Lagomorpha: Leporidae) Fostowicz-Frelik, L., 2003 - Species Distribution and Differentiation of Eurasian Hypolagus (Lagomorpha: Leporidae) - In: Reumer, J.W.F

Lucja Fostowicz-Frelik Institute of Paleobiology, Polish Academy of Sciences

Species distribution and differentiation of Eurasian Hypolagus (Lagomorpha: Leporidae) Fostowicz-Frelik, L., 2003 - Species distribution and differentiation of Eurasian Hypolagus (Lagomorpha: Leporidae) - in: Reumer, J.W.F. & Wessels, W. (eds.) - DISTRIBUTION AND MIGRA- TION OF TERTIARY MAMMALS IN EURASIA. A VOLUME IN HONOUR OF HANS DE BRUIJN - DEINSEA 10: 197-216 [ISSN 0923-9308] Published 1 December 2003

Features of p3 (width, length, AER and PER deepth, and AER-factor and PER-factor), as well as of mandible (diastema length, lower tooth row length, and mandible height) of five Eurasian Hypolagus species were analysed. The "beremendensis-igromovi" morphological group is distin- guished on the basis of similar dimensions and proportions of mandible and teeth. The Pleistocene Central European populations of Hypolagus beremendensis, characterised by larger dimensions, resemble Hypolagus igromovi (Don Region). The division of Hypolagus beremendensis into two, previously proposed, subspecies is maintained. H. beremendensis beremendensis existed from Late Ruscinian to Late Villányian and H. beremendensis brachygnathus was known from Late Villányian to Steinheimian (Q3). Similarity between Hypolagus b. brachygnathus resemble Hypolagus igromovi could be the result of either parallel evolution or closer relationships. Thus, it is possible that the significantly larger, Late Villányian and Early Pleistocene populations of Hypolagus beremendensis are the descendants of Hypolagus igromovi, which invaded Central Europe in the Villányian. Hypolagus schreuderae, described from the Nihewanian deposits, is characterised by a long diastema, a simple enamel pattern and forms a distinct species.

Correspondence: Institute of Paleobiology, Polish Academy of Sciences; Twarda 51/55; 00-818 Warszawa, Poland, e-mail: [email protected]

Keywords: Hypolagus, Lagomorpha, Evolution, Taxonomy, Pliocene, Pleistocene, Turolian, Biharian, Nihewanian, Eurasia

INTRODUCTION land bridge (Gureev 1964; Qiu 1987, Chaline Leporids are known in Asia from the Middle et al. 2000). There are approximately 16 Eocene (McKenna & Bell 1997). During the Hypolagus species known up to now, of Late Oligocene to Late Miocene interval which five (or six) were described from there has been a significant decrease of lepo- Eurasia. Hypolagus became widely distribu- rid abundance in some regions of Asia, espe- ted and abundant in the Pliocene and Early cially in China, where after the Late Pleistocene of Eurasia. The numerous karst Oligocene, and up to the latest Miocene, sites included hundreds of Hypolagus bones leporids have been missing from the fossil and teeth known from Central Europe, chiefly record (Qiu 1987). Hypolagus DICE, 1917 Poland and Hungary (Sulimski 1964; Sych was one of the newcomers, which originated 1965; Jánossy 1986; Wolsan 1989). in North America in the Early Miocene Hypolagus co-occurred in sediments with (Hemingfordian) and migrated to Asia proba- remains of other leporids such as Alilepus bly before the Late Miocene through Beringia DICE, 1931 or Lepus LINNAEUS, 1758. Its dis-

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tribution area bordered and sometimes over- Sokal 1973) was employed for testing diaste- lapped the distribution area of Trischizolagus ma length as a possible specific discrimina- RADULESCO & SAMSON, 1967, Serengetilagus tor. His coefficient of discrimination: DIETRICH, 1941, and Oryctolagus LILLJEBORG, 2 2 1873 (Kurtén 1968; De Bruijn et al. 1970; K= (XiA-XiB) /2 s i Qiu 1987; Sen & Erbajeva 1995; Averianov 2 & Tesakov 1997; Chaline et al. 2000). where s i is the pooled variance for character This study tries to summarise the knowled- i from taxa (here: populations) A and B. The ge on Hypolagus species in Eurasia, basing greater K is, the better i is as a discriminator on the analysis of dimensions and proportions (e.g. K= 7.68 allows 95 % of identifications (metrical characters) of mandibles and lower to be correct). premolars – p3, frequently used in taxonomic investigations on American leporids, also Material used including Hypolagus (White 1991; White & Morgan 1995; Voorhies & Timperley 1997). Hypolagus beremendensis (PETÉNYI, 1864) Localities: Poland, Weze 1: 123 p3, 30 man- MATERIAL AND METHODS dible fragments; Zamkowa Dolna Cave, fauna A: 9 p3; Rebielice Królewskie 1: 27 p3 The material, consisting of p3 and mandibles and 7 mandible fragments, Rebielice of Eurasian Hypolagus species, is listed Królewskie 2: 18 p3; Kadzielnia: 55 p3, 8 below by species, with number of specimens mandible fragments; Kamyk: 136 p3, 10 in brackets. The specimens come from 19 mandible fragments (all coll. ISEZ). Hungary, localities ranging chronologically from the Osztramos 7: 18 mandible fragments (coll. very end of the Miocene to Middle Pleisto- NHMB); Villány 3: 8 p3, 4 mandible frag- cene (Fig. 1). The analysis was based on ments (coll. HGIB). Czech Republic, Chlum: camera lucida drawings of the occlusal sur- 2 p3; Holstejn: 4 p3 (Fladerer & Reiner 1996: face of the p3, from which the measurements fig. 7: 3–5, 7–9). Austria, Deutsch-Altenburg were taken. Three measurements were taken 2C1: 9 p3 and 16 mandibles, and Deutsch- from the mandibular bones: the length of the Altenburg 4B: 5 (Fladerer & Reiner 1996: diastema, the length of the lower tooth row, fig. 7: 10–18, Tab. IV). and the height of the dentary between the p4 and m1. All material used belonged to the Hypolagus igromovi GUREEV, 1964 adult specimens. The teeth were selected Localities: Russia, Rasdorskaya, right bank of having the parallel, not tapering, walls (in Don River, Rostov Region: 8 p3, measure- lateral view), which is typical for adult indi- ments of 9 mandible fragments (Gureev viduals. The anatomical structure of p3 and 1964; Averianov 1996 a: fig. 1, A-F, I-J, Tab. scheme of tooth measurements are shown in I), Kosyakino, Northern Caucasus: 1 p3 Figure 2. The anatomical nomenclature and (Averianov & Tesakov 1998: fig. 1: j). indices after White (1991) and Voorhies & Timperley (1997). Hypolagus multiplicatus (ERBAJEVA IN In order to assess the similarity between the BAZAROV et al., 1976) studied populations, average Euclidean dis- Localities: Russia, Udunga, Western tances were calculated on the basis of seven Transbaikalia: 1 p3 (Erbajeva 1996: fig 2 A); metrical characters (Table 1) for six Beregovaya: 1 p3 (Erbajeva 1996: fig 3 A). Hypolagus populations. The phenogram by unweighted pair-group method using arithme- Hypolagus schreuderae TEILHARD DE tic averages (UPGMA) was then constructed. CHARDIN, 1940 The method of Lubischev (after: Sneath & Localities: China, Inner Mongolia, Loc. 18

