

Alaska

Lilian Creeknear Livengood(Sher, 1967)

9765

30.0 (25.2)*116 def.

93 (94.0)28.0 (24.0)

(68.0)(82.0)(44.0)(47.0)31

Gold Hillnear Fairbanks(Harington, 1981)

29.6

104.427.669.284.848.049.3

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PLEISTOCENE SAIGA IN EURASIA 215

The strong development of the posterior part of the skull and a weak bend betweenthe frontal and temporal planes can be regarded as archaic characters which were moretypical of extinct Siberian Saiga when compared to the recent species (Sher, 1967).

An examination of the fossil materials shows that Saiga from Eastern Siberia differfrom Saiga of West Siberia by a relatively more elongated hind part: the index of hindlength (character 7) to width between horns (character 1) is from 98.5 to 101.7 (x = 101.2,n = 6). This index is much lower in the sample from Western and Middle Siberia(range is 104.6 to 110.0, x = 107.2, n = 4). Fossil Saiga from Alaska resemble that fromEastern Siberia (ratio is 95.9 for Lilian Creek specimens). Saiga from Yakutia alsopossess a lower occiput. This data allows us to assume that there are differencesbetween the West Siberian and East Siberian fossil Saiga at the subspecies level.

The second form, S. borealis borealis, looks slightly more archaic because it hasstronger differences from recent S. tatarica. One skull from the Lena River (ZIAS N13636, Figure 3) preserves all the main characters of 5. borealis, but is smaller in size(neurocranial length = 89.1 mm). However, it is still larger than skulls of the smallestrecent Mongolian Saiga S.b. mongolica Bannikov, 1946, in which the neurocraniallength ranges from 83 mm to 85.6 mm (x = 84.4, n = 3). It is interesting that the frontal-parietal index of S. mongolica is very small (range is 84.7 to 90.9, x = 87.7, n = 3),but is closer to S. borealis than to S. tatarica. The fossil skull from the Lena Riverhas fused sutures and belongs to an adult male of S. borealis, which became smallerat the end of the Pleistocene. This results from the condition of isolation, noted byVereshchagin (1959) and Harrington (1981). Reduction of size at the end of thePleistocene also characterizes horses, bison, mountain sheep, and probably mammoths.

B

Figure 3 Saiga'borealis, from the delta of Lena River. ZIAS N 13636. A - front view, B - side view.

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216 G. BARYSHN1KOV AND A. TIKHONOV

EUROPEAN SAIGA BOREALIS

Pleistocene specimens of Saiga found in Europe from the Urals in the east to Englandin the west and southwards up to Crimean Peninsula are usually refered to S. tatarica.However, the taxonomic position of fossil Saiga from Europe should be defined moreprecisely.

Volga River. - Two skulls of Saiga from the Volga River (Middle Pleistocene,Khazarian theriocomplex, Figure 4A) are of rather large size, with a long neurocranium(Table 2), a large frontal-parietal length index (103.0; 98.9), and also an angle of dipof the frontal plane in relation to that of the parietal bone that correspond to indicesof S. borealis (Figure 6A). The Volga form is characterized by straight, relatively shortand weakly spread, horncores (Vereshchagin, .1959). Perhaps this is a peculiar subspeciesof S. borealis.

Bynagady, Baku. - Saiga binagadensis was described from the asphalt deposits ofBynagady near Baku (Alekperova, 1953, p. 70) which is of late Middle Pleistocene(Riss/Moscow) or last interglacial (Riss-Wurm/Mikulino) age. Vereshchagin (1959,p. 442) assumed that S. binagadensis Alekperova, 1953, from the eastern Transcaucasiais closely related to the Khazarian Saiga. This species is characterized by little spreadbetween horns, but taller horncores than the Khazarian specimens with a more markedbend backwards in the sagittal plane. Thy Bynagady male skull measures 129 mm inwidth between the orbits, 101 mm between the bases of horncores on the outside, and51 mm between the supraorbital apertures. The nasal bones are short and narrow, witha length of 48 mm (Alekperova, 1955, p. 20). Lower jaws and limb bones ofS. binagadensis (ZIAS N 24406) are much smaller than in S. borealis.

In skull proportions the Bynagady Saiga is close to S. tatarica, and is refered hereto the subspecies, S. tatarica binagadensis. This systematic view suggests that thedivergence of the phylogenetic lines of S. tatarica (south steppe and semideserts) andS. borealis (north steppe) began in the Middle Pleistocene.

w••itijtl

A B

Figure 4 The skulls of saiga, front view.A - Saiga borealis ssp., Luchka near Saratov, Volga River, Khosar fauna. ZIAS N 1084.B - Saiga cf. tatarica, Ural River. ZIAS N 24201-10.

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PLEISTOCENE SAIGA IN EURASIA 217

Table 2 The size of fossil saiga skulls from Europe

Measurements

(mm)

1.2.3.4.5.6.7.8.9.10.11.12.13.

Middle

River

LuchkaZIASa>

N°1084

97.566.5

120.728.3

127.250.496.4

27.2,24.674.484.948.748.427

S. borealis

pleistocene

Volga

TungusPIASb)

N°131-277

101.066.4

126.831.5_

53.7104.027.075.690.750.651.2-

Late

Belgium

castZIAS Nc

97.886.2

121.330.5,29.5

125.747.1

100.427.6,27.0

75.490.151.448.429

S. cf. borealis

pleistocene

Poland

Slupianca'5 IASEC)

Z-206/38

93.261.8

118.731.3,31.0

12647.3

100.229.7,29.6

71.086.748.147.2 .26

S. cf. tatarica

The end of pleistocene

ZIASN°24201-10

92.263.7

123.829.7,29.4

13654.6

103.327.5,28.5

74.685.145.249.034

River Ural

ZIASN°24201-8

95.762.5

122.330.3,30.0

134.151.5

107.029.2,28.3

72.386.250.449.8-

ZIASN°24201-29

94.364.2

123.233.4,33.2

133.250.3

103.331.3,31.5

77.288.648.747.8

-

a) - ZIAS - Zoological Institute Academy of Sciencesb) - PIAS - Palaeontological Institute Academy of Sciencesc) - IASE - Institute of Animal Systematics and Evolution, Krakow, Poland

I . . J cm

B

/ /y

Figure 6 The skulls of saiga, side view. A - Saiga borealis ssp., Luchka near Saratov, Volga River.B - Saiga cf. tatarica, Ural River.'

