Pliocene Carnivores (Carnivora, Mammalia) from Ivanovce...

Cour. Forsch.-Inst. Senckenberg 246 15–53 9 Figs, 8 Pls, 19 Tabs Frankfurt a. M., 07. 04. 2004

Pliocene Carnivores (Carnivora, Mammalia) from Ivanovce and Hajnáčka (Slovakia)

With 9 fi gs, 8 pls, 19 tabs

Oldřich FEJFAR & Martin SABOL

Abstract

Carnivores from two late Pliocene Slovak sites – Ivanovce (late MN 15) and Hajnáčka (early MN 16) are described and their taxonomy is discussed. In both sites, the ocurrence of the families Mustelidae and Procyonidae is similarly represented by Lutra cf. bravardi and Parailurus. However, both sites differ by the presence of Parailurus hungaricus, Megantereon sp., Pliohyaena perrieri, Ursidae gen. et spec.indet. in Hajnácka, and by the presence of Parailurus cf. anglicus, Hesperoviverra carpathorum and Viverridae gen. et spec.indet. in Ivanovce. The short temporal occurrence of the large viverrids fi lled the “gap” just since the dissappearance of the Turolian ictitheriids and just before the appearance of the “true” Canids.

The situation cannot be explained simply by paleoecology, because the paleoenvironmental conditions were quite comparable as shown by corresponding occurrence of Amphibians (Anurans), mastodons and tapirs. Both sites represent humid forest belts, along a volcanic lake (in a maar crater) in Hajnacka or along a broad river valley with a karstifi ed limestone massif in Ivanovce; both areas were probably surrounded by an open drier habitat in higher elevations.

Key words: Carnivores, Lutra, Parailurus, Mustela, Hyaena, Hesperoviverra, Megantereon

Zusammenfassung

Raubtiere aus zwei oberpliozänen Lokalitäten der Slowakei – Ivanovce (MN 15b) and Hajnáčka (MN 16a) – werden beschrieben und ihre taxonomische Stellung wird diskutiert. In beiden Fundstellen ist das Vorkommen der Familien Mustelidae und Procyonidae ähnlich repräsentiert von Lutra cf. bravardi und Parailurus.

Beide Fundstellen unterscheiden sich jedoch einerseits durch das Auftreten von Parailurus hungaricus, Megantereon sp., Pliohyaena perrieri, Ursidae gen. et spec.indet. in Hajnácka, einerseits durch die Präsenz von Parailurus cf. anglicus, Hesperoviverra carpathorum and Viverridae gen. et spec.indet. in Ivanovce.

Diese Situation kann nicht paläoökologisch erklärt werden, da die Umweltbedingungen an beiden Fäl-len vergleichbar sind, bestätigt durch das übersinstimmende Vorkommen der Amphibien (Anuren), Masto-donten und Tapire. Paläoökologisch repräsentieren beide Fundstellen feuchte bewaldete Ufer entlang eines vulkanischen Sees (im Maar-Krater) in Hajnacka oder am breiten Flussufer in Ivanovce; in beiden Fällen existierten in der weiteren Umgebung offene locker bewaldete Anhöhen.

Schlüsselworte: Carnivore, Lutra, Parailurus, Mustela, Hyaena, Hesperoviverra, Megantereon

Authors’ addresses: Prof. Dr. Oldřich FEJFAR, Department of Paleontology, Faculty of Sciences, Charles University, Albertov 6, CZ – 128 43 Prague 2, Czech Republic, <[email protected]>; Dr. Martin SABOL, Department of Geology and Paleontology, Faculty of Sciences, Comenius University, Mlynská dolina, SK – 842 15 Bratislava, Slovak Republic, <[email protected]>

FEJFAR & SABOL: Pliocene Carnivores (Carnivora, Mammalia) from Ivanovce and Hajnáčka (Slovakia)

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Introduction

Fossil records of Carnivores from the late Pliocene are rela-tively rare in Europe. Therefore, fossils of all the Pliocene carnivores are very important, because they are our prime source of knowledge on the development of recent Car-nivoran faunas not only in Europe, but also in Asia.

In the territory of Slovakia, some sites are known where the remains of late Pliocene fossil mammals have been collected. However, only two of them – Ivanovce and Hajnáčka – also yield a large amount of carnivore remains and both are also Type localities for some taxa. The general paleoenvironmental surroundings of both localities was similar – a humid, but not swampy, forestal belt along lake or river banks, although they represent quite different types. Hajnáčka lies in the center of a volcanic area and in geological terms is fully connected with the volcanic context; here, the fossiliferous layers (fi ne grained sands overline by coarse tuffs and fi ne bedded tuffi tes with fl ora) were deposited in/or around a small maar lake (its diameter being approximately 1500 m). In contrast, the fauna of Ivanovce comes from reddish clays in the tectonically disturbed karst fi ssures; the skeletal remains here are highly disarticulated and fragmented, and small mammals are much more frequent (FEJFAR & HEINRICH 1985, FEJFAR, HEINRICH & HEINTZ 1990).

The Ivanovce locality was discovered by FEJFAR (1961a: 262–263) in the middle of the last century. The site is situated in western Slovakia approximately 12 km south-westward from Trenčín (longitude (LG) is 17o 54´ and latitude (LA) is 48o 49´) (fi gs A, B). It consists of a reddish sediment fi ll within horizontal and vertical fi ssures of karstic tectonic deformed Triassic limestone. Based on the micromammal fauna (e. g. Mimomys gracilis, M. da-vakosi, Bjornkurtenia canterranensis and Trilophomys depereti), the age of the locality was assigned to the late Early Pliocene (Late Ruscinian, MN 15b) (FEJFAR 2001: 192). Besides some new species of micromammals (FEJFAR 1961b), two new viverrid taxa (Hesperoviverra carpatho-rum and Viverridae gen. et spec.) have also been described from the sediments of the site (KRETZOI & FEJFAR 1982: 980–982, 985). Later, an updated faunal list was presented (FEJFAR & HEINRICH 1985: 220–222).

