Pleistocene Lagomorphs of Eurasia

Home , Daurian pika, Northern pika, Woolly hare

Alexander Ave r i a n ov Zoological Institute, R A S , S t . Pe t e r s b u r g

Pleistocene lagomorphs of Eurasia

Av e r i a n o v, A., 2001 - Pleistocene lagomorphs of Eurasia - D E I N S E A 8: 1-13 [ISSN 0923-9308]. Published 09 November 2001.

In the Pleistocene of Eurasia three species of Prolagidae, 17 species of Ochotonidae and 16 species of Leporidae are known. The species diversity of lagomorphs gradually increased during the Pleistocene and in the late Pleistocene was the same as Recent. Some relict genera became extinct during the early Pleistocene (P l i o p e n t a l a g u s, A l i l e p u s), others in the Middle Pleistocene (O c h o t o n o i d e s, H y p o l a g u s a n d S e r i c o l a g u s). At least two lagomorph species became extinct during the Holocene (P rolagus sard us and Ochotona transcaucasica). The Pleistocene was not a critical period in the evolution of Lagomorpha. The extinction was caused by global cooling and was restricted to relict taxa mostly. During the Pleistocene intensive speciation took place in the genera O c h o t o n a, O ry c t o l a g u s and L e p u s in the Old World, and S y l v i l a g u s in the New World. The Recent time is the period of flourishing and biological progress of phylogenetically young groups of lagomorphs (O c h o t o n a, Leporinae), which started in the late Pliocene and continued during all the Pleistocene.

This paper is presented in the framework of the 2n d International Mammoth Conference, held in Rotterdam, 16-20 May 1999.

Correspondence: Alexander Av e r i a n o v, Zoological Institute, Russian Academy of Sciences, 199034 St. P e t e r s b u rg, Russia. e-mail: [email protected]

Keywords: Lagomorpha, Prolagidae, Ochotonidae, Leporidae, Eurasia, Pleistocene

I N T RO D U C T I O N The Pleistocene is characterised by the most Here we fall sometimes into circular thinking, complete fossil record for many groups of when our determination of Pleistocene forms modern mammals. Here we can trace the ori- is greatly influenced by the current taxonomy gin and early evolution of some recent spe- of recent species and we then use these cies. However, the taxonomic approach paleontological data for the reconstruction of applied to the recent mammals is sometimes the history of recent species and their phylo- d i fficult to apply to Pleistocene forms. T h e genetic relationships. lagomorphs is a good example of such dispa- rity between systematics of Pleistocene and In my report I will briefly review the recent forms. The taxonomy of all recent spe- Pleistocene fossil record for the Eurasiatic cies and subspecies is largely based upon lagomorphs and will try to elucidate how external characters (pelage), and, to a lesser these data really help us to understand the extent, on some cranial and dental characters. evolutionary history of the recent taxa and to The majority of Pleistocene remains of lago- solve some current taxonomic problems in morphs are just isolated and fragmented the field. I provide these data to emphasise bones, determination of which is based exclu- some important gaps in the fossil record and sively on their absolute size and on few den- to settle up the perspectives for the future tal characters, specifically on the structure of work. This review is based on the unpublis- the third lower premolar (p3). However, the hed taxonomic revision of both fossil and variation of these characters in recent lago- recent lagomorphs, carried out by the author. morphs populations is not really understood.

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All the Pleistocene taxa of lagomorphs could terised by a large size, a weak or absent cro- be easily divided into two major groups. T h e chet, and an enlarged protoconulid, exceeding first includes the relict taxa, which became the size of protoconid. A similar but somew- extinct during the Pleistocene or Holocene, or hat smaller lagomorph, P rolagus figaro depe - which are close to extinction nowadays. T h e reti LO P E Z MA RT I N E Z , 1975, was reported other group includes taxa which underwent from the Early Pleistocene locality Bagur 2 in biological progress during the Pleistocene Spain (Lopez Martinez 1989). Otherwise it is and which flourish now. The latter group known from the Ruscinian of France and includes species of the three genera, namely Spain. The P ro l a g u s from the Middle O ry c t o l a g u s, L e p u s and O c h o t o n a . Pleistocene locality Montagnola in Italy, described as P. s a rd u s by Fondi (1970, REVIEW OF SPECIES 1972), probably should be referred to the Villafranchian Italian species P rolagus sava - Relict taxa gei BE R Z I, 1967, contra Lopez Martinez (1989), who referred this material to the A few lineages of the common Mediterranean Ruscinian French and Spanish P. michauxi Neogene ochotonid-like lagomorph P ro l a g u s LO P E Z MA RT I N E Z , 1975. The holotype of P. have survived in the Pleistocene (Fig. 1). s a v a g e i and few other known p3 of P ro l a g u s Prolagus figaro figaro LOPEZ MARTINEZ, 1975 from the Italian Villafranchian (Berzi 1967; is known from the type locality only, Middle Rook & Masini 1990) lack a crochet, which Pleistocene Capo Figaro in Sardinia, Italy is represented in some specimens in the (Lopez Martinez & Thaler 1975). It is charac- Montagnola population and in P. m i c h a u x i .

Figure 1 A p p r oximate distri bution of lagomorph relict taxa in the Pleistocene of Eura s i a : 1 - Prolagus figaro, 2 - Prolagus sardus, 3 - Prolagus sava g e i, 4 - Hypolagus beremendensis bra c h y g n a t h u s, 5 - Caprolagus hispidus, 6 - Caprolagus lapis, 7 - S e ricolagus bra c h y p u s , 8 - Ochotonoides complicidens, 9 - Hypolagus schreudera e, 1 0 - Alilepus zhoukoudianensis, 1 1 - Pliopentalagus progressivus, 1 2 - Pentalagus furn e s s i.