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218 G. BARYSHNKOV AND A. TIKHONOV

Moravia, Czechoslovakia. — Saiga prisca was described by A. Nehring (1891, p. 131)from the Late Pleistocene deposits of Sipka-Hohle in Moravia, Czechoslovakia. Theprimary diagnostic difference of this species is the presence of an alveolus in front ofP/3 on the jaw of the adult individual. As is known, adults of recent Saiga sometimespreserve the dP/2. The size of the alveolus in recent Saiga fully coincides with the sizeof the alveolus in front of P/3 in Moravian specimen. It is clear that this tooth anomalycannot be used as a taxonomic character.

Among 109 adult skulls of S. tatarica, dP/2 is present in front of P/3 on eightspecimens and in three others a single rooted needle-like tooth (reduced dP2?) ispresent. The one skull of S. mongolica has a P/2 (Baryshnikov et al., 1990, p. 11). Lengthof the tooth row in the holotype of S. prisca equals that of Saiga from Pleistocenedeposits of the Crimea, Urals and southern regions of Western Siberia. All are largerin the length of the tooth row than in S. tatarica. The length of P/3 to M/3 (whenM/3 is strongly worn) is 74 mm in S. prisca. This length is 71.2 mm in the specimenfrom grotto Kiik-Koba in Crimea, 75.7 mm in the specimen from Stolbovoi Grotto inthe Urals, and 75 mm in the material from Krasnyi Yar on the Ob River. In the recentspecies the length of P/3-M/3 is less than 71 mm. Thus the form from the Pleistoceneof Moravia resembles the northern Saiga of Eastern Europe and Siberia in length ofthe P3—M3. The name S. prisca in our opinion should be synonymized with S. borealisand reduced to subspecific status. In which case the specimen previously called S. priscashould be considered the subspecies S. borealis prisca from Western Eurasia up toeast to the Yenisei River.

Belgium. — A study of a cast of fossil Saiga from Belgium (ZIAS 5, Leopold Canal)shows that the neurocranium length and the frontal-parietal angle more closely resembleS. borealis than S. tatarica (Table 2). The nasal bones are partly broken (the remainingpart is 49.5 mm) but their total length was apparently 65 mm, and as such the weresomewhat longer than in the recent 5. tatarica. One can assume that 5. borealis is atypical representative of the Mammoth Fauna, living not only in the Late Pleistoceneof Siberia, but also in coeval northern Eurasia west of the Atlantic coast. This assump-tion appears to be confirmed by findings of the large form of Saiga in Germany(Pahren), France (Grotte de Fees), and England (Pratt, 1977; Currant, 1986; Kahlke,1990).

Poland. - One skull from Poland was found in the Slupyanca Cave, Ojcew nearKrakow without accurate geological age. The posterior part is preserved completely asare the frontal bones. Both horncores are present, but the upper third of one is missing.The nasal and maxillary bones are broken. The frontal-parietal angle is 26 degrees. Thisis similar to minimal value of S. tatarica and is common to S. borealis. Neurocraniallength and width between the outer margins of the horncore bases (measurments 1 and7 ) are near the upper limit of corresponding values for 5. tatarica but well within therange of S. borealis. In the scatter diagram plotting absolute values of neurocraniallength against width between horncores (Figure 5), the skull from Slupyanca Cave isintermediate between S. borealis and S. tatarica but is closer to the values forS. tatarica. Therefore we consider this animal as Saiga sp., cf. S. borealis.

A second fossil Saiga from Maszyce cave, Ojcew near Krakow, of Magdalenian ageis related to the one from Slupyanca Cave (Kiernik, 1912). The measurements of thisspecimen are 129 mm width between orbits, 70.5 mm with of neurocranium behind thehorncores, 45.5 mm width between condyles. The length of P/3 to M/3 in the MaszyceCave jaw is 74 mm.

Ural River. — Specimens collected from the sand-banks of the Ural River, may bedated at the end of the Pleistocene by the mineralization of bones (Figure 4B). Thesizes of the skulls are within the range for recent S. tatarica. The posterior part is

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mm105 -

B

100-

95-

90-

oooo Cbo o

PLEISTOCENE SAIGA IN EURASIA

o

0 © •o • •

o o% B •O O

O • •o oo o •

oo

219

o oo. o

• 1Q 2© 3O A

fl85 90 95 100 fn-™

Figure 5 Correlation between upper neurocranial length (A) and breadth between horncores (B) in the skullsof fossil and recent saiga. 1 - S. borealis, 2 - 5 . cf. borealis, Slupianca, 3 - S. cf. tatarica, Ural River,4 - S. tatarica, recent.

relatively weakly elongated (index of neurocranial length is 109.5 to 112.1), and thefrontal angle is large at 34 degrees (Figure 6B). Ratios of neurocranial length and theinterorbital width for the Ural River specimens are more similar to S. tatarica than S.borealis (Figure 5). We think that this form is closely related to S. tatarica and couldbe defined as Saiga sp., cf. S. tatarica. This finding shows that Saiga of recent typeinhabited areas further south in Kazakhstan at the same time as S. borealis.