Fossil mammalian remains from Hajnáčka were fi rst reported in 1863, by KUBINYI (FEJFAR 1964: 7). During subsequent years many paleontologists have reported fossils from this locality; fossil records of carnivores were reported by KORMOS (1917), FEJFAR (1964) and SABOL (2000). KORMOS (1934: 17–18, 26) described new species of the lesser panda (Parailurus hungaricus) from the locality. This record was added to by FEJFAR (1964: 57–59) reporting new records of Parailurus discovered during research in 1955–1958. He also described remains of a fossil hyena Crocuta (Plesiocrocuta) perrieri, and other poorly represented carnivores (machairodontid and lutrine). The Hajnáčka locality is situated approximately 1.5 km SE of the village Hajnáčka near Rimavská Sobota

in Southern Slovakia in the Novohrad Basin (LG is 19o 58´ and LA is 48o 12´) (fi gs A, B). The fossiliferous sequence consists mainly of fi ne grained, tuffaceous and poorly indurated sediments (silty sands, tuffi tes and pyroclastic inclusions) deposited in a small volcanic lake and partly redeposited by Pleistocene land slides and/or solifl uction. The assemblage of fossil rodents (e.g., Mimomys stehlini and M. hassiacus) at Hajnáčka indicates a slightly younger age in comparison with the karst fi llings of Ivanovce. On the basis of these rodent records, the Hajnáčka biocenosis has been correlated with the lower part of the Late Pliocene (Early Villányian (= “Villafranchian”), MN 16a (FEJFAR et al. 1998: 534).

Thus, these sites represent stratigraphically two chronologically distinct faunal phases in time succession (Ivanovce – late Early Pliocene, MN 15, Hajnáčka – early Late Pliocene, MN 16). Paleomagnetic samples in 1980 correlate the Hajnáčka section with the early Gauss mag-netic chron and with other similar Pliocene sites as, e.g., Arondelli-Triversa in Italy and Valdeganga in Spain.

From the sites in the volcanic context of Auvergne in France, three are important for comparison with localities mentioned here: Vialette, Perrier-Les Étouaires and Senèze; they cover the span of the interval of the mammalian stage Villanyian. According to the succession of the index species of arvicolids, the oldest site Vialette well corresponds the level of Hajnáčka (MN 16a), the younger level of Perrier-Les Étouaires corresponds to the karst fi lling Rembielice Królewskie in Poland (MN 16b) and the youngest assem-blage of mammals from Senèze represents the MN 17 level of Saint Vallier. (FEJFAR & HEINRICH 1983, LINDSAY, OPDYKE & FEJFAR 1997, FEJFAR, HEINRICH & LINDSAY 1998).

Material and methods

Fossil remains of carnivores described here were found during FEJFAR´s long research of Hajnáčka and Ivanovce in the 50s organized by Central Geological Survey in Prague, and by excavations in Hajnáčka l during 90s supported by the Faculty of Sciences, Comenius University in Bratislava and the Gemer-Malohont Museum in Rimavská Sobota. The fossiliferous sediments were extensively screenwashed in the fi eld to obtain small mammalian species; during the collection of these sediments some fi nds of carnivore were made. All material studied and reported herein is deposited in the fossil vertebrate collections of the Slovak National Museum – Museum of Natural History (SNM-MNH) in Bratislava and the Gemer-Malohont Museum (GMM).

For this paper, fossils were documented by digital camera Nikon – Coolpix 995 and photos have been con-verted for plates 1–8 and fi g. 5 usinge Corel-Photo Paint 8 software. The measurements of studied carnivoran remains were made with callipers to 0.1 mm calibration precision. The standard deviation of measurements is 0.3 mm with 0.1 mm dispersion and random errorof 4.2 %. All measured data are in millimetres.

Cour. Forsch.-Inst. Senckenberg, 246, 2004

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Fig. A: 1 – Territory of Slovakia with the late Pliocene sites Ivanovce and Hajnáčka, 2 – panoramatic view from the south on the village Hajnacka showing the volcanic (basaltic) cone in the center. The fossilferous sites are in the lower wooded part of the picture.


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Fig. B: 1 – The northern wall of the former limestone quarry at Ivanovc near Trenčín in western Slovakia; several karst fi llings provided rich late Pliocene mammalian fauna. 2 – the schematic sketch of the same view showing the location of different fi llings taken as samples for extensive washing in 1960–1965.


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Abbreviations of collections:

GMM: Gemer-Malohont Museum, Rimavská Sobota, Slovak Republic NMP: National Museum, Prague, Czech RepublicSMF: Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, FRG ZMB: Zoologisches Museum Berlin, FRGSNM-MNH: Slovak National Museum – Museum of Natural History, Bratislava, Slovak Republic

Systematics

Class Mammalia LINNAEUS, 1758Order Carnivora BOWDICH, 1821

Suborder Fel i formia KRETZOI, 1945Family Fel idae FISCHER DE WALDHEIM, 1817Subfamily Machairodont inae GILL, 1872

Tribe Smilodont ini KRETZOI, 1929Genus Megantereon CROIZET & JOBERT, 1828

Megantereon sp.(pl. 1, fi gs 12, 13)

1985 cf. Megantereon sp. – FEJFAR & HEINRICH: 221.1990 Megantereon (?) sp. – FEJFAR, HEINRICH & HEINTZ: 52.

Mater ia l : The fi rst right metacarpus (SNM-MNH, Z 26376) from probe 8/56 of Hajnáčka locality.

Type local i ty: Perrier, France.

The age: Early Villányian (Late Pliocene).

Descr ip t ion: The basis for identifi cation of the fi rst right metacarpus consists of two articulated areas, which are separated from each other by a shallow longitudinal depression (canalis metacarpi proximalis). The sharply restricted larger articulated area is situated on the lateral side and the lesser one is situated on the medial side. The proximal head of the bone is sloped lateral-medially on the distal side, approximately with an angle of 45o. On the dorsal side of the head, a distinct shallow semicircu-lar depression is also present. A distinct sharp crest is a dominant feature of the distal head. The crest runs from the dorsal side of the bone to the palmar one and separates the trochlea on the lateral side. A dorsal longitudinal sulcus runs distinctly from the medial area on the bone base to the crest of the distal head. The palmar longitudinal sulcus is less massive. A distinct laterally compressed sharp crest is present on the lower part of the palmar side of the distal

head for the phalanx PH I. This ridge divides the trochlea into two approximately equally large parts.

The measurements : maximum length (from lateral margin of the canalis metacarpi proximalis to ridge of the distal head) is 33.1 mm, maximum width of the bone base (from lateral margin of the base to its medial margin) is 19.8 mm, maximum width of the distal head (from lateral margin of the head to its medial margin) is 16.2 mm, width in the centre of the bone body is 13.5 mm, and height of the bone body in its centre is 13.3 mm.

Discuss ion: The fi rst metacarpus of Megantereon is shorter than the fi rst metacarpus of the lion (Panthera leo), but more robust.