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Figure 1 Stratigraphic record of some Hypolagus sites bearings from Eurasia, used to the study.The stratigraphic correlation after Qiu (1987), Kretzoi (1987), Qiu & Storch (2000). near Beijing: 4 p3, measurements of 5 mandi- Hypolagus transbaicalicus (ERBAJEVA, in bles (Teilhard de Chardin 1940: fig. 22, BAZAROV et al. 1976) measurements, p. 38; Cai 1989: fig. 1 d); Localities: Russia, Udunga, Western Nihewan: 3 p3 (Cai 1989: fig. 1 a–c); Yushe: Transbaikalia: 1 p3 (Erbajeva 1996: fig. 2 B); 2 p3 (Cai 1989: fig. 1 e–f). Beregovaya: 3 p3 (Bazarov et al. 1976: Pl II, 5, 6; Erbajeva 1996: fig. 3 B) Hypolagus sp. indet. Harr Obo: 2 p3 (Qiu 1987: Text fig. 32; Cai 1989: fig. 1g)

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WHAT IS HYPOLAGUS? Hypolagus is recognised as a genus of fossil rabbit, assigned to the problematic subfamily Archaeolaginae, including two genera intermediate in p3 morphology between Palaeo- laginae and Leporinae (Dice 1929; Dawson 1958). Hypolagus was presumed to be derived from Archaeolagus in the Early Miocene and become widely distributed in North Figure 2 Diagrams of p3: (a) with labelled teeth structures, America and Eurasia (Dawson 1958; 1967; AER – anterior external reentrant, AR – anterior reentrant, PER – posterior external reentrant, PIR – posterior internal Sych 1965). reentrant; and (b) indicating measurements of the occlusal The original diagnosis given by Dice surface, A – transverse width, B – length, C – PER deep, (1917) states that in Hypolagus "p3 has two D – AER deep. anterior re-entrant angles, neither of which extends over half way across the tooth. The Abbreviations and indices posterior angle is much deeper than the anterior. There are no grooves on the inner sur- Sites: Bt – Betfia, DA – Deutsch-Altenburg, face of the tooth. The upper molar has the re- H – Chlum 6 and Holstejn, KA – Kamyk, entrant angle extending about half way across KD1 – Kadzielnia, Os7 – Osztramos 7, RK1 the crown and the enamel in the angle is – Rebielice Królewskie 1, RK2 – Rebielice coarsely folded. Cement is well developed Królewskie 2, V3 – Villány 3, W1 – Weze 1, (…)", and "p3 in Hypolagus differs from the ZDA – Zamkowa Dolna Cave. corresponding tooth in Archaeolagus in being proportionally shorter and broader, although ISEZ: Institute of Systematics and Evolution the enamel pattern is very similar in the two of Animals, Polish Academy of Sciences, genera." This generic diagnosis holds true for Kraków, Poland the most Hypolagus species and specimens as HGIB: Hungarian Geological Institute, a whole. It distinguishes Hypolagus from Budapest, Hungary most of Serengetilagus, Trischizolagus or NHMB: Geological and Paleontological Alilepus species. However, in some species of Department of the Natural History Museum, Hypolagus there are shallow crinkles in the Budapest, Hungary position of the anterior (Hypolagus igromovi, PIN: Paleontological Institute, Russian Hypolagus brachygnathus sensu Fladerer Academy of Sciences, Moscow, Russia 1987) and antero-internal (Hypolagus brachygnathus sensu Fladerer 1987) reentrants ZIN: Zoological Institute, Russian Academy (Fladerer 1987; Averianov 1996 a; Fladerer & of Sciences, St. Petersburg, Russia. Reiner 1996). The crinkles are usually very AER: anterior external reentrant. shallow and could be hardly described as re- PER: posterior external reentrant. entrants, comparing with these observed in AER-factor: AER deep x 100% / width of the Serengetilagus, Trischizolagus or Alilepus. p3 crown On the other hand, in Trischizolagus, PER-factor: PER deep x 100% / width of the Serengetilagus, and some Lepus species there p3 crown are some of p3 teeth (respectively ca. 14.0, Md-ratio 1: diastema length x 100% / lower 3.5, and 0.07 %) showing Hypolagus-like tooth row length crown shape (Erbajeva & Angerman 1983; Md-ratio 2: height of the mandible x 100%/ Averianov & Tesakov 1997; Qiu & Storch diastema length 2000). It is also noteworthy that juvenile specimens of Hypolagus beremendensis (pre-

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viously described as Pliolagus KORMOS, River sediments were described mainly based 1934) frequently display relatively deep ante- on the teeth, on mandibular and maxillar ro-internal re-entrant, which vanishes during features, with minor attention given to the ontogeny (Sych 1965; Averianov & Tesakov postcranial skeleton (Teilhard de Chardin 1997). 1940; Gureev 1964; Bazarov et al. 1976; Cai The skull and postcranial skeleton of 1987, 1989; Qiu 1987; Averianov 1996a; Hypolagus show the conservative and uni- Erbajeva 1996; Qiu & Storch 2000). The best form plan for most fossil leporids (Dawson known Eurasian species appears to be 1967; Lopez-Martinez 1985), the significant Hypolagus beremendensis, which inhabited feature is a short and robust muzzle, caused most of Central and East Europe from Early by shortening of diastema. Limb bones Pliocene to Middle Pleistocene. The investi- express cursorial and fossorial ability inter- gations on numerous material from Poland, mediate between Oryctolagus and Lepus, published by Sych (1965), contributed much with visible tendencies toward increasing cur- to the knowledge about the genus. soriality, especially in the Pleistocene popula- However, Sych did not specify the taxono- tions (Fladerer 1984; Fladerer & Reiner mic features characteristic for Hypolagus 1996; Fostowicz-Frelik 2001). beremendensis in comparison with other Hypolagus species, apart from the relative THE OUTLINE OF HYPOLAGUS length of intermaxilla in the relation to the TAXONOMY nasals, which cannot be regarded as diagnos- The investigations about leporids frequently tic because of its high interspecific variability treated the structure of dentition as the gener- (observed in other leporid genus e. g. Lepus). al diagnostic characteristic (Dice 1929; He was the first author to deal with the pro- Hibbard 1963; Radulesco & Samson 1967; blem of Hypolagus evolution, and put for- White 1991; White & Morgan 1995, Voorhies ward a model about gradual size increase, & Timperley 1997). It is mainly because of conforming to Bergmann's rule, presumably the teeth and mandibular fragments abundan- caused by the climatic cooling at the begin- ce, while the complete skulls and postcranial ning of the Pleistocene. Sych (1965) also bones are relatively scarce in the fossil record revised Pliolagus genus, recognising the spe- (Dawson 1967). Another reason is, that the cimens as juvenile Hypolagus. first premolar teeth in rows – p3 and P2, are Another progress in taxonomy of the particularly useful in studies on the leporid European Hypolagus was the study on p3 and evolution (Fladerer 1987), being the most P2 enamel pattern, using the morphotypes exposed on forces and abrasion during che- concept (Fladerer 1987; Fladerer & Reiner wing activity, and therefore quickly changing 1996). The problem with Hypolagus taxono- in response to vegetation and climatic chan- my, as well as other leporid genera, seems to ges. There are some differences in the post- be in the high intraspecific and ontogenic cranials of different known leporid genera, variability. This, in turn, demands comparing mainly in the limb bones proportions, and the numerous samples of specimens, and thus, cursorial ability (Sych 1965; Campbell 1969; most of the derived features characterise only Fladerer 1984; Fostowicz-Frelik 2001), the population as a whole, which makes it however, they are not very helpful in the dis- hardly useful in case of small samples. crimination between species on intrageneric level (Averianov 1996a; Averianov and RESULTS Tesakov 1997). Chinese Hypolagus schreuderae, as well as Transbaikalian Hypolagus Mandibular characters multiplicatus, Hypolagus transbaicalicus, and Three mandibular dimensions were analysed: European Hypolagus igromovi from Don the length of diastema, length of the lower