2 AVERIANOV: Pleistocene lagomorphs

H o w e v e r, the variation of p3 in P. s a v a g e i i s Province, China, described as O c h o t o n a sp. 1 not adequately known and it is looks prefera- (Qiu et al. 1984), could be referred to this ble to refer all the peninsular Italian species and thus represents its last fossil Pleistocene P ro l a g u s to this species. r e c o r d .

The Pleistocene-Holocene P rolagus sard u s The relict Pliocene-Pleistocene rabbit (WA G N E R, 1829) from Corsica, Sardinia and Hypolagus schreuderae TEILHARD DE CHARDIN, adjacent smaller islands was one of the larg e - 1940 was originally described from the Early st species of P ro l a g u s. It was surviving on the Pleistocene Locality 18 (=Yangshaozun) near small island of Tavolara off the Sardinian Beijing, China, where it was represented by a coast as late as the eighteenth century number of skulls and at least one skeleton (Kurtén 1968). ( Teilhard de Chardin 1940). This species is also known from the Early Pleistocene of Another European relict Pleistocene lago- Hebei Province (Cai 1989). It is characterised morph was a rabbit Hypolagus bere m e n d e n s i s by a large size and a complicated P2 with (PE T É N Y I , 1864). The Pleistocene populations three folds on the anterior side. are referred to the subspecies H. b. brachyg - n a t h u s KO R M O S , 1934, which had gradually The rabbit Alilepus zhoukoudianensis CH E N G , evolved from the Pliocene H. b. bere m e n d e n - CA O, TI A N & LI, 1995 is known from the type s i s (PE T É N Y I , 1864). It differs from the latter locality only, the Early Pleistocene beds of by its larger size and by dominance of more Zhoukoudian near Beijing, China (Cheng e t complicated variants of P2 and p3 (Fladerer a l . 1995). In Europe the last undoubted & Reiner 1996). Apparently all European remains of A l i l e p u s are known from the Early localities with the last H y p o l a g u s r e m a i n s Villafranchian (Odessa catacombs: A. ucraini - (Podumci, Betfia 5, Varbezhnitsa and others) c u s GU R E E V , 1964). The p3 of this species should be dated to the Cromer interg l a c i a l was not figured, so the attribution of the type (Gunz-Mindel); and these rabbits are not material to this genus remains to be demon- known from Mindel-Riss deposits. If so, s t r a t e d . extinction of this species was probably connected with the Mindel (=Oka) Glaciation (IV The Plio-Pleistocene rabbit Sericolagus bra - criosuperclimatem according to Zubakov c h y p u s (YO U N G , 1927) is known from the 1 9 8 6 ) . Early to Middle Pleistocene of northern and north-eastern China (Young 1927, 1935; In Asia a greater number of relict lagomorphs Teilhard de Chardin & Young, 1931; Te i l h a r d have survived during the Pleistocene, espe- de Chardin 1940; Zheng 1976; Zhang et al. cially in southern Asia where the climate was 1993). This species is possibly an offshoot of more favourable than in Europe. The ocho- the A l i l e p u s lineage. It is characterised by the tonid Ochotonoides complicidens (BO U L E & primitive postcranial morphology and a relati- TE I L H A R D D E CH A R D I N , 1928), which was vely derived structure of the p3, where advan- widespread during the Pliocene was living in ced N e k ro l a g u s -like p3 pattern (with the pos- China at least by the Middle Pleistocene. tero-internal fold closed into the enamel lake) Early Pleistocene remains are known from was present together with the usual A l i l e p u s - Qinghai Province (Zheng et al. 1985), Middle like morphotype (Averianov 1996). Pleistocene remains exclusively from Gansu and Shaanxi Provinces (Chow & Li 1965; The rabbit Pliopentalagus pro g re s s i v u s LI U & Zheng 1976; Xue 1981; Wang 1988). T h e ZH E N G , 1997 is known by two teeth from the l a rge ochotonid with complicated p3 from the Late Pliocene - Early Pleistocene fillings in Late Pleistocene of Sanjiacun, Yu n n a n Henan Province, China (Liu & Zheng 1997).

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The genus P l i o p e n t a l a g u s was characteristic O r y c t o l a g u s for the Ruscinian of Eurasia. This lineage is The genus O ry c t o l a g u s includes two extinct nowadays represented by the relict A m a m i species, O. l a c o s t i (PO M E L , 1853) and rabbit Pentalagus furnessi (STO N E, 1900) O ryctolagus laynensis LO P E Z MA RT I N E Z , 1977 inhabiting two small islands of the Ry u k y u (Fig. 3). The former was distributed in the Archipelago, Japan, where its fossil remains Early Villafranchian - Middle Pleistocene of are known from the Late Pleistocene and France, Spain, and Portugal, the latter is res- Holocene deposits (Tomida et al. 1990; tricted to the early Villafranchian of Spain. Tomida & Otsuka 1993). The distinction between these two species is not clear and they may be synonymous. Another recent relict of an ancient lineage is Fossil remains attributed to the Recent the hispid hare Caprolagus hispidus (PEARSON, European rabbit O ryctolagus cuniculus 1839) distributed along the southern (LI N N A E U S , 1758) are known since the Early Himalayan foothills. It is descendant of the Pleistocene in numerous localities in central Villafranchian C a p rolagus sivalensis and southern Europe and possibly in northern FO R S Y T H MA J O R, 1899 from Pakistan. T h e Africa (Donard 1981; Callou 1995). genus was also represented in the Early Pleistocene of Java by C a p rolagus lapis The European rabbits of the Mindel-Riss (HO O I J E R, 1964), known from isolated teeth epoch are attributed to the subspecies O. c. from Sangiran (Hooijer 1964; Dawson 1971). l u n e l l e n s i s DO N A R D , 1981, those of the Riss From the recent species it differs by a epoch to O. c. g re n a l e n s i s DO N A R D , 1981. somewhat less complicated p3 morphology. The Wurmian European rabbits are referred to the recent subspecies O. c. huxleyi HAECKEL,

Figure 2 A p p r oximate distri bution of the O c h o t o n a species in the Pleistocene of Eura s i a : 1 - O. p u s i l l a,2 - , 3 - O. ru fe s c e n s , 4 - O. a ze ri c a and O. t ra n s c a u c a s i c a,5 - O. d o d o g o l i ca and O. z a s u c h i n i,6 - O. t h i b e t a n a, 7 - O. h y p e r b o r e a,8 - O. k o z l ow i, 9 - O. d a u u ri c a, 1 0 - O. w h a rt o n i.