Family Viverr idae GRAY, 1821Subfamily Viverr inae GRAY, 1821

Genus Hesperoviverra (KRETZOI et FEJFAR, 1982)

Hesperoviverra carpathorum (KRETZOI & FEJFAR, 1982)(pl. 2, fi gs 1–8)

1982 Megaviverra carpathorum n. g. n. sp. KRETZOI & FEJFAR: 980–982, text-fi gs 2, 3.1-3, pl. 1 fi gs 1–8.

1985 Hesperoviverra carpathorum (KRETZOI & FEJFAR). – FEJFAR & HEINRICH: 221.

Mater ia l : 1. right mandible fragment with p4, m1 and alveolus of m2, broken in front of p4, and with damaged angular and condylar process (SNM-MNH, Z 26386/65261, Holotype, fi ssure Nr. 6523). 2. P4 right (SNM-MNH, Z 26387/6512108, paratype, fi ssure Nr. 6512), western Slovakia. 3. The left lower canine (SNM-MNH, Z 26386b/651265, holotype, fi ssure Nr. 6512).

Type local i ty: Ivanovce, the horizontal fi ssure fi llings Nr. 6523, 6512, containing an early MN 15 assemblage with prevailing Trilophomys and Bjornkurtenia; western Slovakia: reddish fi lling of karst fi ssures in the Triassic limestone.

The age: Late Ruscinian, MN 15b (Early Pliocene).

Description: The Holotype: the mandible is massive and high with deep and long masseter fossa. The both teeth preserved – p4, m1 – have a massive “infl ated” shape. The lower p4 has oval outline with posterior broader. High massive protoconid is buccally infl ated, its anterior crista connects smal parastylid and posterior crista goes steeply to the sharp notch between the distal buccaly shifted metastylid. The posterior border is fl anked by a circular serrated cingulum which continues as a shallow elevation of cingular character. m1: all cusps are simple and well developed. The trigonid take more than half of the crown. The massive voluminous protoconid cause the buccal ex-pansion of the outline, carnassial blade is high, effective


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and divided symmetrically in the center by a deep notch. The equally long but lower paraconid is anterobuccally surrounded by a weak cigulum. The lower metaconid is slightly caudally shifted and medioanteriorly delimited by a thin slit; the trigonid dip opens steeply lingually. The gentle junction between trigonid and talonid is oblique indicated by sharp notches separating procotonid from hypoconid and metaconid from entoconid, respectively. The shallow lingually inclined talonid is distally bounded by three well developed low connected cusps (the lowest hypoconulid, entoconid, and a strong hypoconid). By the demage of the lingual part of dentary the roots of both p4 and m1 are exposed; m1 has two sligthly divergent later-ally fl at roots, p4 has roots thinner and less divergent. The alveole of the m2 is dubble strangled and indicates two roots fused together.

The juvenile upper P4 is not yet worn; the strong pro-toconid is shifted mesially, its distal crest joins a narrow denticulated cingulum, mesial crest points to the anterior buccal parastyl. Buccally, the highest paracone and low metacon are divided by a deep slesh of the carnassial blade buccally opened by a broad depression. The mesial crest of the high paracon joins the distal crest of the strong parastyl; lingually towards the protocone is a deep hollow. The buccal base of the crown is fl anked by a shallow edge of cingular character.

The left lower canine has a damaged top of the crown; the root is complete. A small wear facet for contact with the upper canine is situated on the posterior buccal side of the crown. On the posterolingual side, a weak crest is situated which probably ran from the top to the crown base. In dorsal view the tooth has an S-like shape.

D i s c u s s i o n : similar roughly contemporaneous (Ruscinium, MN 15 – early Villányium, MN 16a) viverrids are known from following localities: 1. Yassiören, north-west of Ankara, Turkey: Hesperoviverra intertuberculata (OSANSOY, 1965), the largest species of the viverrids originally classifi ed as Ictitherium; it is very similar and may be conspecifi c with H. carpathorum. 2. Arondelli-Triversa, Villafranca d’Ásti, north Italy: Hesperoviverra (?) apennina KRETZOI & FEJFAR, 1982, and 3. East Africa: Hesperoviverra (?) leakeyi (PETTER, 1963). According to KRETZOI and FEJFAR (1982) these late Pliocene large viverids represent a dispersal of one genus of south Asian origin.

Viverrinae gen. et spec. indet.(pl. 2, fi gs 9–11)

1982 Megaviverra (?) sp. KRETZOI & FEJFAR: 985, text-fi gs 5, pl. 2 fi g. 11.

Material : incomplete fragment of the left mandible with alveoli for m1 and m2 (SNM-MNH, Z 26389/65264, Holo-type) from Ivanovce (fi ssure Nr. 6523).

Type loca l i ty : Ivanovce, fi ssure Nr. 6523, western Slovakia.

The age: Late Ruscinian, MN 15b (Early Pliocene).

Description: Fragment of the left mandible (introduced by KRETZOI & FEJFAR (1982), with front part of mandible to the back wall of the alveolus of the anterior root of m1is

Sites Teeth etc.

Ivanovce, Slovakia H. carpathorum

(Type and Paratype)

Yassi ren , Turkey H. intertuberculata

Arondelli-Triversa, Italy H. apennina

p2 L W

/ / 9,90 4,20

p3 L W

/ / 11,77 4,95

p4 L W

14,0 7,4

17,4 8,8

13,42 5,76

m1 L W

18.8 11.7 [*7,6]

20,2 10,0

16,23 8,42 [*7,24]

mandible: height (under m1)

27.2 / 19,0

c sag trv

10,3 6,8

/ 7,42 5,64

C sag trv / / 9,17

5,71 P4 L W

19,5 11,0

24,0 13,8

17,1 +10,25

Table 1: The measurements of Hesporoviverra


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missing. Posterior part of the mandible is fl at on the lingual side, whereas its buccal side is expanded arc-like. The coronoid process is high and relatively narrow, concave on the top; the condyle and angular process are broken off. Presence of a blunt subangular process is situated in the area of the mandible base, below the front of the external temporal fossa boundary. The external temporal fossa is deep, extending by its front margin to below the m2 alveo-lus. Mandibular foramen is small and semicircular. Two alveoli of the molar m 1 are present: the anterior alveolus is preserved as a back wall with a blunt crest in the centre, the oval posterior alveolus is complete; it has a weak crest on the lingual side. The alveolus of m2 is similar in shape but smaller and narrower.

The measurements : mandible height is 54.6 mm, (height measured between condylar and angular process is 19.3 mm); mandible height measured of m1 is 20.1 mm.