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Figure 3 Relationships between diastema length and lower tooth row length (Md-ratio 1), for various Eurasian Hypolagus species and populations (centroids). Symbols as follows: ˼ - Hypolagus beremendensis-igromovi group (white – Pliocene populations, gray – the earliest Villánian-Pleistocene populations, black – H. igromovi), ̆ - H. schreuderae (Nihewanian).

Figure 4 Relationships between diastema length and mandible height (Md-ratio 2) in different Eurasian Hypolagus species (centroids). Symbols as in Figure 3.

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tooth row, and the height of the mandibular P < 0.02, Student’s t-test, for comparison bone, measured between p4 and m1 (Table between H. schreuderae and H. igromovi). 1). The most discriminating dimension is the The height of the mandible is much less diastema length. The longest diastema is variable factor, and its values are very close observed in Hypolagus schreuderae and its for all species analysed. There is a tendency length seems to have the highest value among towards elongation of the mandible, while its most of Hypolagus species, closely compara- height remains virtually unchanged in the ble with that of Oryctolagus cuniculus Pleistocene Hypolagus beremendensis popu- (Hibbard 1969; White 1991, White & Morgan lations ( Kamyk, Deutsch-Altenburg, Betfia; 1995; Voorhies & Timperley 1997). This Fladerer & Reiner 1996), and in Hypolagus feature allows to distinguish two major igromovi. groups: a monospecific "schreuderae" group and a "beremendensis–igromovi" group, p3 characters including all Central European populations Four metrical characters of p3 (length, width, (Figs. 3, 4). The Md-ratio 1 values for this and depth of PER and AER) were measured dimension for Kamyk population were taken in the studied species (Table 1). The most from Sych (1965) because of the poor mate- discriminating feature seems to be the AER rial preservation. The Md-ratio 1 seems to be depth. The largest values of each measure- highly statistically significant feature (0.01< ment are observed in Hypolagus igromovi

Figure 5 Relationships between AER and PER depth and the p3 length (centroids). Symbols as in Figure 3, additionally: ̅ - Hypolagus sp. of Harr Obo (Early Pliocene).

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and Pleistocene populations of Hypolagus generally uniform in all species. The most beremendensis, which are grouped with the square-shaped p3 are observed in Hypolagus former, and are distinct from the smaller transbaicalicus and Hypolagus multiplicatus, Hypolagus schreuderae and Pliocene popula- with W/L index 97 % and 99 %, respectively. tions of Hypolagus beremendensis (Fig. 5, 6). The PER and AER-factors show the relations The middle position is occupied by Kamyk between the re-entrant depth and the width of and Villány 3 populations, which display the p3 crown. The highest values of both fac- relatively small p3 width and length, but a tors are observed in H. beremendensis from significant increase in re-entrant depth. The Kamyk and in Hypolagus igromovi. The dis- width to length ratio (W/L) is similar and tinctively lowest values of both factors are

Figure 6 Comparison of the AER and PER-factors distribution among Hypolagus species, expressed in percentages; observed range in white, standard deviations in black (values for North American species from White 1991;White & Morgan 1995; Voorhies & Timperley 1997).

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observed in the Nihewanian populations of group into two subgroups. The first of them, Hypolagus schreuderae and of the AER-fac- characterised by a lower value of the AER- tor in the Hypolagus transbaicalicus and factor, includes all Pliocene populations of Hypolagus multiplicatus (Fig. 7). Hypolagus beremendensis. The second one, Significantly, the Early Pliocene Chinese with a higher AER-factor value, groups specimen from Harr Obo, presumed to belong Pleistocene Hypolagus beremendensis popu- to Hypolagus schreuderae, displays dimen- lations together with material from Villány 3, sions very similar to those of the Pliocene with Early Pliocene Hypolagus sp. from Harr Hypolagus beremendensis, but proportions Obo (China, Inner Mongolia), and with similar to the Pleistocene populations. Hypolagus igromovi (Fig. 7). Thus, there is a Nihewanian Hypolagus schreuderae forms an visible morphological trend in Hypolagus obviously distinct unit (Fig. 5, 7). beremendensis to deepening the antero- The AER-factor is the most discriminating, external fold, which started probably in the dividing the "beremendensis - igromovi" very Early Pleistocene. A rather unexpected

Figure 7 Relationships between PER and AER-factors for different species of Eurasian Hypolagus (centroids). Symbols as in Figures 3 and 5. Additionally: ˿ - Hypolagus transbaicalicus, ࡯ - Hypolagus multiplicatus.