4 AVERIANOV: Pleistocene lagomorphs

1874 (Donard 1981). All these subspecies Fladerer 1987). All these remains belong to d i ffe r mostly in size. According to the tradi- small-sized hares similar in size with the tional taxonomic works there are from two to recent L. c a p e n s i s . The species attribution of six recent subspecies of O. c u n i c u l u s, all dif- these materials is not clear. fering in size and pelage characteristics. T h e recent craniometric analysis of O. c u n i c u l u s The fossil record of variable hare Lepus timidus showed that variation in size occurred in a LI N N A E U S , 1758 can be traced since the continuous cline, so there is no evidence for Middle Pleistocene (Kurtén 1968; Vo n separation of this species into subspecies Koenigswald 1972). Nowadays it inhabits the (Sharples et al. 1996). However, the mito- forest and Arctic zones of Eurasia. The recent chondrial DNA analysis of 13 rabbit popula- range of L. t i m i d u s d i ffe rs considerably from tions reveals two geographically separated its Pleistocene range. In Western Europe the maternal lineages, the divergence of which variable hare is now restricted to the northern goes back to 2.5 My (Monnerot et al. 1994), latitudes (Fenno-Scandia, Ireland, Scotland) so at least two recent geographic subspecies and to the Alps (subspecies L. t. v a rro n i s ) . of O. c u n i c u l u s could be maintained. This distribution pattern is apparently connected both with the climatic change of the L e p u s Holocene and with the competitionary displa- The reliable Early Pleistocene records of the cing of this species by the brown hare, L. genus L e p u s , with p3 or P2 described, are e u ro p a e u s. During the Pleistocene the distri- known from Moldavia, Ukraine, Russia and bution of L. t i m i d u s was confined to Europe Austria (Sukhov 1976; Shushpanov 1977; (Fig. 4), but since the Holocene its range greatl y

Figure 3 A p p r oximate distri bution of the O r y c t o l a g u s species in the Pleistocene of Eura s i a : 1 - O. l a c o s t i,2 - O. l a y n e n s i s, 3 - O. c u n i c u l u s.

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extended to the east up to the Pacific Ocean; Wurmian 3 (Sartanian) of Yakutia, Eastern this was connected with the disappearance of Siberia (Averianov & Kuzmina 1993; the periglacial steppe and its replacement by Averianov 1995, 1999). These subspecies f o r e s t s . d i ffe r mostly in size.

The Middle Pleistocene European records of The Early Pleistocene Lepus valdarnensis the variable hare can probably be referred to We i t h o f e r, 1889 from Italy was a relatively the subspecies L. t. p r a e t i m i d u s KR E T Z O I i n l a rge hare, possible close to L. e u ro p a e u s a n d J a n o s s y, 1969, which was characterised by L. c a p e n s i s (subgenus E u l a g o s GR AY, 1867), very large size (Janossy 1969). In the Late d i ffe ring from them mostly in less developed Pleistocene remains of L. t i m i d u s are known ability for fast running (it has relatively short- from more than hundred localities in Spain, er distal limb bones). By some authors it was Portugal, France, Belgium, Great Britain, referred to the genus O ry c t o l a g u s and syno- I t a l y, Switzerland, Germany, Austria, Czech, nymised with O. l a c o s t i (PO M E L , 1853). T h e Poland, Yugoslavia, Hungary, Moldavia, wes- species is reliably known only from the Late tern Ukraine and Crimea. Sometimes they are Villafranchian of Italy (Weithofer 1889; found in the great abundance, like in the Bosco 1900; Forteleoni 1974). Madlenian beds of the Kesslerloch cave in Switzerland where remains of at least 1000 The Late Pliocene - Recent cape hare L e p u s individuals were collected (Heierli 1907; c a p e n s i s LI N N A E U S, 1758 inhabits savannahs, Koby 1960). Apparently all continental deserts, steeps, and mountain regions of European records of variable hare could be Africa, Spain, the Near East, and Central referred to the subspecies L. t. w u e r m e n s i s Asia. The recent distribution of L. c a p e n s i s, KO B Y , 1960. The Late Pleistocene variable compared to the Pleistocene range, is restrict- hare from Great Britain was somewhat larg e r ed only in Europe (now it is absent from (Sanford 1869; Hinton 1909) and could be I t a l y, Germany, and Hungary). This may be separated as the subspecies L. t. a n g l i c u s connected with the disappearance of the HI N TO N, 1909. steeps here in the Holocene. It cannot be excluded that Pleistocene remains from Italy The Late Pleistocene L. t. ponticus AVERIANOV, referred to L. c a p e n s i s are actually belonging 1994 is known from several localities on the to L. c o r s i c a n u s D E WI N TO N, 1898, which is Crimea peninsula (Averianov 1994). Late now considered as a distinct species (Palacios Pleistocene records of the variable hare from 1996). Similarly, all Spanish Pleistocene Moldavia can probably be referred to the records of the cape hare could be referable to same subspecies. L. g r a n a t e n s i s RO S E N H A U E R , 1856. The oldest fossil records of L. c a p e n s i s are known from The extinct Late Pleistocene Don hare L e p u s the Upper Pliocene of Eastern Africa. T h i s t a n a i t i c u s GU R E E V , 1964 inhabited the perig- species was reported from more than 20 loca- lacial zone of Eastern Europe and Northern lities in Spain, Italy, Germany, Hungary, nort- Asia. It differs from L. t i m i d u s by the structu- hern Caucasus, Kazakhstan, Middle A s i a , re of dentary and p3 (Averianov 1999). T h e Altai Mountains, southern Western Siberia, species could be divided into three subspe- Transbaikalia and China. Two fossil subspecies: L. t. g m e l i n i AV E R I A N O V & KU Z M I N A, cies of the Cape hare can be currently recog- 1993 from the Wurmian 2/3 of central Russia, nised. L. c. t e r r a e r u b r a e KR E T Z O I, 1956 is a L. t. t a n a i t i c u s GU R E E V , 1964 from the poorly known subspecies to which a few iso- Wurmian 3 of Central Russia, Ukraine, Ural lated teeth from the Middle Pleistocene of Mountains and northern Kazakhstan, and L. t. G e r m a n y, Hungary, Rumania, Italy, and v e re s c h a g i n i AV E R I A N O V, 1995 from the Croatia could be referred (Kormos 1934;