Discussion:

Family Hyaenidae GRAY, 1821Subfamily Hyaeninae GRAY, 1821

Genus Pliohyaena KRETZOI, 1938

Pliohyaena perrieri (CROIZET & JOBERT, 1828)(pl. 1, fi gs 1–11)

1828 Hyaena perrieri CROIZET & JOBERT: 175–178, pl. I fi g. 12, pl. II fi gs 1, 2, 3, 5, 6, 7, pl. IV fi gs 3, 5, 6.

1828 Hyaena arvernensis CROIZET & JOBERT. – CROIZET & JOBERT: 178–180, pl. I fi g. 4, pl. III fi gs 1, 2, pl. IV: fi gs 1, 2, 4.

1883 Hyaena topariensis FORSYTH MAJOR. – FORSYTH MAJOR: 2.1885 Hyaena sp. FORSYTH MAJOR. – FORSYTH MAJOR: 2.1889 Hyaena topariensis FORSYTH MAJOR. – WEITHOFER: 342–346,

pl. I: fi gs 1–4, pl. II fi gs 1, 2, pl. III: fi g. 3, pl. IV: fi gs 3, 4.1889 Hyaena robusta WEITHOFER. – WEITHOFER: 346–356, pl. II:

fi gs 3–5, pl. III: fi gs 1, 2, pl. IV: fi gs 1, 2.1930 Hyaena sinensis TEILHARD & PIVETEAU. – TEILHARD & PIVETEAU:

101–104, pl. XX fi gs 1, 1a, 2, 2a.1934 Hyaena licenti PEI. – PEI: 120–121.1936 Hyaena perrieri CROIZET & JOBERT. – HELLER: 123–124, pl. IX:

fi g. 2.1938 Pliocrocuta perrieri (CROIZET & JOBERT). – KRETZOI: 118–119.1953 Hyaena arvernensis CROIZET & JOBERT. – LEHMANN: 441,

fi g. 3, 4.1954 Pachycrocuta robusta KRETZOI. – KRETZOI: 246.1954 Crocuta (Plesiocrocuta) perrieri (CROIZET & JOBERT). – VIRET:

46–52, pl. 5 fi gs 1, 2, pl. 6 fi gs 1–8, pl. 7 fi gs 1–5, pl. 8 fi gs 1, 2.

1957 Crocuta perrieri (CROIZET & JOBERT). – LEHMANN: 69, taf. 4 fi gs 7–9.

1957 Crocuta perrieri (CROIZET & JOBERT). – FEJFAR: 73.1961 Crocuta perrieri (CROIZET & JOBERT). – FEJFAR: 262.1964 Crocuta (Plesiocrocuta) perrieri (CROIZET & JOBERT). – FEJFAR:

59–60, text-fi g. 37, pl. VIII fi gs 4–6.1970 Pachycrocuta perrieri (CROIZET & JOBERT). – FICCARELLI &

TORRE: 18–20, 30, fig. 1; pl. I–III, VIII, XVI, XIX, XX, XXI: 1, XXII: 5,

1985 Hyaena cf. perrieri (CROIZET & JOBERT). – FEJFAR & HEINRICH: 221.1987 Pliohyaena perrieri (CROIZET & JOBERT). – QIU ZHAN XIANG:

50–52, 67–68, 79–80, fig. 15, tab. VIII, pl. 10, 11: 1. 1990 Hyaena cf. perrieri (CROIZET & JOBERT). – FEJFAR, HEINRICH &

HEINTZ: 52.

Ma te r i a l : 1. incomplete right mandible with c and p2– p4 in situ (SNM-MNH, Z 26379; surface fi nd), left upper juvenile canine (SNM-MNH, Z 26378; probe 8/56), 2. right p2 (GMM, No. B-4063; probe 1/96), and 3. left calcaneus (SNM-MNH, Z 26377, probe 8/56) from Hajnáčka I.

Type local i ty: Perrier, Auvergne, France.

The age: Early Villányian, Late Pliocene (s. c. level of Les Étouaires, the biozone MN 16b).

Descript ion: 1. Right mandible with canine and p2 – p4 and the left upper canine have been introduced by FEJFAR (1964: 59–60). The longitudinal axis of the symphysis form with the horizontal axis of the dentary an angle 110o; the length of diastema is 10 mm; the rounded foramen mentale is 15 mm below the p2. The teeth preserved are middle-worn with fl at distally inclined facets. The typically screw shaped canine has the medial edge at the higher elevagted crown basis and a posterolingual wear facet. In occlusal view, the three premolars are arranged in an arch curved roughly similar as the lingual body of the mandible. The p2 is smallest, oval, with relatively low central protoconid; its both crests divide the crown sym-metrically, the anterior connetcting the reduced parastylid fl anked lingually with a short singular edge; the posterior crest point to the low metastylid separated by shallow notch. The two premolars p3 and p4 are similar in size and oclusal outline (p3 being little more oval and expanded) but quite different in morphology. P3 shows the strongest conical protoconid without parastylid – ther anterior crest ends blind on the mesial border. The metastylid has similar shape as in p2, weak serrated cingulum is indicated on the mesial border. In lateral views the axis of p2 displays a typical “hyenid” posterior inclination. In occlusal view p3 and p4 take a mutual “coulisse” position. In contrast to the previous premolars p4 has all three cusps: less expressed central protoconid fl anked by nearly symmetrically worn para- and metastylid, separated by shallow nicks.

2. The right p2 with two broken roots was found during the washing of fossiliferous sediments from probe 1/96. The crown of the premolar is slightly damaged by grooves on the surface. The most distinct cusp at the crown is the protoconid with two crests, anterior and more distinct posterior crests run from the top to the base of the crown. Small indistinct paraconid is situated at the mesial ridge of the crown and is separated from the protoconid by an indistinct notch. At the distal crown surface a low sharp cusp is situated posterior to the protoconid, separated from it by a distinct notch, which passes into a shallow depression on the lingual side of the tooth. Behind the posterior cusp is another low indistinct accessory cusp attached to the cingulum which is distinctly developed, especially on the lingual side of the tooth.

The measurements: the tooth length is 14.3 mm, tooth width is 8.9 mm, and crown height at the protoconid is 8.8 mm.