DISCUSSION

Taxonomy

Usually the Pleistocene Saiga is defined as 5. tatarica (Vereshchagin, 1959; Harrington,1981) or as a subspecies S. tatarica borealis (Bannikov, 1963, Kahlke, 1975, 1990,Baryshnikov, 1981). Our original researches of the material shows that S. borealis(including S. prisca, S. ricei) is an independent species. Thus, we confirm Sher's (1967)opinion of the distinct status of the northern Saiga.

Morphometric comparisons of the indices of the skulls of S. borealis (including theskull from Slupyanca Cave) and recent S. tatarica demonstrate a number of differencesbetween these species with a high degree of significance (Table 3). Particularly significant

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220 G. BARYSHNIKOV AND A. TIKHONOV

Table 3 Comparative

Measurements (mm)

Index (%)

size

n

ot skulls Saiga borealis

Saiga borealis

lim x

and

a

Saiga

n

tatarica

lim

Saiga

X

tatarica

G- T

1. 15 98.1-101.0 96.2 2.716 33 S2.8-94.8 88.4 3.580 +++2. 15 60.5-68.2 64.9 2.177 33 51.5-66.3 56.7 3.460 +++3. 14 117.2-129.1 121.4 1.833 33 104.7-122.6 116.5 4.310 +++4.* 46 26.5-34.7 30.3 1.737 46 25.0-32.7 29.6 2.0575. 13 119.7-137.2 128.9 5.193 33 119.0-136.3 128.4 5.6106. 14 40.5-54.6 50.2 3.840 33 42.2-50.6 47.0 2.360 +7. 16 90.6-110.1 99.6 4.850 33 89.2-102.3 96.5 2.833 +8* 50 24.6-32.9 28.6 1.898 56 24.7-31.7-".-• 27.5 2.675 +9. 13 65.1-77.5 73.2 2.470 33 65.3-73.3 70.3 1.555 +10. 11 83.1-92.8 87.6 2.644 32 74.0-93.2 83.2 5.089 ++11. 11 45.7-53.0 49.5 2.690 32 43.0-49.3 46.6 1.74212. 11 45.0-51.2 47.1 1.522 32 41.3-51.2 46.7 2.302 +13. 12 22 -24 28 12 26-38 33

7:1 14 95.9-110.0 102.6 3.874 23 103.1-116.9 109.9 4.230 +5:1 13 120.5-143.6 133.7 7.280 33 130.3-153.0 145.1 6.27 ++

10:12 11 167-201 187 9.72 32 164-197 178 9.27 +

* - Includes separate horncores from Crimea, Ural mountains and Siberia. In modern saiga we measuredleft and right horncores from one skull.

are the length of neurocranial part of the skull (measurements 1-3), and the width ofthe skull (measurments 7-9). S. borealis is characterized by large skull and teeth, andlonger nasals. Study of long bones shows that on average the size of S. borealisexceeded that of S. tatarica (Baryshnikov et al., 1990).

There are some differences in the structure of the astragalus. Material fromPalaeolithic sites of the Crimea shows that in most specimens the astragalus has astronger developed upper lateral-dorsal hillock and short lateral surface of distal blockas compared to recent Saiga. These morphological characteristics as well as the longbones point to peculiar morpho-functional adaptations of S. borealis and confirm itsspecies status.

Small recent S. mongolica is similar to S. borealis by the index of neurocranium.The length of nasals of the Mongolian Saiga are above the average of this measurementin S. tatarica. In our opinion the Mongolian Saiga should be defined as a subspecies,5. borealis mongolica. As such, it is a Pleistocene relict which survived in semidesertsof the hollow of great western lakes in Mongolia.

Distribution

The earlier finds of S. borealis in the territory of Russia are famous from the MiddlePleistocene (Khazarian fauna). It may be described as a separate subspecies of S. borealis.Saiga that also occurs in France and probably in Alaska (Harrington, 1981).

Fossil remains of Saiga from the Volga River localities are very rare. It was neverdescribed from the lower Kama River (Gromova, 1932). Only a single horncore fromthe Chogra River was found in museum collections of Pugachevsk (Belvaeva, 1935).Saiga remains become more numerous on the Lower Volga (collections of ZIAS). Fromthe type locality of the Khazarian complex (Chernyi Yar in Volgograd region) weredefined the following species: Mammuthus chosaricus; Equus chosaricus; Camelusknoblochi; Megaloceros giganteus; Bison priscus longicornis; Saiga tatarica and Canislupus (Alekseeva, 1990, p. 55).

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In the late Pleistocene (Wiirm/Waldayi) the fossil remains of Saiga become morenumerous. It inhabited arctic and boreal zones from the British Isles on the west upto northwestern Canada on the east (Figure 7). We can separate two subspecies fromthis vast area: S. borealis prisca Nehring (Europe, Ural, West Siberia up to the YeniseyiRiver) and 5. borealis borealis Tchersky (East Siberia, Alaska).

The remains of Saiga from the Pleistocene of West Europe and Alaska are small innumbers and as a rule without exact geological age, so it is not easy to retrace thedynamics of range and density of Saiga in the Late Pleistocene in the territory of Russia.However, Saiga lived in Western Europe to the upper Magdalenian (Harrington, 1981;Currant, 1986).