6 AVERIANOV: Pleistocene lagomorphs

Kowalski 1958; Kretzoi 1965; Terzea & n i c u s AV E R I A N O V , 1994 is known from the Jurcsak 1969; Van der Meulen 1973). Some Late Pleistocene and Early Holocene of remains from the Late Pleistocene of China Crimea (Averianov 1994). In the Late were referred to the recent subspecies L. c. Pleistocene L. e u ro p a e u s was quite common t o l a i PA L L A S , 1778 (Jin et al. 1984). in the southern Europe and in Asia. Its remains were reported, but usually not descri- The Early Pleistocene - Recent brown hare bed, from more than 60 localities of Spain, Lepus euro p a e u s PA L L A s, 1778 is distributed France, Italy, Yugoslavia, Bulgaria, Hungary, now in Europe (except for Scandinavia and Moldavia, Crimea and southern Ukraine, Ireland), the Near East, Transcaucasia, and European Russia, the Caucasus, and Kazakhstan. Early Pleistocene records of this Kazakhstan. The Pleistocene records of L. species were reported from the Caucasus e u ro p a e u s do not extend the recent geogra- ( Averianov & Baryshnikov 1993). Middle phic range of the species. Pleistocene remains are known from Germany and Czechoslovakia (Freudenberg The Middle Pleistocene - Recent Manchurian 1914; Schirmeissen 1927; Ziegler 1995). T h e hare Lepus mandshuricus RA D D E , 1861 from relatively numerous remains of the extinct the Russian Far East and north-eastern China subspecies of brown hare, L. e. g u re e v i was reported from some Late Pleistocene GR O M O V , 1952, were described from the Late localities nearby Amur River in Russia and Pleistocene of Binagady in A z e r b a i j a n described from the Middle Pleistocene of (Gromov 1952; Vereschagin 1959; Av e r i a n o v China (Zhang et al. 1993). & Baryshnikov 1993). The extinct L. e. e u x i -

Figure 4 A p p r oximate distri bution of the L e p u s species in the Pleistocene of Eura s i a : 1 - L. t i m i d u s,2 - L. e u r o p a e u s, 3 - L. t a n a i t i c u s, 4 - L. c a p e n s i s,5 - L. o i o s t o l u s, 6 - L. m a n d s h u ri c u s.

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The Early Pleistocene - Recent woolly hare z a s u c h i n i ER B A J E VA, 1988 are known from Lepus oiostolus HO D G S O N, 1840 now inhabits the Early Pleistocene. The former species is the Tibetan Plateau (eastern India, Nepal, and based on relatively numerous material, inclu- China). All remains of this species from the ding skull fragments (Polyakova & Erbaeva Early - Middle Pleistocene of China are pro- 1974), the latter, larg e r-s ized, is based on one bably referrable to the extinct subspecies L. isolated p3 only (Erbaeva 1988). o. w o n g i YO U N G, 1927, which was characterised by its larger size relative to the modern The Pleistocene American W h a r t o n ’s pika subspecies. The remains of this hare are Ochotona whart o n i GU T H R I E & MAT T H E W S , described from a number of Chinese locali- 1971 was reported from the Middle ties (Young 1927, 1930; Colbert & Hooijer Villafranchian of Yakutia (Erbaeva & 1953; Chow & Li 1965; Zhang et al. 1993; Belolyubskii 1993). This species may have Cheng et al. 1995). existed in Siberia during the Pleistocene.