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3. The left calcaneus has a fl at knob (tuber calcanei) divided by shallow depression on the top with a larger medial segment and lesser lateral segment. On the lateral and medial sides of the shaft below the knob, a distinct tuberosity is present; it continues on the dorsal and plantar-distal sides. The calcaneus shaft extends slightly from the knob towards the distal surface; its plantar side is broad and faintly concave, whereas the dorsal side is narrow becoming crest-like. Three areas for articulation (facies articulares talares) are present on the dorsal side. The largest articulation is situated on the damaged blunt coracoid process. The second largest articulation is circular in shape and is situated on the third and sustentaculum tali. These two articulations are separated by a distinct narrow sulcus calcanei. Below this sulcus, the smallest articulate area is situated which is medially oriented and damaged. Sustentaculum tali extends towards the medial side, it is triangular in shape with a faintly round top. On the plantar side, the sustentaculum tali is separated from the calcaneus body by a deep, distinctly limited muscular furrow (for the sulcus tendinis m. fl ex. digiti I longi). Two distinct crest-like rugosities are situated on the lateral side near the bone base, limiting the narrow sulcus calcanei. Base of the calcaneus consists of a large articulate surface for the central tarsus bone (os tarsi centrale).

The measurements : maximum length is 62.3 mm, width of the proximal part of calcaneus is 16.5 mm, width

of the distal part is 20.9 mm, maximum width is 23.5 mm, maximum diameter is 24.0 mm, maximum length of the area on the sustentaculum tali is 10.1 mm.

Infraorder Arctoidea FLOWER, 1869Superfamily Ursoidea FISCHER DE WALDHEIM, 1817

Family Ursidae FISCHER DE WALDHEIM, 1817

Ursidae gen. et spec. indet.(pl. 3, fi gs 4–6; table 3)

Mater ia l : front premolar (upper right P1 or ? lower left p3) (GMM, B-4077) from probe 2/97 of Hajnáčka I.

Descr ipt ion: Crown of the front premolar (probably P1 or p3) is complete. It is almost unabraded and slightly damaged at the base of the distal margin. A blunt cusp (paracone/protoconid) with two distinct crests dominates the middle part of the crown. The anterior and posterior crests continue from the cusp top to the base of the crown. The cusp passes into a shallow depression or shelf on the distal-lingual side of the tooth limited by the cingulum that is well developed especially on the lingual base of the crown. The root is preserved, slightly thinner on the distal and lingual side.

Table 2: The measurements of Pliohyaena

sites teeth etc. Val´d Arno Les Étouaires/Perrier Gundersheim St. Vallier

p4 L B

23,113,5

2314

23,9 24,6 23,3 14,5 14,7 15,0

--

23,3 22,4 22,7 22,7 23,5 15,0 14,9 14,3 14,8 15,0

p3 L B

20,513,6

2214

20,3 22,2 20,5 14,9 15,1 14,8

2115

20,5 20,8 20,9 20,4 20,9 14,8 15,0 14,8 14,5 14,9

p2 L B

14,710,5

1610

15,2 14,7 17,4 10,3 11,2 11,6

1610

15,2 15,2 15,3 16,0 15,7 10,3 10,3 9,8 10,2 11,2

c inf. sag. trsv.

18,513,7 - 18,3

14,91713

17,7 17,7 17,4 14,7 15,0 13,6

height of the mandible behind m1 - 48 43 - 45

height of the mandible below p4 38,5 44 39 - 40

height of the mandible below p3 19,5 16 17 - -

length of diastema 10 11 10 ? 8 12

sites teeth etc. Villaroya Yushe, China Nihovan

China p4 L B

23,113,5

23,0 24,0 13,6 -

22,8 22,6 14,5 14,6

--

p3 L B

20,513,6

20,8 21,6 13,5 13,7

20,1 20,3 14,0 14,1

21,310,9

p2 L B

14,7 14,310,5 8,9

15,0 17,0 9,4 10,1

15,2 14,7 11,1 11,1

22,715,2

c inf. sag. trsv.

18,513,7 - 16,9 17,3

13,6 14,0 -

áHajn ka

áHajn ka


23

T h e m e a s u r e m e n t s : the length = 6.2 mm, the width = 4.5 mm, and the crown height at the main cusp = 3.4 mm.

Discuss ion: From the morphological point of view, the bear front premolar is similar to the left p3 of Ursus minimus, published by DEPÉRET (1892: pl. XI fi gs 2 and 2a), where this premolar is fi gured in the fragment of the left mandible. However, the premolar is closer in the shape to the right P1 of the species Ursus (Selenarctos) thibetanus (skull of the Tibetan Black Bear male, No. 64935, Senckenberg Museum in Frankfurt a. M.). Also, the measurements of the Hajnáčka ursid premolar corre-spond to measurements of this recent Tibetan Bear. There are only small differences between both these teeth in the shape of the crown base, which is more concave on the lingual side of the Tibetan Bear P1 than by the bear front premolar from Hajnáčka. The shape of the posterior part is different too, when the paracone of the Ursus thibetanus P1 faintly passes into the distal cingulum, but the main cusp of the Hajnáčka fossil passes into a shallow depres-sion or shelf resp. Cingulum of the recent bear P1 is more robust as well. The Hajnáčka bear front premolar differs from other preserved unabraded front premolars of the measured Ursus thibetanus skull either in its morphology (P3 and p3) or measurements (p1 and p3).

The fossil representatives of the Tibetan Black Bear (U. thibetanus mediterraneus) have been present in the territory of Slovakia and elsewhere in Europe from the beginning of the Biharian (MQ 1 zone), and during the Villányian (MN 17 zone). However, the age of the Hajnáčka I site is the Early Villányian (MN 16a zone). Two Pliocene bear taxa – Ursus ruscinensis and Ursus minimus – are important for species determination of the Hajnáčka fi nding. Both of these taxa are placed at the beginning of the so-called black bear phylogeny. Though, the species U. ruscinensis is a typical element of the older mammal assemblages from the Ruscinian (MN 15 zone). On the other hand, the species U. minimus is a typical representative of the Late Pliocene mammal assemblages, especially of the Villafranchian (MN 16 zone), known from

Table 3:The measurements of Ursus

Ursus thibetanus Ursidae sp. Ursus minimus Erdbrink,

1953

Ursus etruscus

Erdbrink, 1953

Erdbrink, 1953

No. 64935 Seckenberg

Museum B-4077

length 4.8 - 9.0 5.0 - 7.0 4.5 - 7.0 6.3 - 6.4 6.2 P1width 3.0 - 5.3 3.0 - 5.0 3.5 - 5.5 4.5 4.5 length 6.5 - 7.0 5.0 - 7.0 3.5 - 6.0 4.0 - 4.8 - p3 width 4.5 4.0 - 5.0 2.5 - 4.5 3.2 - 3.8 -

áHajn ka

many European and Asian sites (Montagne de Boulade, Perrier, Montpellier, Perpignan (ERDBRINK 1953); Chagny (ARGANT 1991); Senéze; Osztramos 7, Kisláng (JÁNOSSY 1986); Odessa, Kvabebi, Karakum, Kuruksaj (GROMOVA & BARANOVA 1981) etc.). Also, the measurements of the U. minimus P1 and p3 (in ERDBRINK 1953) correspond to that of the Hajnáčka bear front premolar.