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divergent tendency is observed in Chinese Choanae broader than in Oryctolagus cunicu- populations (Cai 1989: Fig. 2). lus and narrower than observed in Lepus. The outline of p3 in H. b. beremendensis clearly CLASSIFICATION triangular, with narrow pointed anterior part of the trigonid and angular internal side of Lagomorpha BRANDT, 1855 the talonid, in H. b. brachygnathus the out- Leporidae FISCHER DE WALDHEIM, 1817 line of p3 trapezoidal, with broader anterior Hypolagus DICE, 1917 part of the trigonid and rounded internal margin. AER of p3 of moderate depth (average Type species Lepus vetus KELLOGG, 1910 17-20 % of crown width, not exceeding 30%); PER ca. 50 % of crown width (not Holotype Left mandible having incisor and exceeding 65%). H. b. brachygnathus occu- p3-m2 (Collection of University California, pies higher values of the observed range. The Vertebrate Palaeontology No. 12567), from buccal lobe of P2 smooth or expressing a U.C. locality No. 1100, Thousand Creek shallow groove (external anterior re-entrant), beds, Humboldt County, Nevada; Middle the lingual lobe of P2 in H. b. beremendensis Pliocene. smooth and rounded or flattened at the front, in H. b. brachygnathus it may express a shal- Emended diagnosis Medium to large-sized low groove. The postcranial skeleton expres- rabbit, with significant shortening of the dias- sing better cursorial ability than Oryctolagus, tema (Md-ratio, in most species, between 85 but lesser than Lepus. The ulnar olecranon and 105 %, the only exception is Hypolagus shorter and wider than that of Oryctolagus, schreuderae where it is ca. 140 %). Orbital but longer than that of Lepus. process of maxilla is at the right or slightly obtuse angle according to the zygomatic pro- Stratigraphic range Most of continental cess. p3 with two external folds (re-entrants): Europe and Sicily. H. b. beremendensis: Early shallower AER and deeper PER, the former to Late Pliocene (Late Ruscinian, MN 15 to extending up to 38%, the latter extending up Late Villányian, MN 17). H. b. brachygnathus to 65% of the crown width. from Late Villányian to Middle Pleistocene (Steinheimian, Q3). Studied Asian and European species Hypolagus beremendensis, Hypolagus igro- History Kormos (1934) described Hypo- movi, Hypolagus multiplicatus, Hypolagus lagus brachygnathus from the Hungarian site schreuderae, Hypolagus transbaicalicus. of Villány 3. Kretzoi (1962), who continued the work on Hungarian fossil localities, put Hypolagus beremendensis (PETÉNYI, both taxa into synonymy, and retained the 1864) species name Hypolagus beremendensis due to its priority. The author of this name was Holotype A left mandible with teeth, No. the Hungarian 19th century researcher S.J. Ob/3689 (HGIB), from Villány 3, latest Petényi. However, Petényi did not complete Villányian. his work on fossil mammals from Beremend before he died; in the posthumous edition of Emended diagnosis Middle sized his monography, completed by his co-worker Hypolagus having short and stocky muzzle. F. Kubinyi, the mandible of Hypolagus bere- Diastema is short, in H. b. beremendensis mendensis was illustrated (Petényi 1864: generally shorter than the lower tooth row Table II: 1 a, b). The drawings were not (Md-ratio 1: 85-95%), in H. b. brachygnathus accompanied by any diagnosis or description. slightly longer (Md-ratio 1: 95-105%). Remarks on skeletal findings assigned by

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Figure 8 Enamel patterns of the occlusal surfaces of p3. A – Hypolagus beremendensis beremendensis from Weze 1, Poland (from left: MF 2220/T/7, 23– reversed, 21, 54), B – Hypolagus beremendensis brachygnathus from Kamyk, Poland (from left: MF 2160/T/13, 8, 153, 166– reversed), C – Hypolagus igromovi (from Averianov 1996a: Figure 1. A, C, D– reversed, I, J), D – Hypolagus schreuderae from Loc. 18 near Beijing (from Teilhard de Chardin 1940: Figure 22: A, B), E – Hypolagus sp . from Harr Obo (V 7990, from Qiu 1987:Text fig. 32), F – Hypolagus multiplicatus (from Udunga – left and Beregovaya – right; after Erbajeva 1996: Fig. 2: A, 3: A), G – Hypolagus transbaicalicus (from Udunga – left and Beregovaya – right; after Erbajeva 1996: Fig. 2: B, 3: B).The dotted areas - cement.

Petényi to Lepus come only from his letters Sciences, and should be accepted as lectotype to Dr. K.C. Leonhard and Dr. H.G. Bronn of Hypolagus beremendensis. Further study enclosed in this edition (Petényi 1864). The on these populations, based on p3 and P2 original specimen illustrated by Petényi is morphotypes, made a distinction into two missing, but the type specimen of Hypolagus species: Hypolagus beremendensis and brachygnathus, the left mandible with teeth, Hypolagus brachygnathus, that were subse- No. Ob/3689, from Villány, described by quently changed into two subspecies: Kormos (1934), is kept in the Geological Hypolagus b. beremendensis and Hypolagus Museum of the Hungarian Academy of b. brachygnathus. The former was thought to

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be mainly a Pliocene form, characterised by a Holotype A fragment of maxilla with p3 of simple shape without any tracks of P2–M2, No. 56311 (ZIN), Beregova farm, anterior or internal folds, with a talonid River Czika valley, Baikal Region, Russia. slightly concave at the buccal side, and for- ming a sharp edge at the margin of PER, the Diagnosis A leporid of intermediate dimen- so-called "beremendensis edge" (Fladerer sions. P2 (width x length–1.6 x 3.0 mm) with 1987; Fladerer & Reiner 1996). The latter two anterior re-entrants; p3 rounded, with subspecies was characterised by the more nearly equal width and length, and extended complicated tooth outline, with traces of AR, internal margin; the re-entrants shallow (AER AIR, or PIR (see Fig. 2), and is believed to ca. 12% and PER 50-52% of the crown arise from a beremendensis population in the width). Upper cheek teeth possessing highly late MN 17. The specimen No. Ob/3689 was crenulated internal folds. Hypolagus multipli- stated as a holotype for this form (Fladerer & catus differs from the others Hypolagus spe- Reiner 1996). cies by possessing higher crenulated upper cheek teeth. It differs from H. beremendensis Hypolagus igromovi GUREEV, 1964 and H. schreuderae in being smaller, and from Hypolagus transbaicalicus in being larger. Holotype A fragment of left maxilla with P3-M1, No. 48636 (ZIN) from Rasdorskaya, Stratigraphic range Late Pliocene and the right bank of Don River, Rostov Region, Early Pleistocene of Baikal Region, Udunga, Russia; Upper Miocene, Lower Pontian beds. Beregovaya, Dodogol, Zasuchino 2 (Erbajeva 1996). Diagnosis Dimensions of big mountain hare (Lepus timidus). Mandible robust with high Hypolagus schreuderae TEILHARD DE alveolar part and broad angular process. CHARDIN, 1940 Mandibular diastema short, nearly equal to the tooth row (Md-ratio 1 ca. 100%). Incisors Holotype Almost complete specimen from robust like in modern mountain hare. Locality 18 near Peking, China, Late Maximal breadth of upper molars 1.5–1.75 Pliocene. times more than length. Upper teeth (P4-M2) re-entrants strongly crenulated. Re-entrants of Diagnosis According to Teilhard de Chardin p3 deep (AER ca. 19% of the crown width, (1940): "A large sized hare, with the lower p3 PER 52-61% of the crown width); the crown of a typical Hypolagus pattern (posterior outline usually triangular, with narrow and outer cleft incomplete, no anterior furrow). pointed anterior part of the trigonid, rounded First upper incisors relatively large. Auditory internal margin. In over 50% of the teeth a bullae relatively small and narrow. Posterior crinkle occupies the anterointernal part of the tip of the maxillary extending back of the crown observed. fronto-nasal suture. Triangular extension of the frontal between the nasalia large and Stratigraphic range Late Turolian (MN 13) broad." Diastema significantly long to ?Early Ruscinian, in Don River Region, (21,5–26,0 mm; Md-ratio 1 ca. 140%); p3 of foothills of the Caucasus (Gureev 1964; relatively shallow re-entrants depth (AER 15- Averianov 1996a; Averianov & Tesakov 17% of the crown width, and PER ca. 47-48%). 1998; Bajgusheva et al. 2001). Stratigraphic range Late Pliocene - Early Hypolagus multiplicatus (ERBAJEVA in Pleistocene (Villafranchian/Nihewanian) of BAZAROV et al., 1976) China, typical in such sites as Yushe III or Loc. 18 near Beijing.