O c h o t o n a A few Recent Eurasiatic pika species have a Another successful group of recent lago- known fossil record in the Pleistocene. T h e morphs is the genus O c h o t o n a, the last repre- northern pika Ochotona hyperbore a (PA L L A S, sentative of the family Ochotonidae. T h e r e 1 8 11) was described from the Middle are several fossil species of O c h o t o n a d e s c r i - Pleistocene of China (Zhang et al. 1993) and bed from the Pleistocene (Fig. 2). the Late Pleistocene of Yakutia (Agadjanian 1972). The Daurian pika Ochotona dauurica Ochotona azerica GA D Z H I E V & AL I E V i n (Pallas, 1776) was reported from some Erbajeva, 1988 is based on the common Middle and Late Pleistocene localities in material from the Middle Pleistocene of China, but the material was not described. A n Azykh Cave in Azerbaijan. From the extinct subspecies, O. d. g u re e v i ER B A J E VA, geographically close species O. ru f e s c e n s i t 1966, is known from the Early Pleistocene of d i ffe rs by a relatively larger anterior segment Transbaikalia (Erbaeva 1988). of p3. Another Caucasian extinct ochotonid is Ochotona transcaucasica (VE K U A , 1967), Two fragmented skulls from the Middle ranging from the Early Pleistocene till probably Pleistocene Zhoukoudian site in China were Holocene (Averianov & Baryshnikov 1993). referred to as O. k o s l o w i and O c h o t o n a c f . It has some similarity with O. ru f e s c e n s ( t h e k o s l o w i ( Young 1934; Pei 1936). A l t h o u g h anterior segment of p3 relatively small, bro- these skulls have some differences from the adly fused to the posterior segment). Recent Kozlov’s Pika O. k o s l o w i (BU E C H N E R, 1894) (see Erbaeva 1988), they share the The type material for Ochotona polonica c o n vex frontals and are similar in size. SY C H , 1980 comes from the Late Ruscinian O. koslowi was reported (but not described) Zamkova Dolna locality in Poland. To this also from the Late Pleistocene of China. species all pikas remains from the Late Pliocene - Early Pleistocene of Poland are The Late Pliocene - Recent steppe pika usually attributed (see review in Wo l s a n Ochotona pusilla (PA L L A S, 1769) was widely 1989, 1990), however, the majority of these distributed in the Pleistocene. Its remains are materials are not described and such determi- known from Great Britain, the Netherlands, nation needs to be established. France, Switzerland, Italy, Yu g o s l a v i a , G e r m a n y, Hungary, Poland, Czechoslovakia, A number of extinct O c h o t o n a species were Austria, Rumania, Ukraine, Russia, and described from Transbaikalia. O c h o t o n a Kazakhstan (it nowadays lives only in the d o d o g o l i c a ER B A J E VA, 1966 and O c h o t o n a steppe zone of Russia and Kazakhstan, from

8 AVERIANOV: Pleistocene lagomorphs

the Volga River till the Ural Mountains). lagus cuniculus and Lepus timidus lived here There are several extinct subspecies of the at least since the Early and Middle steppe pika. O. p. l a z a r i (KR E T Z O I, 1941) is Pleistocene, respectively. Lepus corsicanus known by the holotype only from the Early (Apennines, Corsica) and L. g r a n a t e n s i s Pleistocene locality of Gombasek in (Spain) possibly represent a more ancient Czechoslovakia. The name "O. p. v e t e r i o r" radiation of hares and are now peripherally KR E T Z O I in Janossy, 1969 [nomen nudum] restricted in distribution. In contrast, in A s i a was applied to the material from Solymar, we see an intensive speciation in the popula- Uppony 1, and Tarkü in Hungary (Janossy e t tions of Lepus timidus and Ochotona hyper - a l . 1968; Janossy 1969). O. p. s p e l a e a b o re a , which invaded a huge territory of (OW E N , 1846) was originally described from Siberia in the Holocene, leading to the forma- the Late Pleistocene of Great Britain. tion of many subspecies in Asia and at least Apparently this name could be applied to all one species in North America (the Arctic hare Late Pleistocene steppe pikas from Europe Lepus arcticus ROSS, 1819). A similar intensive (except Crimea and the Caucasus). O. p. speciation process can be expected in the l i u b i n i ER B A J E VA & BA RY S C H N I K O V i n future for the brown hare, which considerably ER B A J E VA, 1988 is known from the type extended its range to the East during past few locality only, Moustierian layers of the centuries. This was connected with the Barakaevskaya Cave in the northern declining forest in this area. Caucasus. O. p. t a n a i t i c a ER B A J E VA, 1988 is known from several Late Pleistocene sites in The recent centre of the species diversity for Crimea (Erbaeva 1988). pikas is located in Tibet and the Himalayas. U n f o r t u n a t e l y, any fossil record for these ter- The fossil remains of the Afghan pika ritories is virtually absent. However, it can be Ochotona ru f e s c e n s (GR AY, 1842) are known hypothesised that intensive speciation of from the Middle Pleistocene sites of Sel’ these mountain pikas took place in the post- Ungur in Kirghisia (Erbaeva 1988) and Pleistocene epoch. During the Pleistocene Emirkaya-2 in Turkey (Montuire et al. 1994). their ancestors apparently lived at lower alti- The Moupin pika Ochotona thibetana tudes. Having become adapted to the cold (MI L N E- E D WA R D S, 1871) was described from Asiatic steppes they had to migrate to higher the Early and Middle Pleistocene of Gansu mountains with the Holocene warming and and Henan Provinces, China (Teilhard de subsequent speciation was connected with the Chardin 1940; Zheng 1976). range fragmentation and geographic isolation by the mountain ridges. This may be the main D I S C U S S I O N reason for the high taxonomic diversity of The Pleistocene epoch was not a crucial Tibetan and Himalayan pikas. This scenario period for the lagomorphs’ h i s t o r y. During looks more suitable than the hypothesis of this epoch the background extinction of some Mitchell (1981), who considered Tibet as the relict taxa, connected with the global cooling, centre of origin for the genus O c h o t o n a . took place. During this time an intensive speciation occurred in phylogenetically young Concluding, we may see that the known fos- groups of lagomorphs: in the genera O c h o - sil record for the Pleistocene lagomorphs is t o n a, O r y c t o l a g u s, and L e p u s in the Old incomplete in many respects and really does World and in the genus S y l v i l a g u s in the New not help much for understanding of the evo- Wo r l d . lutionary history of the Recent species. For the better understanding of this history we The Recent fauna of lagomorphs in Europe need more Pleistocene records, more nume- has, in general, a relict appearance. O r y c t o - rous fossil samples, we need detailed descrip-