In early Wtirm (Waldayi) the remains of Saiga in Russia are noted mainly fromsouthern regions. Particularly, they were numerous in the Crimea and dominatedMousterian sites. The accompanying fauna in the Crimea includes the following species:Mammuthus primigenius; Coelodonta antiquitalis; Equus caballus; Equus hydruntinus;Cervus elaphus; Megaloceros giganteus; Rangifer tarandus; Bison priscus; Canis lupus;Vulpes vulpes; Vulpes corsac; Alopex lagopus; Ursus spelaeus crimaeus; Crocutaspelaea; and Panthera spelaea. Saiga also inhabitated the north, penetrating EasternSiberia. The skull of Saiga from the Balyktiach River was refered to deposits of theKarginskyi level (Lazarev Tomskaya, 1987, pp. 114-129). Harrington (1981) supposedthat dispersal of Saiga was in waves, beginning during the Middle Pleistocene (Illinoian).The radiocarbon date of a horncore from Usuktuk River, 37,000 ± 990 BP (GSC-3050),shows that Saiga penetrated to northern Alaska in the Middle Wisconsin (Harrington,1981, p. 225).

In the late Wiirm (Waldayi) the remains of Saiga are known from the Crimea,Russian Plane, Ural, Altayi, and East Siberia. They are small in number in most areas,but are very numerous in the Crimea. It is remarkable that the fossil remains of Saigaare never found in the Baikal region (Ermolova, 1987; Ovodov, 1987). The report ofSaiga remains from the Upper Palaeolithic site of Suchotino near Chita are erroneous(Kasparov, 1988, p. 99).

The northern Saiga S. borealis in the Late Wiirm is an index element of theMammoth Fauna. Its remains have been found together with Mammuthus primigenius;Coelodonta antiquitatis; Equus latipes; Equus lenensis; Alces dices; Rangifer tarandus;Bison priscus; Ovibos moschatus; Canis lupus; Gulo gulo; Crocuta spelaea andPanthera spelaea.

Further south in the Transcaucasus, Kazakhstan and Middle Asia, S. tataricaapparently existed in arid steppe and semidesert areas. S. t. binagadensis had smallmetacarpal and metatarsal bones that are relatively narrow in the distal part. Probablythe Apscheron Saiga inhabited hot dry steppes. Other animals thought to indicate warmclimate are found together with Saiga in the Bynagady site, such as Equus caballus;Equus hydruntinus; Rhinoceros binagadensis; Sus apscheronicus; Cervus elaphus;Megaceros euryceros; Bos mastanzadei; Canis lupus; Vulpes corsac; Vulpes vulpes;Ursus arctos; Vormela peregusna; Meles meles; Panthera spelaea and Acinonyx jubatus(Vereshchagin, 1959, p. 141).

Pleistocene Saiga are rare in Kazakhstan. Remains of Saiga sp., cf., S. tatarica arefound on beaches of the Ural River together with bones of Mammuthus primigenius,Rhinoceros tichorhinus, Equus caballus, Rangifer tarandus, Alces alces, Cervuselapphus, and Bos primigenius (Vereshchagin, Gromov, 1952). Such a grouping of largeanimals is characteristic of the southern zone of the Mammoth Fauna. A transition tothe fauna of the Kazakhstan steppes is observed in the composition of the rodent fauna:Citellus pygmaeus; Allactaga jaculus; Cricetus cricetus; Meriones tamariscinus;Lagurus luteus; Lagurus lagurus and Ellobius talpinus. Here in the region of the Ural

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Figure 7 The Pleistocene finds of Saiga.Triangles - fossil skulls : A - Leopold Canal (Belgium), B - Slupianca Cave (Poland), C - Tungus (Volga River near delta of Cheremshan River), D - Luchka (nearSaratov), E - Ural River, F - Tura River, G - Uschaika River (near Tomsk) H - Kuzbass, I - Aidorach River, (near Minusinsk), I - Yenisei River (near Krasnoyarsk),K - Vilui River, L - Olenek River, M - Delta of Lena River, N - Kolyma River, O - Lillian Creek (near Livengood), P - Gold Hill (near Fairbanks).Circles - Localities of the Pleistocene remains of saiga (numbers near circles are similar to number of the localities from appendix).

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PLEISTOCENE SAIGA IN EURASIA 223

steppes, S. borealis and 5. tatarica could have been sympatric. The remains of Saigaare found in the upper palaeolithic site of Ushbas Cave in southern Kazakhstan (Dzhambulregion) together with Rhinoceros sp, Bison sp, (Alpysbaev, 1962). There are no fossilremains of Saiga in Middle Asia. The definition of Saiga from Samarkandskaya (upperPalaeolithic) are erroneous (Dzhurakulov et al., 1980).

Paleoecology

It is commonly held that species represented by fossils have similar habitat require-ments to those of their closest recent species. S. tatarica today lives in dry semidesertswith low sparse vegetation and thin snow cover. It forms large herds which migratewidely. The main food includes the following: Kochia prostata; Artemisia; Ephedra;and Anabasia (Bannikov, 1963: Zhirnov, 1982).

S. mongolica inhabits lowlands of the great western lakes of Mongolia in aridconditions. Small herds of these animals do not migrate long distances. It feeds on:Stipa gobica, Allium, and Anabasia. Artemisia is not a common food of the mongoliansaiga.

S. borealis inhabitated vast grassy planes together with other herbivorous animals -Mammuthus, Coelodonta, Equus, and Bison. In addition it ranged with Rangifer andOvibos on the north. The above species of proboscideans and ungulates are thought tohave lived in large herds in grasslands of the periglacial steppes, tundra-steppes andarctic cryogenic savannah. Winter is assumed to have been very cold with little snow,and the summers were probably short and dry because morphology of northern Saigasuggests they were best adapted to open country that lacked deep snow.

We can suggest that the morphological similarity between S. borealis and S. mongolicais closely related to their ecological similarity. Investigation of fossil remains of Saigafrom the Palaeolithic of the Crimea shows that Crimean Saiga inhabitated there all yeararound and bred (Baryshnikov et al., 1990). Probably their food included cereals andnot Kochia and Artemisia, as in S. tatarica. Confirmation of this lies in the proportionsof lower molars Ml and M3 of Saiga from the Palaeolithic of the Crimea (Baryshnikovet al., 1990).