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Hypolagus transbaicalicus (ERBAJEVA data and small samples. in BAZAROV et al., 1976) The clustering places Hypolagus igromovi among "H. beremendensis" populations, Holotype Left P2, No. 56310 (ZIN, collec- grouping it very close with the populations tion of M.A. Erbajeva 1963); River Czika from Kamyk and Villány 3 (Jánossy 1986). valley, Beregova farm, Russia. This suggests that the status of this species needs revision. The external position of the Diagnosis Small leporid. P2 with single Weze 1 population in relation to the anterior reentrant of a depth equal 1/3 of Villányian and Early Biharian H. beremen- tooth length. Upper cheek teeth possessing densis stresses the previously suggested sub- variably crenulated reentrants. Crenulation specific differentiation within H. beremen- developed weaker than in Hypolagus multi- densis. The external position of H. schreude- plicatus towards the back of the tooth row. rae supports the distinction of this species The smallest of all Eurasian Hypolagus. Both from the European populations, marked pre- p3 reentrants relatively shallow (AER: 8- viously by the significant longer diastema. 18%, and PER: 49-56 % of the crown width). The "beremendensis-igromovi" group Stratigraphic range The same as for The data on H. beremendensis included in the Hypolagus multiplicatus, co-occurring in present study, show mosaic evolution of the sediments. dental and mandibular features. From a morphological point of view, the "beremendensis- Note The coexistence of small and large igromovi" group forms a continuous group, Hypolagus (H. multiplicatus & H. transbai- with H. igromovi and Biharian populations of calicus) which share a unique derived charac- H. beremendensis at the edge of the highest ter (plications of the hypostria) suggests juve- dimensions. Intermediate positions are occup- nile and adult specimens of the same species. ied by the Late Villányian and earliest The large individual variability within the Biharian populations. lagomorph populations (see the Alilepus A statistical search for some metrical annectens study by Qui 1987) can be mislea- features as discriminators between the diffe- ding. Thus, to keep separately these two spe- rent populations within the "beremendensis- cies would need a further study on their com- igromovi" group was performed. The most mon range of variability against that expected useful feature proved to be the diastema for intraspecific variation. length. According to the Lubischev factor K, there is a significant difference between H. DISCUSSION beremendensis from Weze 1 on the one hand, and both H. igromovi (K=8.40) and the The dendrogram (Fig. 9) shows one complex Kamyk H. beremendensis population cluster, with an inner group consisting of (K=7.68) on the other hand. There are no Villányian to Early Biharian populations of such differences between H. igromovi, and Hypolagus beremendensis with externally Kamyk or Villány 3 populations of H. bere- attached Hypolagus igromovi and the Early mendensis, nor between H. igromovi and the Pliocene population of Hypolagus beremen- Rebielice Królewskie 1 H. beremendensis densis from Weze 1. The Nihewanian population. Thus, a morphological dis- Hypolagus schreuderae is placed externally continuity is observed between Weze 1 and of this "beremendensis-igromovi" group. The the remaining populations of the "beremen- Transbaikalian species Hypolagus multiplica- densis-igromovi" group. In this study, follow- tus and H. transbaicalicus were not included ing Fladerer & Reiner (1996), the Central into the dendrogram because of inadequate European H. beremendensis populations are

209 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

divided into two subspecies: H. b. beremen- H. b. brachygnathus is thought to exist in densis (Ruscinian-Early Villanian), which is Europe from Early to Middle Pleistocene small and has shallower re-entrants in p3, and (Fladerer 1987; Fladerer & Reiner 1996). Its H. b. brachygnathus (Late Villányian- remains are known from Poland (Kamyk), Steinheimian), which is larger and has a p3 Czech Republic (Chlum, Holstejn, Stránská with more complicated enamel pattern and Skála), and Austria (Deutsch-Altenburg). The deeper reentrants. abundance of this species decreased quickly H. b. beremendensis was abundant in in Biharian, due to the arrival and spreading Northwestern and Central European Pliocene. of Lepus (Kormos 1934; Sych 1965; Wolsan Its distribution extended over parts of France, 1989). The presumably last Eurasian through the Netherlands, Germany, Austria, Hypolagus survived in Moravian refuges the Czech Republic to Slovakia, and Poland (Stránská Skála 2 and Mladec 3) untill the in the north, and Hungary, Serbia, Romania, earliest Middle Pleistocene (Fladerer & and probably Greece in the south (Schreuder Reiner 1996). 1936; Sulimski 1964; Sych 1965; Koenigs- Hypolagus igromovi occurs in the Latest wald 1974; Jánossy 1986; Wolsan 1989; Miocene or, more plausibly, the Earliest Chaline et al. 2000). It is also known from Pliocene (Averianov 1996 a; Averianov & the Late Pliocene of Sicily, Monte Pellegrino Tesakov 1998). The original material of this site (Thaler 1972). The oldest and most species is referred to be of Sarmatian or numerous remains assigned to this subspecies Meotian (Late Miocene) origin (Gureev 1964; come from Polish sites: Weze 1, 2 (Sulimski Averianov 1996 a). The mandible of a suba- 1962, 1964; Sych 1965). Late Ruscinian H. b. dult Hypolagus (corresponding "well in the beremendensis is known also from the karst morphology of p3 and the state of preserva- sites of Slovakia (Ivanovce 1, Fladerer 1996), tion with the type material") was found in and Hungary (Csarnóta 1 and 3, Jánossy 1986). situ in the beds of Early Pontian age, overlain

Figure 9 Phenogram of six populations of Hypolagus from Eurasia, based on UPGMA clustering.