Besides this Pliocene bear species, the front premolar measurements of U. etruscus, especially those of p3, cor-respond to measurements of the Hajnáčka bear, too. How-ever, U. etruscus evolved during the Early Pleistocene, or during the Plio-Pleistocene period (but some authors mention the fi rst appearance of this bear during the Late Pliocene). It is a typical representative especially for the Early Biharian fauna assemblages. Thus, a circle of the potential bear species is narrowed to only one taxon –U. minimus. However, on the basis of the relatively faint morphologic diversity of ursid front premolar crowns and weak morphologic features of the Hajnácka bear, the pos-sibility of assigning this fossil to the another bear taxon is not excluded. Accordingly, the ursid front premolar from Hajnáčka is determined only to taxon Ursidae gen. et spec. indet.. In the event that this premolar belongs to the species Ursus minimus, it will be earliest fi nding of this bear species in the Hajnáčka fauna and in the territory of Slovakia too.

Family Mustel idae FISCHER DE WALDHEIM, 1817Subfamily Lutr inae BONAPARTE, 1838

Tribe Lutr ini BONAPARTE, 1838Genus Lutra BRUNNICH, 1772

Lutra cf. bravardi POMEL, 1843(fi g. 1:1, fi g. 2:1–4, fi g. 3, fi g. 4:1, 3,

pl. 3: fi gs 1–3, 7–11, pl. 4: fi gs 1–3, pl. 5, tab. 4–9)

1964 Lutrinae sp. (aff. Cyrnaonyx antiqua (BLAINVILLE)). – FEJFAR: 23.1985 Lutra cf. bravardi POMEL. – FEJFAR & HEINRICH: 221.1990 Lutra cf. bravardi POMEL. – FEJFAR, HEINRICH & HEINTZ: 52.

Mater ia l : 1. partial skeleton of a single individual from Hajnáčka I (probe 3/56) consists of broken skull fragments,


24

right fragment of upper jaw with P4 and M1 in situ (SNM-MNH, Z 26374/3), left fragment of upper jaw with P4 and M1 in situ (SNM-MNH, Z 26374/4), right fragment of lower jaw with c and p3 – m2 in situ (SNM-MNH, Z 26374/1), left mandible fragment with c and m1 – 2 in situ (SNM-MNH, Z 26374/2), left C (SNM-MNH, Z 26374/10), right P 2 (SNM-MNH, Z 26374/8), left P 2 (SNM-MNH, Z 26374/7), right P 3 (SNM-MNH, Z 26374/5), left P 3 (SNM-MNH, Z 26374/9), part of backbone (min. 16–17 vertebras) (SNM-MNH, Z 26374/23), right humerus (SNM-MNH, Z 26374/18), left humerus (SNM-MNH, Z 26374/17), right ulna (SNM-MNH, Z 26374/19), left ulna (SNM-MNH, Z 26374/20), right radius (SNM-MNH, Z 26374/21), left radius (SNM-MNH, Z 26374/22), right scapholunatum (SNM-MNH, Z 26374/15), right Mc I (SNM-MNH, Z 26374/16), right Mc II. (SNM-MNH, Z 26374/11), right Mc III (SNM-MNH, Z 26374/12), right Mc V (SNM-MNH, Z 26374/14), and left Mc III (SNM-MNH, Z 26374/13); 2. fragment of the left mandible branch with p4 and m1 in situ, and m2 alveolus (GMM, No. B-3598) from Hajnáčka I (probe 5/98).

Descr ipt ion: Fossil remains, which have been found at the bottom of an erosive ravine near probe No. 3/1956, probably belong to one male individual. The skeleton was washed out from an original position in the nearby fos-siliferous sediments. From several skull fragments only fi ve elements have been exactly determined – left auditory bulla and part of right one, part of robust left zygomatic arc and part of right zygomatic arc, which is still situated in a bit of the original sediment, and a posterior skull fragment, consisting of low sagittal crest, part of both parietals, and part of the occipital bone.

Upper dent i t ion: right and left fragment of an upper jaw consists of part of a maxilla with P4 and M1 (fi g. 1: a, b, pl. 4, fi gs 1–3), and posterior portion of the right palate. The upper premolars are nearly unworn, especially a dam-aged left P4, its lingual part with protocone being broken off. Distinct paracone has three crests – two anterior and one posterior. The fi rst of anterior crest runs antero-lin-gually from the top and the second crest runs anteriorly toward a small blunt parastyle in the antero-buccal margin of a triangular crown as part of the anterior cingulum. The posterior crest (the “cutting edge”) is orientated oblique-longitudinally, running from the top toward the metacone. The metacone is a small blunt cusp in the postero-buccal crown margin, separated from the paracone on the buccal side by a distinct valley, which is either damaged (right P4) or faintly abraded (left P4). A low protocone is included in the antero-lingual margin of the crown as a crest-like wall running from the protocone toward the metacone; it forms the lingual border of one or two shallow depressions situ-ated between the protocone, paracone, and metacone. The depressions are separated from each other by a transverse wall. The external margin of the crown consists of a low cingulum visible in buccal view (pl. 4, fi g. 1). Also, an

indistinct cingulum is situated below the parastyle of the right premolar. Two roots are present, the posterior one is more robust.

The both fi rst upper molars are complete and show typi-cal fl at wear. The joint paracone-metacone is developed as a wide blunt medio-posterior crest. The protocone is a fl at, abraded, narrow longitudinal/parasagittal cusp elongated longitudinally/parasagittaly and connected mesially with an abraded accessory cusp (= paraconulus). The distinct lingual and posterior cingulum includes a blunt metaconu-lus on the postero-lingual crown margin. Also a distinct cingulum developed on the buccal side of the tooth forms a marked antero-buccal parastyl near the paracone. The central part of the crown consists of a shallow depression clearly separating the buccal cusps/crests from the lingual ones. The broad lingual root is transversely compressed.

Besides upper jaw fragments with teeth, some isolated upper teeth of this same individual have also been found. The top of the left upper canine (pl. 3, fi gs 7, 8) is fl at abraded. The canine crown contains two crests – distal and lingual. They run from the top towards the thicker crown base. Between both crests, one shallow ridge is well developed, also running from the top towards the crown base. On the lingual side a distinct cingulum forms the crown base. In mesial view, the canine shape ia a slightly elongated S-shape.