Thus the wise distribution of Pleistocene Saiga in Eurasia, can be explained bycorresponding distribution in the Pleistocene tundra-steppe or cryogenic savannah, inareas which now are tundra and taiga zonal landscapes. Along with the mammoth, woollyrhinoceros, reindeer and musk-ox, the northern S. borealis was a natural part of ecologicalstructure of the mammoth biome.

General Appearance

S. borealis was a rather large antelope. Adult males had a height in shoulders140-150 cm and weight near 65-70 kg. They had a massive head with a large horns.The skull was characterized by elongated posterior part and nasals. In comparison withS. tatarica, by the frontal-parietal angle, S. borealis had a small hump-nose profile. Bythis feature it resembles the recent S. mongolica. Firstly this was noted by Bannikov(1963, p. 282). Our suggestion about small "hump-nose" profile of the PleistoceneSaiga are confirmed by the Palaeolithic painting of the Saiga's head from French caves(Bequen et al. 1986, Figure 5)

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224 G. BARYSHNKOV AND A. TKHONOV

Acknowledgements

We are especially grateful to Dr. A. Nadachowski (Institute of Animal Systomaticsand Evolution, Krakow, Poland) for permission to investigate the fossil skull of Saigafrom Poland, and to Dr. P. Sartonaer (Institut Royal des Sciences Naturelles deBelgique, Bruxelles) for information about Saiga material from Belgium. The help ofA.K. Kasparov and O.R. Potapova (Zoological Institute of Russian Academy of Sciences,Leningrad) is gratefully acknowledged.

The paper's improvement was kindly made by the editing of Drs. A. Barnosky(University of California, Berkely, USA), and E. Anderson, T. Hardy, and R. Stucky(Denver Museum of Natural History, USA). We are also thankful to Carol Carpenterfor typing the edited manuscript.

Drawings were made by G.I. Baranova, and photos by G.A. Apanovich.

References

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Alpysbaev, Kh. A. (1962) The finds of stone age sites in the Karatau Ridge. Trudy Instituta istorii,archeologii i ethnografii AN Kaz SSR, Alma-Ata, 14, 12-37 (in Russian)

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Bannikov, A.G. (1954) Mammals of Mongolian People's Republic. Trudy Mongolskoi komissii,Moscow, 53, 669 p. (in Russian)

Bannikov, A.G. (1963) Die Saiga-Antelope (Saiga tatarica). Wittenberg, A. Ziemsen Verlag. 143pBaryshnikov, G.F. (1981) Order Artiodactyla Owen. 1848. In: Gromov, I.M. and Baranova,

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Begouen, R., Clottes, J., Giraud, J-P. and Rouzard, F. (1986) Le propulseur au saiga d' Enlene.Bulletin de la Sociéte Prěrristorique. Ariège-Pyrénèes. 41, 11-22

Bibikova, V.I. and Starkin, A.V. (1989) Description of the osteological matherial from the LatePalaeolithic site Anetovka 2. In: Stanko, V.N., Grigorieva, G.V., Shvaiko, T.N.,Pozdnepaleoliticheskoe poselenie Anetovka 2. Prilozhenie., Kiev. Naukova Dumka, 127-131(in Russian)

Chersky, I.D. (1876) Antilope (Saiga) borealis n. spec. fossilis. Izvestia Sibirskogo otdelaImperatorskogo Geographicheskogo Obschestwa. St. Petersburg, 7, 4-5, 145-151 (in Russian)

Currant, A.P. (1986) The Late glacial mammal fauna of Gough's Cave. Cheddar Somerset.Proceedings of University of Bristol Speleological Society, Bristol, 17, 3, 286-304

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Dzurakulov, M.D., Kholushkin, Yu. P., Kholushkina, V.A. and Batyrov, B.H. (1980) Samarkandskayasite with comment of its position within the Late Palaeolithic of Middle Asia. In: Larichev,V.E., ed.. Paleolit Srednei i Vostochnoi Asii. Novosibirsk. Nauka, 51-95 (in Russian)

Ermolova, N.M. (1978) The theriofauna of Angara Valley in Late Anthropogene. Novosibirsk.Nauka, 223 p. (in Russian)

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Harrington, C.R. (1981) Pleistocene saiga antilopes in North America and their paleo-environmental implications. In: Mahaney W.C., ed., Quaternary paleoclimate. Norwich.University of East Anglia. 193-225

Hoffecker, J.F., Baryshnikov, G.F. and Potapova, O.R. (1989) Mousterian bison hunters of thenorthern Caucasus: Analysis of faunal remains from Ilskaya 1. Current Research in thePleistocene., 6, 69-72

Kahlke, H.D. (1975) Der Saiga-Fund von Bottrop/Westfalen. Quartar, 26, 135-146Kahlke, R.D. (1990) Der Saiga-Fund von Pahren. Ein Beitrag zur Kenntniss der Paläarktischen

Verbreitungsgeschichte der Gattung Saiga Gray 1843 unter besonderer Berücksichtigung desGebeites der DDR. Eiszeitalter und Gegenwart. Hannover, 40, 20-37

Kasparov, A.K. (1986) Mammal remains from the Late Palaeolithic site Sukhotino 1 inTransbaikalia. Trudy Zoologicheskogo Instituta AN SSSR. 149, 98-106 (in Russian)

Kiemik, E. (1912) Materyaly do paleozoologii dyluwialnych ssakow Ziem Polskich. Cześć 3.Szczatki suhaka (Antilope saiga) z jaskini Maszyckiei kolo Ojcowa. Rozprawy wydzialymatematychno-pryyrodnichego Akademii Umiejftnosci. Krakow, ser, 3, 12, 399-434