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tions of all relevant materials and especially Villafranchian of Villafranca d' Asti (Italy). we need the study of skull and dentition Preliminary note - Giornale di Geologia 35 (1): 1-16 variation in the Recent and fossil lagomorph Bosco, A., 1900 - I roditori pliocenici del Val d'Arno populations. This will allow us to make more superiore - Palaeontographica Italica 5: 84-104 precise determination of the fossil materials Cai, B., 1989 - Fossil Lagomorphs from the late Pliocene at the species and subspecies levels and to of Yangyuan and Yuxian, Hebei - Vertebrata reconstruct the real history of the Recent spe- Palasiatica 27 (3): 170-181 c i e s . Callou, C., 1995 - Modifications de l’aire de répartition du lapin (O ryctolagus cuniculus) en France et en AC K N OW L E D G E M E N T S Espagne, du Pléistocène a l’Époque actuelle. État de I am grateful to Dr. Andrew Currant for the la question - Anthropozoologica 21: 95-11 4 help in studying collections in the British C h o w, M. & Li, C.-k., 1965 - Mammalian fossils in Museum (Natural History), London. My work association with the mandible of Lantian man at was supported by the Russian Fund of Basic Chen-Chia-Ou, in Lantian, Shensi - Vertebrata Research (RFBR) grant 00-15-99355. Palasiatica 9 (4): 377-394 Colbert, E.H. & Hooijer, D.A., 1953 - Pleistocene R E F E R E N C E S mammals from the limestone fissures of Szechwan, Agadjanian, A.K., 1972 - Rodents from the Pleistocene China - Bulletin of the American Museum of Natural deposits of Mamontovaya Gora - in: Theriofauna of History 102: 1-134 the Pleistocene - pp. 24-69. Moscow State University, Cheng, J., Cao, B., Tian, M. & Li, L., 1995 - A new early Moscow [In Russian] Pleistocene Mammalian fauna from Zhoukoudian - Av e r i a n o v, A.O., 1994 - Pleistocene hares (L e p u s, Earth Science (Wuhan) 20 (5): 497-504 L a g o m o r p h a ) of Crimea - Trudy Zoologicheskogo Dawson, M.R., 1971 - Fossil mammals of Java. I. Notes Instituta RAN 256: 69-91 [In Russian] on Quaternary Leporidae (Mammalia, Lagomorpha) Av e r i a n o v, A.O., 1995 - Late Pleistocene hare, L e p u s from central Java - Proceedings of the Koninklijke t a n a i t i c u s (Lagomorpha, Leporidae) of Siberia - Nederlandse Akademie van Wetenschappen, Series B Trudy Zoologicheskogo Instituta RAN 263: 121-162 74 (1): 27-32 [In Russian] Donard, E., 1981 - O r yctolagus cuniculus dans quelques Av e r i a n o v, A.O., 1996 - On the systematic position of gisements quaternaires francais - Quaternaria 23: the rabbit ‘C a p ro l a g u s ’ b r a c h y p u s Young, 1927 1 4 5 - 1 5 7 (Lagomorpha, Leporidae) from the Villafranchian of Erbaeva, M.A., 1988 - Pikas of the Cenozoic (taxonomy, China - Trudy Zoologicheskogo Instituta RAN 270: systematics, phylogeny) - Nauka, Moscow: 222 pp. 148-157 [In Russian] [In Russian] Av e r i a n o v, A.O., 1999 - Late Pleistocene hares (L e p u s ) Erbaeva, M.A. & Belolyubskii, I.N., 1993 - First record of the Russian Plain - in: Saunders, J.J., Styles, B.W. of the W h a r t o n ’s pika on the Kolyma lowland - & Baryshnikov, G.F. (eds.) - Quaternary Geologiya i Geofizika 34 (6): 142-144 [In Russian] Paleozoology in the Northern Hemisphere - pp. 33- F l a d e r e r, F.A., 1987 - Beitrag zur Entwicklung von 61. Illinois State Museum Scientific Papers, H y p o l a g u s und L e p u s (Lagomorpha, Mammalia) im S p r i n g f i e l d Pliopleistozän von Mitteleuropa - Sitzungsberichten Av e r i a n o v, A.O. & Baryshnikov, G.F., 1993 - Pleistocene der österreichischen Akademie der Wissenschaften, hares (genus L e p u s, Lagomorpha) of Great Caucasus Mathematisch-naturwissenschaftliche Klasse, - Trudy Zoologicheskogo Instituta RAN 246: 4-28 Abteilung 1 196 (1-4): 123-138 [In Russian] F l a d e r e r, F.A. & Reiner, G., 1996 - Evolutionary shifts in Av e r i a n o v, A.O. & Kuzmina, I.E., 1993 - The Don hare, the first premolar pattern of Hypolagus beremendensis Lepus tanaiticus G u r e e v, 1964 from the Paleolithic (Petényi, 1864) (Lagomorpha, Mammalia) in the Kostenki sites - Trudy Zoologicheskogo Instituta Plio-Pleistocene of Central Europe - Acta zoologica RAN 249: 66-92 [In Russian] Cracoviensia 39 (1): 147-160 Berzi, A., 1967 - Lagomorphs from the type Fondi, R., 1970 - P rolagus sard u s Wagner (Ochotonidae,