The second upper premolars are preserved only as fragments. They consist of either the anterior part with paracone and front root (right P2) or an almost complete crown with the posterior root (left P2). The paracone is fl at abraded on top, from which three crests (anterior, posterior, and lingual) run toward the crown base. A damaged cingu-lum was developed around the crown base. The anterior root of right P2 is slender, anteriorly bowed and tapering to the base, where it is again faintly expanded. The posterior root of left P2 is robust, posteriorly bowed and tapering to the base and there again slight expanded.

Only the right P3 is preserved completely together with a bit of bone tissue between the roots. The left P3 consists of rest the crown part and posterior root. From the morphological point of view, these teeth are similar to P2 but are larger. The crown is oval in shape, with greatest width in the central part. A prominent character is a dis-tinct paracone, anteriorly situated. Three crests (anterior, posterior, and lingual) are developed on its surface from the cusp apex toward the crown base formed by a distinct cingulum. The mesial margin of the crown has an indis-tinct blunt accessory cusp between the lingual and buccal connection with the cingulum. The anterior root is slender and longer than the robust posterior one. The directions of the roots diverge: anterior root is bowed anteriorly, the posterior root is bowed posteriorly.

Lower dent i t ion: All lower teeth (fi g. 4, pl. 3) are preserved in three mandible fragments. Both complete ca-nines, right and left, of the same individual are fl at abraded on the top, and slight bowed posteriorly with an indistinct


25

crest on the antero-lingual and lingual side. The crown is slightly wrinkled on the labial surface, with a distinct cingulum on the lingual and distal side. The canines are slightly bowed S-like. A single circular alveoli of the both right and left p2 are present; their position indicates that these premolars had a oblique orientation (the s.c. “coulisse position”) in the tooth row. The average size of the right p2 alveolus is 2.1 mm.

Only the right p 3 is preserved. Its crown is slightly expanded centrally; mesially, a distinct protoconid is lo-cated with a weak anterior crest directed toward a small accessory cusp located on the anterior cingulum. Lingual and posterior crests are better developed, descending from the protoconid apex toward the crown base, where they merge with distinct lingual and posterior cingula. The buc-cal cingulum is interrupted in the center of the tooth. The left p3 has only a posterior root and a damaged anterior alveolus preserved.

The largest lower premolar is p4; its occlusal outline is oval to fi gure 8 shape, caused by a slight constriction between the mesial and larger posterior segment. A fl at abraded protoconid is the principal cusp; its anterior crest descends toward an indistinct accessory cusp on the front cingulum; its posterior crest descends from a small blunt accessory cusp on the posterior side of the protoconid toward the crown base, where it passes into a narrow and shallow depression separating the protoconid and posterior cingulum with a small accessory cusp. A third crest de-scends on the lingual side toward a small blunt accessory cusp on the lingual cingulum. The low buccal cingulum is in the middle interrupted. Two roots are preserved, the posterior one is more robust.

The m1 is the largest lower cheek tooth. A conical paraconid is located in the anterolingual area of trigonid. On the buccal side of the paraconid, a crest ascends toward protoconid terminating as distinct a notch, that separates the paraconid buccal crest from the short blunt anterior crest of the protoconid. A blunt ridge is located on the lingual side of paraconid, descending from the apex to the base of the paraconid. A distinct crest descends from the apex of the protoconid medially to the metaconid. On the posterior side of the protocinid a short blunt crest near its base is directed labially toward a small indistinct blunt accessory cusp. Metaconid is approximately as large as the paraconid, but faintly taller; it has two crests, a medial crest directed toward the protoconid and a posterior crest directed lingually toward an indistinct notch that sepa-rates the metaconid from the talonid. Similarly as distal protoconid, the posterior metaconid wall perpendicularly directs to the talonid basin, forming a distinct sheer face separating the trigonid from the talonid. Trigonid is opened to the lingual side. Only the hypoconid and hypoconulid are well developed on the talonid, the hypoconid is a low cusp abraded longitudinally on the buccal side to became crest-like. The hypoconulid is smaller than the hyopoconid and located posterior to it, on the posterolabial marin of the tooth.

The entoconid and entoconulid are not developed, or indistinct on the crest-like lingual wall of the talonid. Shape of the talonid semilunate to oval. A labial cingulum is developed on two areas below the paraconid/protoconid notch and along the hypoconid, with well preserved ce-mentum fi rmly attached to the enamel walls. In occlusal view, the buccal shape of the crown is a pointed semilunar arch from paraconid over the both protoconid and hypoco-nid to hypoconulid.

The small m2 is typically oriented diagonally relative to the occlusal level. The occlusal outline is triangular to circular. The paraconid is a small indistinct cusp in the front of the crown. The protoconid and larger metaconid are better developed than the paraconid. Talonid cusps are reduced; in forms a semicircular low crest from the protoconid to the metaconid. A cingulum is faintly hinted on the buccal side. The m2 forms a cylindrical shape. It has a single root.

Both mandible branches are damaged, with only fragments of mandible preserved. Canine and molars are present only in the left mandible, the third and fourth premolars are preserved in the right mandible. Two ap-proximately same large oval mental foramina are pre-served on the right dentary together with the front part of a shallow masseteric fossa and lesser mandible foramen preserved on the lingual side. The anterior mental foramen is located below p3 and slightly higher than the posterior foamen, which is located below p4. Fragments of the right and left posterior dentaries are preseved; they consist of a damaged cylindrical condylar process and part of the coronoid process.

A left mandible fragment found in probe No. 5/98 (pl. 3, fi gs 10, 11) probably belongs to another, smaller individual of the same taxon (female?). Slight abrasion of teeth in the mandible indicate a young adult individual. Front of this left mandible fragment is broken off from the incisors to p4. Below this p4, one of mental foramina is located. The coronoid process together with the condylar and angular processes are also missing. Only the anterior part of the masseteric fossa and the mandible foramen are preserved on the lingual side of the posterior mandible. An indistinct blunt ridge is preserved in the posterior end in front of the vascular notch tips of the mandible body. Also, a shallow depression is located between m1 and m2 alveolus on the buccal side of the mandible fragment. Average of length the alveolus is 3.7 mm and the transverse width is 2.6 mm.