Kolossov, Yu. G. (1983) The Mousterian sites in Belogorsk Region (the problems of the CrimeanLate Palaeolithic periodization). Kiev. Naukova Dumka, 208 p. (in Russian)

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Kosintsev, P.A. and Borodin, A.V. (1990) The theriofauna of eastern slope of Northern Ural inthe Late Pleistocene and Holocene. Trudy Zoologicheskogo Instituta AN SSSR, 212, 120-134(in Russian)

Kuzmina, I.E. (1971) Forming of theriofauna of Northern Ural in late anthropogene. TrudyZoologicheskogo Instituta AN SSSR, Leningrad, 49, 44-122 (in Russian)

Kuzmina, I.E. (1975) Some data on the Late Pleistocene mammals from the Middle Urals. BulletinKomissii po izucheniu chetvertichnogo perioda, 43, 63-77 (in Russian)

Lazarev, P.A. and Tomskaya, A.I. (1987) Mammals and biostratigraphy of late cainozoic ofNorthern Yakutia, Yakutsk, Izdatelstwo Yakutskogo filiala SO AN SSSR, 172 p. (in Russian)

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Ovodov, N.D. (1987) Fauna of the Palaeolithic sites Tolbaga and Varvarina Gora in WesternTransbaikalia. In: Resanov I., Basarova L. Khamsina E. ed. Prirodnaya sreda i drevnii chelovekv pozdnem anthropogene. Ulan-Ude. Geologicheskii Institut, 122-140 (in Russian)

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Pidoplichko, I.G. (1956) Matherials for studying of fossil faunas of the Ukraine, Vol. 2. Kiev,AN USSR, 236 p. (in Ukrainen).

Prat, F. (1966) Deuxieme partie. Chapter 9. Les antilopes, Atlas de prehistoire, Faunes at floresprehistoriques de l'Europe occidentale, 3, 323-336, Paris

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Sher, A.V. (1971) Mammals and stratigraphy of the Pleistocene of far north-east of the USSRand Northern America, Moscow, Nauka, 312 p (in Russian)

Vereshchagin, N.K. (1959) Mammals of Caucasus. History of forming of the fauna. Moscow-Leningrad, Izdatelstwo AN SSSR, 704 (in Russian)

Vereshchagin, N.K. and Baryshnikov, G.F. (1980) Mammals of northern Crimea foothills in thePalaeolithic (on the base of food remains from the caves Chokurcha, Staroselye and Mamat-Koba. Trudy Zoologicheskogo Instituta AN SSSR, Leningrad, 93, 26-49 (in Russian)

Vereshchagin, N.K. and Gromov, I.M. (1952) History of fauna of vertebrates from the regionof lower part of River Ural. Trudy zoologicheskogo Instituta AN SSSR, Leningrad, 9,1226-1269 (in Russian)

Vereshchagin, N.K. and Kolbutov, A.D. (1957) Animal remains from the mousterian site atStalingrad and stratigraphical position of the Paleolithic stratum. Trudy ZoologicheskogoInstituta AN SSSR, Leningrad, 22, 75-111 (in Russian)

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APPENDIX

Fossil remains of in the Pleistocene assamblages with Proboscidea. Perissodactyla andArtiodactyla in the former USSR"'2'

Middle Pleistocene1. Bynagady, Baku, Transcaucasia Riss or Riss-Wiirm (Moskow or Mikulino)

(Vereshchagin, 1959, p. 141)

Equus caballus ssp. 154Equus aff. hydruntinus 73Rhinoceros binagadensis 31Sus apscheronicus 11Cervus elaphus binagadensis 52Megaceros cf. euryceros 2Bos mastanzadei 6Saiga tatarica binagadensis 82Ovis cf. ammon . 1

2. Ural River, Sandy banks Pleistocene (mixed material) (Vereshchagin, Gromov. 1952,p. 1244)

Elephas cf. primigenius 17Equus caballus 166Equus cf. hydruntinus 9Equus hemionus 1Rhinoceros cf. tichorinus 11Elasmotherium sibiricum 6Camelus knoblochi 12Cervus elaphus 25Megaceros euryceros 2Alces alces 2Rangifer tarandus 1Bos primigenius 58Bison priscus 57Saiga tatarica 91

IJ According to Alpine and East-European stratigraphical scales2) Number of individuals in localities NN°l,10 in other cases the number of remains.

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Upper Pleistocene, Lower Wiirm (Lower Waldai)

3. Buzduzhany 1 Grotto Rakovets River, Moldova Mousterien (David, 1980, p. 38)

Mammuthus primigenius 67Equus latipes 418Equus (Asinus) hydruntinus 6Coelodonta antiquitatis 28Cervus elaphus 38Megaceros euryceros 7Rangifer tarandus 54Cervinae 26Bison priscus 214Saiga tatarica 3

4. Iliinka Grotto Odessa, Ukraine Mousterien (Pidoplichko, 1956, p. 99)

Equus equus 14Rhinoceros antiquitatis 1Cervus elaphus 11Capreolus capreolus 2Bison priscus 24Saiga tatarica 3

5. Staroselie Bakhchisarai, the Crimea Mousterien (Vereshchagin, Baryshnikov, 1980,p. 39)

Mammuthus primigenius 136Equus caballus 134Equus hydruntinus 17360Coelodonta antiquitatis 18Cervus cf. elaphus 61Capreolus cf. capreolus 4Megaceros giganteus 10Rangifer tarandus 22Bison priscus 140Saiga tatarica 404

6. Shaitan-Koba Grotto Bodrak River, the Crimea Mousterien (Gromova, Gromov, 1937,p. 92)

Elephas sp. (cf. primigenius, trogontherii) 1Equus (Equus) caballus foss 11Equus (Asinus) sp. 50Cervus elaphus 7Bos sp.? (cf. Bison priscus) 2Saiga tatarica 177