210 FOSTOWICZ-FRELIK: Eurasian Hypolagus

by clays of Villafranchian age, at the type tionships. locality (Averianov 1996 a, following person- Parallel evolution may be explained by the al communication V.V. Titov & A.S. environmental similarity. The longer muzzle Tesakov). In the original work by Gureev observed in the Pleistocene populations of H. (1964) this species is described as being lar- b. brachygnathus and in the Late Miocene H. ger than H. beremendensis and having a more igromovi (Fig. 4), could be explained by the complicated enamel pattern. However, need of longer nasal ducts to moisturize and Gureev (1964) did not know the Kamyk or warm inhaled air. Actually, at the Miocene/ Deutsch-Altenburg populations of H. bere- Pliocene transition, a global vegetation change mendensis, because they were not yet descri- was observed (Cerling et al. 1997). It resulted bed. On the other hand, Averianov (1996a), in the rapid expansion of open habitats, with emending the diagnosis, stated that H. igro- domination of C4 grasses, indicating drier movi differs from the European Pliocene H. climate. Also, the development of steppe- beremendensis and Pleistocene H. brachyg- tundra areas in the Early Biharian (Fladerer nathus in being larger (mandible and postcra- & Reiner 1996) was related to increasing ari- nial skeleton), comparing with material from dity and the arrival of tougher vegetation. Weze, Rebielice Królewskie, Kadzielnia, and This triggered selection towards a more com- Kamyk. In fact, H. igromovi is not signifi- plicated and thus more resistant p3 enamel cantly larger than Hypolagus from Kamyk pattern, with deep re-entrant folds, as obser- and most of its dimensions are well within ved in both species. A tentative relationship the observed range for Hypolagus from could be explained by supposing that H. b. Deutsch-Altenburg (Fladerer 1984) and some brachygnathus is derived from H. igromovi, specimens from Pleistocene Hungarian sites. which was invading Europe since the Early According to the enamel pattern, the p3 pre- Villányian (MN 16) and which gradually for- sented by Averianov (1996 a, Fig. 1) closely ced out H. beremendensis. This "relationship resembles those of H. beremendensis, except hypothesis" seems to be more acceptable on for the teeth G and H, which may either be the basis of the close morphological similari- aberrant or belong to Serengetilagus or ties observed, and because it is conceivable Trischizolagus. Moreover, the teeth of H. that H. beremendensis and H. igromovi origi- igromovi show an intermediate shape nated from a single stock. However, the taxo- between those of Weze 1 (presumed to be H. nomic position of H. igromovi and H. bere- b. beremendensis with a clearly triangular mendensis needs additional study. outline), and those of Kamyk and Villány 3 (supposed to be H. b. brachygnathus with a Chinese Hypolagus and more trapezoidal outline and with distinct Transbaikalian species antero- and postero-internal re-entrants; see The first problem to be dealt with is the sta- Figure 8). tus of Hypolagus brachypus (YOUNG, 1927). Thus, the problem of the relationship of H. Young (1927) described Caprolagus brachy- igromovi and H. beremendensis arises. First, pus from the Late Villafranchian of Loc. 60 a hypothesis that H. b. brachygnathus origi- (Sanchiatien near Beijing, China). It was nated from H. b. beremendensis in the latest characterised by "small dimensions and pos- Pliocene (Fladerer 1987; Fladerer & Reiner terointernal reentrant". Later, more remains 1996) is one of the possibilities; these sub- were assigned to this species (Teilhard de species form one evolutionary lineage. On the Chardin & Young 1931; Young 1935). other hand, H. igromovi and H. b. brachyg- Schreuder (1936) considered this taxon to be nathus share close similarities e.g. in meas- Hypolagus. Bohlin (1942) examined the col- urements of p3, which could be the result of lection, including the type specimen of either parallel evolution or of their close rela- "Caprolagus" brachypus, kept in the

211 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Paleontological Museum in Uppsala, noting less probable, although the shape of p3 from the great similarity of these specimens to Harr Obo (a narrow anterior border, absence Alilepus, without however ascribing these of a furrow or extended flattening), which is remains to Alilepus. Averianov (1996b) revi- typical for Hypolagus, is also known to occur sed the collection, which resulted in ascribing in some specimens of Trischizolagus the studied material of "Caprolagus" brachy- (Averianov & Tesakov 1997; Qiu 1987; Qiu pus to a new genus: Sericolagus AVERIANOV, & Storch 2000). The tooth from Harr Obo 1996. shows close similarity in measurements and Three Hypolagus species of Asian origin proportions to the "beremendensis-igromovi" are known, but their relationships remain group (Figs. 5, 7). However, until the mandi- obscure. These are: Chinese H. schreuderae ble of the species is known, its status is not and two species from Transbaikalian sites, H. certain. The Asian record is completed by a transbaicalicus and H. multiplicatus, all number of Hypolagus sp. remains from known from deposits of Villafranchian (or its Kazakhstan, Altai Region, and Bashkiria, that Chinese equivalent, Nihewanian) age may either belong to H. igromovi or to H. (Teilhard de Chardin 1940; Bazarov et al. beremendensis (Averianov 1996a; Erbajeva & 1976; Kretzoi 1987; Qiu 1987; Erbajeva Tutkova 1997). 1996; Erbajeva & Alexeeva 2000). The best known taxon is H. schreuderae (Teilhard de CONCLUDING REMARKS Chardin 1940; Qiu 1987; Cai 1989). This Comparison of the Eurasian species of Hypo- species is characterised mainly by a relatively lagus with the North American record of the long diastema and simple enamel pattern, genus shows the uniformity of the tooth mor- with shallow re-entrants (Fig. 3-8, Table 1). phology in the Eurasian populations. The The long diastema of H. schreuderae seems North American species seem to be more dif- to be a derived feature, characteristic also for ferentiated in the depth of the reentrants (Fig. evolutionary younger Leporinae, such as 6). It is not definitely known which American Oryctolagus or Lepus. In comparison, most taxa could be considered ancestral to Eurasian North American Hypolagus species, as well ones. Hypolagus parviplicatus, treated as one as some Alilepus and Pratilepus, have shorter of the more primitive and ancestral North diastemata and an Md-ratio 1 oscillating to c. American species (Dawson 1958; White 1991; 100.0% (Hibbard 1969; White 1991; White & Voorhies & Timperley 1997), displays the Morgan 1995; Voorhies & Timperley 1997). closest similarity in p3 with Eurasian species, However, the mandibles of both Trans- especially with H. b. beremendensis, accor- baikalian species are unknown up to now, ding to AER and PER - factors and general making a detaile comparison difficult. The crown outline. The trend of deepening the re- analysis of the p3 reveals some resemblance entrants in more advanced North American between H. schreuderae and the Trans- species (Voorhies & Timperley 1997) is also baikalian species H. multiplicatus and H. pronounced in Eur-asian Hypolagus, especial- transbaicalicus (Fig. 8): but the dental featu- ly in H. beremendensis. It seems, however, res of H. schreuderae seem to be highly that the oldest known Eurasian species (H. primitive. igromovi and H. sp. from Harr Obo) show an The earliest Asian Hypolagus fossil (a advanced stage of this feature. H. schreuderae, single p3) comes from the Chinese site of with rather primitive features of p3 and a sig- Harr Obo (Inner Mongolia, Early Jinglian, nificantly longer diastema, is distinct, so far MN 14), and was assigned to Hypolagus sp. being known only from the Chinese Early (Qiu 1987; Cai 1989). However, Qiu & Pleistocene. Storch (2000) suggested that it could be the The tentative routes of dispersal of Hypo- tooth of Trischizolagus maritsae. This seems lagus in Eurasia could be as follows: the genus