10 AVERIANOV: Pleistocene lagomorphs

Lagomorpha, Mammalia) de una breccia ossifera Kretzoi, M., 1965 - Die Nager und Lagomorphen von della Montagnola senese - Atti della Societa Toscana Voigstedt in Thüringen und ihre chronologische di Scienze Naturali, Serie A 77: 260-288 Aussage - Paläontologische Abhandlungen, A b t e i l u n g Fondi, R., 1972 - Fauna cromeriana della Montagnola 4: Paläozoologie 2 (2/3): 585-660 senese - Palaeontographia Italica 68: 1-27 Kurtén, B., 1968 - Pleistocene Mammals of Europe - Forteleoni, G., 1974 - The Upper Valdarno lagomorph, Aldine Publishing Company, Chicago: 317 pp. Lepus valdarnensis Weithofer - Palaeontographia Liu, L. & Zheng, S., 1997 - Note on the late Cenozoic Italica 67: 55-68 lagomorphs of Danjiang Reservoir area in Hubei and F r e u d e n b e rg, W., 1914 - Die Säugetiere des älteren Henan - Vertebrata Palasiatica 35 (2): 130-144 Quartars von Mitteleuropa - Geologische und Lopez Martinez, N., 1989 - Revision sistematica y Paläontologische Abhandlungen Jena, Neue Folge 12 biostratigrafica de los lagomorpha (Mammalia) del (4/5): 455-670 Terciario y Cuaternario de Espana - Memorias del G r o m o v, I.M., 1952 - Rodent fauna (Rodentia) of the Museo Paleontologico de la Universidad de Zaragosa Binagady Pleistocene and its nature - Trudy 3 (3): 1-350 Estestvenno-Istoricheskogo Museya imeni G.Zardabi Lopez Martinez, N. & T h a l e r, L., 1975 - Biogéographie , 5 (2): 203-349 [In Russian] évolution et compléments à la systématique du Heierli, J., 1907 - Das Kesslerloch bei Thaingen - Neue groupe d’Ochotoinides P i e z o d u s - P ro l a g u s Denkschriften der allgemeinen schweizerischen (Mammalia, Lagomorpha) - Bulletin de la Societé Gesellschaft für die gesammten Naturwissenschaften G é o l o g i q u e de France, 7serie 17(5): 850-866 43: 1-214 Mitchell, R.M., 1981: The O c h o t o n a (Lagomorpha: Hinton, M.A.C., 1909 - On the fossil hare of the Ochotonidae) of Asia - in: Proceedings of ossiferous fissures on Ightham, Kent, and on the Symposium on Qinghai-Xizang (Tibet) Plateau recent hares of the Lepus variabilis group - Scientific (Beijing, China) 2. Geological and ecological studies Proceedings of the Royal Dublin Society, New Series of Qinghai-Xizang Plateau - pp. 1031-1038. Science 12: 225-265 Press and Gordon and Breach, Beijing, New Yo r k H o o i j e r, D.A., 1964 - New records of mammals from the Monnerot, M., Casane, D., Hardy, C. & Vigne, J.-D., Middle Pleistocene of Sangiran, central Java - 1994 - Evolution of O r y c t o l a g u s : relationships within Zoologische Medelingen 40 (10): 73-88 the Lagomorpha system and access to population J a n o s s y, D., 1969 - Stratigraphische Auswertung der history by ancient DNA - Polish Evolutionary Studies europaischen mittelpleistozanen Wirbeltierfauna. Teil 20 (3-4): 543-551 II - Berichte deutschen Gesellschaft geologische Montuire, S., Sen, S. & Michaux, J., 1994 - The Middle Wissenschaften, Serie A 14 (5): 573-643 Pleistocene mammalian fauna from Emirkaya-2, J a n o s s y, D., Krolopp, E. & Brunnacker, K., 1968 - Die Central Anatolia (Turkey): systematics and Felsnische Uppony I (Nordungarn) - Eiszeitalter und paleoenvironment - Neues Jahrbuch für Geologie und Gegenwart 19: 31-47 Paläontologie, Abhandlungen 193(1): 107-144 Jin, C., Xu, Q. & Li, C., 1984 - The Quaternary Palacios, F., 1996 - Systematics of the indigenous hares mammalian faunas from Qingshantou site, Jilin of Italy traditionally identified as Lepus euro p a e u s Province - Vertebrata Palasiatica 22 (4): 314-323 Pallas, 1778 (Mammalia: Leporidae) - Bonner K o b y, F.E., 1960 - Sur l’extension maximale vers le zoologische Beiträge 46 (1-4): 59-91 sud-ouest de quelques representants de la faune Pei, W.C., 1936 - On the mammalian remains from froide würmienne - Antropos, Mammalia, locality 3 at Choukoutien - Palaeontologica Sinica, pleistocenica. Brno, Moravské Museum 1: 101-108 Series C 7 (5): 1-108 Kormos, T., 1934 - Zur Frage der Abstammung Polyakova, R.S. & Erbaeva, M.A., 1974 - On the eurasiatischer Hasen - Allatani közlemenyek 31: d i ffer ences in the skeletons of the extinct Dodogol 6 5 - 7 8 and Recent Daurian pikas - Trudy Zoologicheskogo Kowalski, K., 1958 - Altpleistozäne Kleinsäugetierfauna Instituta AN SSSR 54: 180-189 [In Russian] von Podumci in Norddalmatien - Palaeontologia Qiu, Z., Li, C. & Hu, S., 1984 - Late Pleistocene Jugoslavica 2: 5-30 micromammal fauna of Sanjiacun, Kunming -