7. Chokurcha Grotto Simpheropol, the Crimea Mousterien (Vereshchagin, Baryshnikov,1980. p. 32)

Mammuthus primigenius 3041Equus caballus 147

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Equus hydruntinus 730Coelodonta antiquitatis 4Cervus elaphus 59Megaceros giganteus 3Rangifer tarandus 7Bison priscus 813Bison aut Bos 38Saiga tatarica 69

8. Kosh-Koba Grotto Zuya River, the Crimea Mousterien (Gromova, Gromov, 1937,p. 92)

Elephas sp.? (cf. primigenius, trogontherii) 5Equus (Equus) caballus foss 86Equus (Asinus) sp.? 37Rhinoceros antiquitatis 19Sus scrofa 13Cervus elaphus 26Cervus euryceros 8Rangifer tarandus 6Bos sp.? (cf. Bison priscus) 2Saiga tatarica 116

9. Kiik-Koba Zuya River, the Crimea Mousterien (Gromova, Gromov, 1937. p. 92)

Elephas sp.? (cf. primigenius trogontherii) 43Equus (Equus) caballus foss 103Equus (Asinus) sp.? 3Rhinoceros antiqiutatis 5Sus scrofa 2Cervus elaphus 16Cervus euryceros 232Rangifer tarandus 2Bos sp.? (cf. Bison priscus) 14Saiga tatarica 144Ovis sp.? (cf. argaloides) 1

10. Volchii Grotto Beshterek River, the Crimea Mousterien (Bader, Bader, 1979, p. 25)

Mammuthus primigenius 14Equus caballus 3Equus hydruntinus 11Coelodonta antiquitatis 1Sus scrofa 1Cervus elaphus 1Megaceros giganteus 2Rangifer tarandus 1Bison priscus 3Saiga tatarica 24Ovis sp. 1

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11. Zaskalnaya 5 Grotto Biuk-Karasu River, the Crimea Mousterien (Kolossov, 1983,p. 28)

Mammuthus primigenius 212Equus caballus 77Cervus elaphus 6Megaceros giganteus 1Bison priscus 6Saiga tatarica 351

12. Zaskalnaya 6 Grotto Biuk-Karasu River, the Crimea Mousterien (Kolossov, 1986,p. 11)

Mammuthus primigenius 382Equus caballus 542Equus hydruntinus 33Coelodonta antiquitatis 1Cervus elaphus 6Megaceros giganteus 4Rangifer tarandus 54Bison priscus 9Saiga tatarica 483

13. Prolom 2 Kuchuk-Karasu River, the Crimea Mousterien (Kolossov, 1986. p. 11)

Mammuthus primigenius 77Equus cf. latipes 589Equus (Hydruntinus) hydruntinus 128Coelodonta antiquitatis 63Sus scrofa 8Cervus elaphus 25Megaceros giganteus 16Rangifer tarandus 92Bison priscus 157Saiga cf. borealis 3093Capra aut Ovis 1

14. Adzhi-Koba Cave (stratum 3) Karabi-Yaila, the Crimea Mousterien. (Gromova,Gromov, 1937 p. 92)

Equus (Equus) caballus 1Equus (Asinus) sp.? 1Rhinoceros antiquitatis 6Cervus elaphus 6Bos sp.? (cf. Bison priscus) 2Saiga tatarica 130Ovis sp.? (cf. ammon) 10

15. Mamat-Koba Cave Karabi-Yaila, the Crimea Mousterien? (Vereshchagin, Baryshnikov,1980, p. 40)

Equus caballus 2Equus hydruntinus 3

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230 G.BARYSHNKOVANDA.TKHONOV

Coelodonta antiquitalis 1Cervus elaphus 12Capreolus capreolus 5Rangifer tarandus 2Saiga tatarica 577

16. Ilskaya 1 II River. Northern Caucasus Mousterien (Hoffecker et al., 1989, p. 70)

Mammuthus cf. chosaricus 7Equus (Equus) cf. mosbachensis 21Equus (Hydruntinus) hydruntinus 6Cervus elaphus 16Megaceros giganteus 25Bison priscus 1334Saiga tatarica 2

17. Sukhaya Mechetka Volgograd, Volga Rover Mousterien (Vereshchagin, Kolbutov,1957, p. 84)

Elephas primigenius 51Equus caballus 42Cervus elaphus 15Rangifer tarandus 1Bison priscus 366Saiga tatarica 36

Upper Wurm (Upper Waldai)

18. Starye Duruitory Grotto (stratum 2) Duruitory, Moldova Upper Palaeolithic (David,1980, p. 26-27)

Mammuthus primigenius 50Equus latipes 4969Equus sp. 36Coelodonta antiquitatis 54Sus scrofa 1Cervus elaphus 74Capreolus capreolus 28Megaceros giganteus 18Alces alces 2Rangifer tarandus 1249Bison priscus 564Saiga tatarica 3

19. Anetovka 2 Yuzhnyi Bug River, Ukraine Upper Palaeolithic: 18040+150 (LE-2424)(Bibikova, Starkin, 1989, p. 129)

Equus latipes 60Cervus elaphus 1Rangifer tarandus 434Bison priscus 21739 .Saiga tatarica 5

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PLEISTOCENE SAIGA IN EURASIA 231

20. Siuren 1 Grotto Belbek River, the Crimea Upper Palaeolithic (Gromova, Gromov,1937, p. 92)

Equus (Equus) caballus foss. 61Sus scrofa 25Cervus elaphus 50Cervus euryceros 170Rangifer tarandus 18Bos sp.? (cf. Bison.primigenius) 58Saiga tatarica 703