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Figure 10 Distribution of the Hypolagus species in Eurasia during the Late Miocene to Early Pleistocene, and the probable migration tracks. entered Asia probably no later than the Early REFERENCES Turolian, and then migrated to Europe (Fig. Averianov, A., 1996a – The Neogene rabbit Hypolagus 10). Regional species evolved in the Baikal igromovi Gureev, 1964 (Lagomorpha, Leporidae) Region and in Inner Mongolia. The European from southern European Russia – Acta zoologica record is partly the result of this first Neogene cracoviensia 39 (1): 61-66 migration event. A second migration from the Averianov, A., 1996b – O sistematiceskim polozenii Don Region into Central Europe could have krolika "Caprolagus" brachypus Young, 1927 taken place in the Late Villányian. (Lagomorpha, Leporidae) iz villafranka Kitaâ – Trudy Zoologiceskogo Instituta RAN 270: 148-157 ACKNOWLEDGEMENTS Averianov, A. & Tesakov, A.S., 1997 – Evolutionary I would like to thank M. Dawson and N. trends in Mio-Pliocene Leporinae, based on Lopez-Martinez for critically reading the Trischizolagus (Mammalia, Lagomorpha) – manuscript. Special thanks are extended to L. Paläontologische Zeitschrift 71 (1/2): 145-153 Kordos (Geological Institute of Hungary) and Averianov, A. & Tesakov, A.S., 1998 – The Lagomorpha M. Gasparik (Paleontological Department of (Mammalia) from the Early Pliocene Kosyakino the Natural History Museum, Budapest) for Locality of the Northern Caucasus – Paleontological access to specimens and assistance. I am Journal 32 (3): 305-309 indebted to M. Wolsan for reading the draft Bajgusheva, V.S., Titov, V.V. & Tesakov, A.S., 2001 – of this paper and encouragement. Ms A. The sequence of Plio-Pleistocene mammal faunas Holda-Michalska made the map and some from the south Russian Plain (the Azov Region) – in: drawings. Rook, L. & Torre, D. (eds.) – Neogene and Quater- nary continental stratigraphy and mammal evolution – Bollettino della Società Paleontologica Italiana 40 (2): 133-138 Bazarov, D.B., Erbajeva, M.A. & Rezanov, I.N., 1976 – Geologiâ i fauna opornych razrezov antropogena

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zapadnogo zabajkal’â – Nauka, Moscow Pleistocene biostratigraphic succession of Trans- Bohlin, B., 1942 – A Revision of the fossil Lagomorpha baikalia with emphasis on small mammals – in the Palaeontological Museum, Upsala – Bulletin of Quaternary International 68-71: 67-75 the Geological Institute Upsala 30: 117-154 Erbajeva, M.A. & Angermann, R., 1983 – Das Cai, B., 1987 – A preliminary report on the Late Pliocene Originalmaterial von Serengetilagus praecapensis micromammalian fauna from Yangyuan and Yuxian, Dietrich, 1941 – ergänzende Beschreibung und ver- Hebei – Vertebrata Palasiatica 25 (2): 124-136 gleichende Diskussion – Schriftenreiche für geologi- Cai, B., 1989 – Fossil lagomorphs from the Late sche Wissenschaften 19/20: 39-60 Pliocene of Yangyuan and Yuxian, Hebei – Vertebrata Erbajeva, M.A. & Tutkova, L.A., 1997 – Paleogene and Palasiatica 27 (3): 170-181 Neogene lagomorphs from Kazakhstan – in: Aguilar, Campbell, K.E., 1969 – Comparing postcranial skeletons J.-P., Legendre, S. & Michaux, J. (eds.) – Actes du of Pliocene rabbits – Papers of the Michigan Congrès BiochroM’97, Memoires et Travaux de l’ Academy of Science, Arts and Letters 1 (1): 99-115 E.P.H.E., Institut de Montpellier 21: 209-214 Cerling, T.E., Harris, J.M., MacFadden, B.J., Leakey, Fladerer, F.A., 1984 – Das Vordergliedmassenskelett von M.G., Quade, J., Eisenmann, V. & Ehrlinger, J.R. Hypolagus beremendensis und von Lepus sp. 1997 – Global vegetation change through the (Lagomorpha, Mammalia) aus dem Altpleistozän von Miocene/Pliocene boundary – Nature 389: 153-158 Deutsch-Altenburg (Niederösterreich) – Beiträge zur Chaline, J., Erbajeva, M.A. & Montuire, S., 2000 – Paläontologie von Österreich 11: 71-148 Upper Pliocene leporids (Mammalia, Lagomorpha) Fladerer, F.A., 1987 – Beitrag zur Entwicklung von from Montoussé-5 (Hautes-Pyrénées, France) – Hypolagus und Lepus (Lagomorpha, Mammalia) im Neues Jahrbuch für Geologie und Paläontologie, Pliopleistozän von Mitteleuropa – Sitzungsberichte Monatshefte 2000 (2): 93-106 der Österreichischen Akademie der Wissenschaften, Dawson, M., 1958 – Later Tertiary Leporidae of North Mathematisch-naturwissenschaftliche Klasse, America – University of Kansas Paleontological Abteilung I 196: 123-138 Contributions 6: 1-75 Fladerer, F.A. & Reiner, G., 1996 – Evolutionary shifts Dawson, M., 1967 – Lagomorph history and the strati- in the first premolar pattern of Hypolagus beremen graphic record – pp. 287-316, in: Teichert, C. & densis (Petényi, 1964) (Lagomorpha, Mammalia) in Yochelson, E.L., (eds) – Essays in Paleontology and the Plio-Pleistocene of Central Europe – Acta zoolo Stratigraphy, Raymont C. Moore Commemorative gica cracoviensia 39 (1): 147-160 volume – University Kansas, Department of Geology Fostowicz-Frelik, L., 2001 – Limb biomechanics of the Special Publications 2 Plio-Pleistocene lagomorph Hypolagus beremenden De Bruijn, H., Dawson, M. & Mein, P., 1970 – Upper sis and its behavioral implications – Sixth Pliocene Rodentia and Insectivora (Mammalia) from International Congress of Vertebrate Morphology – the isle of Rhodos (Greece) – Proceedings Konink- Jena, Germany, July 21-26, 2001. Abstract volume, lijke Nederlandsche Akademie van Wetenschappen Journal of Morphology 248 (3): 230 B 73 (5): 568-584 Gureev, A.A., 1964 – Fauna ZSRR. Mlekopitaûsie, Dice, L.R., 1917 – Systematic position of several Zajceobraznye (Lagomorpha) III (10) – Nauka, American Tertiary lagomorphs – Bulletin of the Leningrad Department of Geology, University of California Hibbard, C.W., 1963 – The origin of the p3 pattern of Publications 10 (12): 179-183 Sylvilagus, Caprolagus, Oryctolagus and Lepus – Dice, L.R., 1929 – The phylogeny of the Leporidae, with Journal of Mammalogy 44 (1): 1-15 description of a new genus – Journal of Mammalogy Hibbard, C.W., 1969 – The rabbit (Hypolagus and 10 (4): 340-344 Pratilepus) from the Upper Pliocene, Hagerman Erbajeva, M.A., 1996 – Lagomorphs from Villafranchian Local Fauna of Idaho – Papers of the Michigan sequence in Transbaikalia and their paleoenvironmen- Academy of Science, Arts, and Letters 1 (1): 81-97 tal implications – Acta zoologica cracoviensia 39 (1): Jánossy, D., 1986 – Pleistocene Vertebrate Faunas of 131-135 Hungary – Developments in Paleontology and Erbajeva, M.A. & Alexeeva, N.V., 2000 – Pliocene and Stratigraphy 8, Elsevier, Amsterdam: 208 pp.