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Vertebrata Palasiatica 22 (4): 281-293 National Science Museum [Tokyo] 23: 173-183 [In Rook, L. & Masini, F., 1990 - P ro l a g u s from the Upper J a p a n e s e ] Valdarno (faunal assotiations of the Olivola and Ta s s o Van der Meulen, A.J., 1973 - Middle Pleistocene smaller Units, late Villafranchian) - Bollettino della Societa mammals from the Monte Peglia (Orvieto, Italy), Paleontologica Italiana 29 (3): 347-360 with special reference to the phylogeny of M i c ro t u s Sanford, W.A., 1869 - On the Rodentia of the Somerset (Arvicolidae, Rodentia) - Quaternaria 17: 1-144 caves - Quarterly Journal of the Geological Society Vereschagin, N.K., 1959 - Mammals of Caucasus. of London 26: 124-131 History of the fauna development - Izdatel’stvo AN Schirmeissen, K., 1927 - Altdiluviale Mahlzeitreste aus SSSR, Moscow, Leningrad: 703 pp [In Russian]. dem Lateiner Berge bei Brünn - Verhandlungen des Von Koenigswald, W., 1972 - Sudmer- B e rg 2, eine Fauna naturforschenden Vereines in Brünn 60: 29-51 des frühen Mittelpleistozäns aus dem Harz - Neues Sharples, C.M., Fa, J.E. & Bell, D.J., 1996 - Jahrbuch fur Geologie und Paläntologie, Geographical variation in size in the European rabbit Abhandlungen 141 (2): 194-221 O r yctolagus cuniculus (Lagomorpha: Leporidae) in Wang, H., 1988 - Early Pleistocene mammalian fauna western Europe and North Africa - Zoological Journal from Dali, Shaanxi - Vertebrata Palasiatica 26 (1): of the Linnean Society 117: 141-158 5 9 - 7 2 S h u s h p a n o v, K.I., 1977 - Hares remains (genera We i t h o f e r, A., 1889 - Ueber die tertiären P l i o l a g u s and L e p u s) from the Upper Pliocene of the Landsäugethiere Italiens - Jahrbuch der Kaiserlich- southern Moldavia and Ukraine - Izvestiya AN Königlichen Geologischen Reichsanstalt, Wien 39: Moldavskoi SSR, Seriya biologiya i khimiya 6: 59-63 5 5 - 8 2 [In Russian] Wolsan, M., 1989 - Zajeczaki - Lagomorpha - Folia S u k h o v, V. P., 1976 - Small mammals of the Tiraspol Quaternaria 59-60: 145-150 faunistic assemblage of the Belaya River (on the Wolsan, M., 1990 - Lower Pleistocene Lagomorphs of Chui-Atasevo section) - in: Questions of stratigraphy Poland - Quartärpaläontologie 8: 273-276 and correlation of the Pliocene and Pleistocene Xue, X., 1981 - An early Pleistocene mammalian fauna deposits of northern and southern parts of Predural’e and its stratigraphy of the River You, Weinan, Shensi - pp. 4-40. Izdatel’stvo Bashkirskogo filiala AN - Vertebrata Palasiatica 19 (1): 35-44 SSSR, Ufa [In Russian] Young, C.C., 1927 - Fossile Nagetiere aus Nord-China - Teilhard de Chardin, P., 1940 - The fossils from Locality Palaeontologia Sinica, Series C 5 (3): 1-82 18 near Peking - Palaeontologia Sinica, New Series Young, C.C., 1930 - On the mammalian remains from (C) 9 (124): 1-94 Chi Ku Shan near Chou Kou Tien - Palaeontologia Teilhard de Chardin, P. & Young, C.C., 1931 - Fossil Sinica, Series C 7 (1): 1-19 mammals from the late Cenozoic of Northern China - Young, C.C., 1934 - On the Insectivora, Rodentia and Palaeontologia Sinica, Series C 9 (1): 1-67 Primates other than Sinanthropus from Locality 1 at Terzea, E. & Jurcsak, T., 1969 - Contributii la cunoaste Choukoutien - Palaeontologia Sinica, Series C 8 (3): rea faunelor pleistocene medii de la Betfia (Romania) 9 - 1 3 9 - Lucrârile Institutuli de Spéologie ‘Emile Racovitza’ Young, C.C., 1935 - Miscellaneous mammalian fossils 8: 201-213. from Shansi and Honan - Palaeontologia Sinica, Tomida, Y. & Otsuka, H., 1993 - First discovery of fossil Series C 9 (2): 1-42 amami rabbit (Pentalagus furnessi) from Zhang, S. et al. [other authors not indicated], 1993 - Tokunoshima, Southwestern Japan - Bulletin of the Comprehensive study on the Jinniushan Paleolithic National Science Museum [Tokyo], Series C 19 (2): site - Memoires of the Institute of Vertebrate 7 3 - 7 9 Paleontology and Palaeoanthropology, Academia Tomida, Y., Otsuka, H., Ueno, T., Hajime, S. & Baba, H., Sinica 19: 1-163 1990 - A synopsis of a vertebrate paleontological and Zheng, S., 1976 - Small Mammals of Middle Pleistocene paleoanthropological survey of the island of Tokuno in Heshui, Gansu - Vertebrata Palasiatica 14 (2): 11 2 - and the big island of Amami - Memoirs of the 11 9

12 AVERIANOV: Pleistocene lagomorphs

Zheng, S., Wu, W., Li, Y. & Wang G., 1985 - Late Cenozoic mammalian faunas of Guide and Gonghe Basins, Qinghai Province - Vertebrata Palasiatica 23 (2): 89-134 Z i e g l e r, R., 1995 - Pleistozäne Säugetierfaunen von Genkingen bei Reutlingen (Baden-Württemberg) - Stuttgarter Beiträge zur Naturkunde, Series B 234: 1- 4 3 Z u b a k o v, V.A., 1986 - Global climatic events of the Pleistocene - Gidrometeoizdat, Leningrad: 288 pp [In Russian].

re c e i ved 09 Ju ly 1999 accepted 27 December 1999

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DEINSEA - ANNUAL OF THE NATURAL HISTORY MUSEUM ROTTERDAM P. O . B o x 2 3 4 5 2 , N L - 3 0 0 1 K L R o t t e r d a m T h e N e t h e r l a n d s