21. Adzhi-Koba Cave (stratum 2) Karabi-Yaila, the Crimea Upper Palaeolithic (Gromova,Gromov, 1937, p. 92)

Equus (Equus) caballus foss. 18Equus (Asinus) sp.? 16Cervus elaphus 9Rangifer tarandus 9Bos sp.? (cf. Bison priscus) 5Saiga tatarica 617Capra (Ibex) sp.? (aff. sibirica) 1Ovis sp.? (cf. ammon) 1Ovis sp.? (cf. argaloides) 1

22. Kostenki 1 (stratum 3) Kostenki, Don River Upper Palaeolithic: stratum 1 -21300+400 (GIN-2534); 22300+230 (GIN-1870); 23000+500 (GIN-2528);24100+500 (GIN-2524). (Vereshchagin, Kuzmina, 1977, p. 100)

Mammuthus primigenius 5Equus caballus latipes 71Rangifer tarandus 69Saiga tatarica 3

23. Kostenki 4 (Vereshchagin, Kuzmina. 1977, p. 102)

Mammuthus primigenius 272Equus caballus latipes 421Coelodonta antiquitatis 2Sus scrofa 1Cervus elaphus 3Rangifer tarandus 76Bos aut Bison 5Saiga tatarica 6

24. Kostenki 8 (stratum 2) 27700+750 (GIN-10509) (Vereshchagin, Kuzmina, 1977,p. 104)

Mammuthus primigenius 67Equus caballus latipes 62Coelodonta antiquitatis 18Cervus elaphus 1Megaceros giganteus 2Rangifer tarandus 9

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232 G. BARYSHNKOV AND A. TIKHONOV

Bos aut Bison 11Saiga tatarica 7

25. Kostenki 14 (stratum 2) 19300+200 (LE-1400): 26400+660, 28200+700 (LU-59)(Vereshchagin, Kuzmina, 1977, p. 107)

Mammuthus primigenius 6Equus caballus latipes 2083Coelodonta antiquitatis 16Cervus elaphus 41Rangifer tarandus 11Bos aut Bison 1Saiga tatarica 1

26. Bliznetsov Grotto Chanva River, Middle Urals Upper Palaeolithic (Kuzmina, 1975,p. 64)

Mammuthus primigenius 391Equus caballus latipes 79Coelodonta antiquitatis 17Alces alces 1Rangifer tarandus 21Bison priscus 1Saiga tatarica 15Ovibos moschatus 6

27. Stolbovoi Grotto Usva River, Middle Urals Upper Palaeolithic (Kuzmina, 1975,p. 64)

Equus caballus uralensis 19Coelodonta antiquitatis 5Rangifer tarandus 109Bison priscus 2Saiga tatarica 8

28. Medvezhya Cave Pechora River, North Urals Upper Palaeolithic: 17980+200(LE-233) (Kuzmina, 1971, p. 56)

Mammuthus primigenius 171Equus caballus 429Coelodonta antiquitatis 109Capreolus capreolus 28Alces alces 18Rangifer tarandus 7669Bison priscus 44Saiga tatarica 41Ovibos moschatus 216

29. Shaitan Grotto Ivdel River, Middle Urals Upper palaeolithic (Kosintsev, Borodin.1990, p. 123)

Mammuthus primigenius 1Equus cf. uralensis 155

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Coelodonta antiquitatis 67Cervus elaphus 4Rangifer tarandus 248Bison priscus 65Saiga tatarica 4Ovibos moschatus 12

30. Shikaevka 2 Shikaevka, Tobolo River Upper Palaeolithic (Petrin, Smimov, 1975,p. 84-85)

Mammuthus primigenius 223Rangifer tarandus? 2Saiga tatarica 3

31. Novoselovo 7 Popovka River, Yenisei River Upper Palaeolithic (Abramova, 1979,p. 151)

Equus caballus 5Rangifer tarandus 887Bison priscus 5Saiga tatarica 3

32. Tashtyk 1 (stratum 3) Tashtyk River. Yenisei River Upper Palaeolithic: stratum 1- 12180+120 (LE-771) (Zeitlin, 1979, p. 98)

Equus caballus 4Cervus elaphus 1Rangifer tarandus 31Bison aut Bos . 3Saiga tatarica 2

33. Kokorevo 2 Yenisei River Upper Palaeolithic: 13330+100 (GIN-90) (Zeitlin, 1979,p. 112)

Mammuthus primigenius 148Equus caballus 93Cervus elaphus 162Alces alces 12Rangifer tarandus 11Bison priscus 15Saiga tatarica 11Ovis ammon 21

34. Aphontova Gora 2 Krasnoyarsk. Yenisei River Upper Palaeolithic: 20900+300(GIN-117) (Gromov, 1948, p. 322)

Elephas primigenius 6Equus (Equus) sp. 2Equus hemionus 1Cervus elaphus 3Capreolus pygargus 2Rangifer tarandus 46Bos aut Bison 2

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234 G.BARYSHNIKOVANDA.TIKHONOV

Bos sp. 1Saiga tatarica 2Capra sibirica 1Capra sp. 2Ovis ammon 3

35. Aleshkinskaya Zaimka Kolyma River 14980+100 (MAG-470): 15000+200(MAG-468) (Sher, 1971, p. 95)

Mammuthus primigenius 17Equus caballus subsp. 18Rangifer tarandus 8Bison priscus subsp. 4Saiga ricei 3

36. Northern Yakutia Bolshoy Liachovsky Island, Delta of Lena River (Chersky, 1891.p. 68-69)

Elephas primigenius 71Equus caballus 659Rhinoceros tichorhinus 32Cervus canadensis 4Alces palmatus 7Rangifer tarandus 794Bison priscus 344Colus saiga 4Ovibos moschatus 159Ovis nivicola 2

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