214 FOSTOWICZ-FRELIK: Eurasian Hypolagus

Kormos, T., 1934 – Az Euráziai nyulak származástani from Weze 1 (Poland) – Acta Paleontologica problémája – Allattani Közlemények 31(1-2): 65-78 Polonica 9 (2): 149-262 Kretzoi, M., 1962 – Fauna und Faunenhorizont von Sych, L., 1965 – Fossil Leporidae from the Pliocene and Csarnóta – A Magyar Állami Földtani Intézet Évi the Pleistocene of Poland – Acta zoologica cracovien- Jelentése Az. 1959: 309-395 sia 10 (1): 1-88 Kretzoi, M., 1987 – Remarks on the correlation between Teilhard de Chardin, P., 1940 – The fossils from Locality European and Asian Late Cenozoic local biostrati 18 near Peking – Palaeontologica Sinica, New Series graphies – Vertebrata PalAsiatica 25(2): 145-157 C 9: 1-101 Kurtén, B., 1968 – Pleistocene Mammals of Europe – Teilhard de Chardin, P. & Young, C.C., 1931 – Fossil Weidenfeld and Nicolson, London: 317 pp. Mammals from the Late Cenozoic of Northern China Lopez-Martinez, N., 1985 – Reconstruction of ancestral – Palaentologia Sinica, New Series C, 9 (1): 1-89 cranioskeletal features in the order Lagomorpha - in: Thaler, M.L., 1972 – Les rongeurs (Rodentia et Luckett, W.P. & Hartenberger, J.-L. (eds.) – Evolu- Lagomorpha) du Monte Pellegrino et la question des tionary relationships among rodents: A multidiscipli anciens isthmes de la Sicile – Comptes Rendus des nary analysis – NATO ASI Series, Series A, 92: Séances de l'Académie des Sciences, Paris, Serie D 227–276. Plenum Press, New York 274: 188-190 McKenna, M.C. & Bell, S.K., 1997 – Classification of Young, C.C., 1927 – Fossile Nagetiere aus Nord-China – mammals above the species level – Columbia Palaeontologia Sinica, New Series C 5 (3): 1-82 University Press, New York: 631 pp. Young, C.C., 1935 – Miscellaneous Mammalian Fossils Petényi, S.J., 1864 – Hátrahagyott munkái – Pest from Shansi and Honan – Palaeontologia Sinica – Qiu, Z., 1987 – The Neogene faunas of Ertemte and Harr New Series C 9 (2): 1-57 Obo in Inner Mongolia (Nei Mongol), China – 6. Von Koenigswald, W., 1974 – Solnhofen 5, eine Hares and pikas – Lagomorpha: Leporidae and villafranchische Spaltenfüllung aus Bayern – Ochotonidae – Senckenbergiana Lethaea 67 (5/6): Mitteilungen der Bayerischen Staatssammlung für 375-399 Paläontologie und historische Geologie in München Qiu, Z. & Storch, G., 2000 – The early Pliocene 14: 39-48 Micromammalian Fauna of Bilike, Inner Mongolia, Voorhies, M.R. & Timperley C.L., 1997 – A new China (Mammalia: Lipotyphla, Chiroptera, Rodentia, Pronotolagus (Lagomorpha: Leporidae) and other Lagomorpha) – Senckenbergiana Lethaea 80 (1): 173- leporids from the Valentine Railway Quarries 229 (Barstovian, Nebraska), and the Archaeolaginae- Radulesco, C. & Samson, P., 1967 – Contribution à la Leporinae transition – Journal of Vertebrate connaissance du complexe faunistique de Malusteni- Paleontology 17 (4): 725-737 Beresti (Pléistocène inférieur), Roumanie I. Ord. White, J.A., 1991 – North American Leporinae Lagomorpha, Fam. Leporidae – Neues Jahrbuch für (Mammalia: Lagomorpha) from Late Miocene Geologie und Paläontologie, Monatshefte 9: 544-563 (Clarendonian) to latest Pliocene (Blancan) – Journal Schreuder, A., 1936 – Hypolagus from the Tegelen clay; of Vertebrate Paleontology 11 (1): 67-89 with a note on recent Nesolagus – Archives Neér- White, J.A. & Morgan, N.H., 1995 – The Leporidae landaises de Zoologie 2 (2/3): 225-239 (Mammalia: Lagomorpha) from the Blancan Sen, S. & Erbajeva, M. 1995 – Early Pliocene Leporids (Pliocene) Taunton Local fauna of Washington – (Mammalia, Lagomorpha) from Afganistan – Journal of Vertebrate Paleontology 15 (2): 366-374 Comptes Rendus de l’Académie des Sciences, Paris, Wolsan, M., 1989 – Zaj_czaki, Lagomorpha – in: serie IIa 320: 1225-1231 Kowalski, K., (ed.) – Historia i ewolucja ladowej Sneath, P.H.A. & Sokal, R.R., 1973 – Numerical fauny Polski – pp. 145-150, Folia Quaternaria 59–60, Taxonomy – W. H. Freeman and Company, San Kraków Francisco: 593 pp. Sulimski, A., 1962 – O nowym znalezisku kopalnej Received 19 May 2001 fauny kregowców w okolicy Dzialoszyna – Przeglad Revision received 17 June 2002 Geologiczny 10: 219-223 Accepted 08 September 2003 Sulimski, A., 1964 – Pliocene Lagomorpha and Rodentia

215 DISTRIBUTION AND MIGRATION OF TERTIARY MAMMALS IN EURASIA DEINSEA 10, 2003

Table 1 Measurements (in mm) of p3 and mandibles for Hypolagus species (approximated values italicized).

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