26

Table 4: Measurements of the lutrine lower dentition and dentaries from Hajnáčka

Lutra sp. 1 Lutra sp. 2Ivanovce Ivanovce

Z 26392 Z 26393 Z 26374/1 Z 26374/2 B-3598 length measured from tip to root tip 25.0 - 22.5 - - longitudinal average of the crown base 7.1 - 6.8 6.4 - transverse average of the crown base 5.3 - 4.8 4.9 - crown height 9.8 - 8.5 8.6 - root length in front 17.0 - 16.2 - -

c

root length at the back - - 13.4 - - maximum length 4.5 - - - - maximum width 2.8 - - - - p2

crown height in the place of the protoconid 2.9 - - - - maximum length 5.8 - 5.8 - - maximum width 3.2 - 3.2 - - p3

crown height in the place of the protoconid 3.5 - 3.3 - - maximum length 6.7 6.0 8.0 - 6.2 maximum width 3.6 3.0 3.8 - 3.5 p4

crown height in the place of the protoconid 4.7 4.0 4.2 - 3.9 tooth length 12.0 11.4 13.9 14.2 10.9 trigonid length 6.7 5.4 6.7 6.2 5.5 talonid length 5.3 5.0 6.7 7.3 4.4 width at the back reduction of the tooth 4.9 4.8 6.5 - 5.4 crown height in the place of the protoconid - 5.1 5.4 5.8 5.0 width in the place of the protoconid 4.7 4.4 6.3 6.2 5.4 width in the place of the hypoconid 5.0 4.8 7.2 6.9 5.0 front part length on the buccal side 8.5 8.4 8.6 - 8.0 back part length on the buccal side 4.5 4.2 6.6 - 4.0 paraconid length 2.8 3.1 4.0 3.9 3.4 metaconid length 3.5 2.8 3.2 - 4.2 paraconid height 2.9 3.8 3.5 3.8 4.0

m1

metaconid height - 3.0 4.8 5.1 3.6 tooth length - 3.8 5.8 5.7 - m2 tooth width - 3.5 5.2 5.3 - length of c - m1 35.3 - 37.4 37.3 - length of c - m2 - - 40.7 41.2 - length of p2 - m1 26.7 - 28.3(a) 28.4(a) - length of p4 - m1 17.3 17.4 21.2 - 17.6 length of p3 - m2 - 25.4(a) - - - length of p4 - m2 - 20.0 24.9 - 21.4(a) maximum mandible height - 30.8 - - - height between condylus and angular process - 11.8 - - -

mand.

mandible height in the place of m1 12.9 10.5 11.8 12.4 11.6

Lutra cf. bravardi, Hajná ka


27

Table 5: Measurements of the lutrine upper dentition from Hajnáčka

Lutrini sp., Ivanovce Lutra cf. bravardi, H

Z 26394 Z 26395 Z 26374/3 Z 26374/4 Z 26374/10 Z 26374/7 Z 26374/5 Z 26374/9

length measured from tip to root tip 28.6 29.0 - - 25.0 - - - longitudinal average of the crown base 7.0 6.4 - - 5.8 - - - transverse average of the crown base 4.8 5.0 - - 4.8 - - - crown height 12.0 12.7 - - 9.6 - - - longitudinal average of the root 7.1 6.3 - - 6.0 - - - transverse average of the root 5.0 4.6 - - 4.6 - - - root length in front 17.0 18.6 - - 6.6 - - -

C

root length at the back 15.3 15.7 - - 14.8 - - - maximum length - - - - - 4.9 - - maximum width - - - - - 3.0 - - P2

crown height in the place of the paracone - - - - - 4.0 - - maximum length - - - - - - 6.8 - maximum width - - - - - - 3.9 3.9 P3crown height in the place of the paracone - - - - - - 4.6 4.4 maximum length - - 11.4 11.6 - - - - maximum width - - 8.5 - - - - - crown height in the place of the paracone - - 6.0 6.0 - - - -

P4

paracone length - - 7.7 7.6 - - - - maximum length - - 8.7 9.2 - - - - width of the front part - - 9.7 10.7 - - - - width of the back part - - 8.9 9.2 - - - - width in the central part - - 9.4 9.6 - - - - medial length of the front part - - 4.0 3.6 - - - - medial length of the back part - - 3.5 4.0 - - - - paracone length - - 4.8 4.5 - - - - metacone length - - 3.7 3.9 - - - - paracone height - - 2.0 2.1 - - - -

M1

metacone height - - 2.6 2.8 - - - -

áajn ka

The postcranial skeleton (f igs 3 , 4 , pl . 5) : Pre-served vertebrae consist of minimally from 16 to 17 ver-tebrae. Four (two pairs) of them are situated in the original position, suggesting little dorso-ventral postmortal trans-port. Single vertebrae are preserved as fragments in four vertebrae, parts of zygapophyses are also preserved.

The right and left humeri (fi g. 3: 1a,b, pl. 5, fi gs 1–4) are complete. The humerus head has a robust beak-like shape, limited by minor tubercle on the medial side. A dorso-medial surface of this tubercle is coarsened. The tuberculus maior area forms a wide wall on the latero-cranial side of the head; it is not as well developed as the tuberculus minor area. An oval area for attachment of the infraspinatus muscle is located slightly below the caudal beginning of the greater tubercle. Below this area, a tuberosity for attachment of the musculus minor teres is located. The intertubercular sulcus is shallow and wide.

The humerus body is relatively smooth, cranially bowed and faintly tapering. Posteriorly on its cranial side, the crest of the greater tubercle is located. The crest is lim-ited by an indistinct deltoid tuberosity along the lateral side and by shallow sulcus on the medial side, which is still limited by two distinct short ridges. The distal end of humerus has well developed trochlea with a deep longi-tudinal olecranon fossa, a robust medial epicondyle, and a small lateral epicondyle. Above the medial epicondyle, a longitudinal supratrochlear foramina are located. From lateral epicondyle, a robust faintly bowed crest is directed medio-dorsally from the lateral epicondyle toward the shaft above the distal end of humerus. The radius fossa is in-distinct. A large longitudinal depression is situated on the right humerus below the supratrochlear foramen on the cranial side, whereas on the left humerus only a tuberosity is located in this area.


28

Measurements of the humeri from Hajnáčka: 1 – maxi-mum height on the medial side is 66.3 mm (right) – 67.4 mm (left), 2 – maximum height on the lateral side is 64.3 mm (right) – 66.2 mm (left), 3 – width of the proximal part is 18.9 mm (right) – 19.0 mm (left), 4 – width of the distal part is 23.1 mm (dex.) – 23.3 mm (sin.), 5 – medio-lateral width of the bone body in the middle is 7.1 mm (right) – 7.0 mm (left), 6 – maximum trochlea length is 15.0 mm (right) – 15.6 mm (left), and 7 – width of trochlea in the middle is 6.7 mm (right) – 6.8 mm (left).

The right ulna (fi g. 4: 1a,b; pl. 5, fi gs 7, 8) is completely preserved, whereas only the proximal body is preserved from the left ulna. The olecranon is fl attened medio-lat-erally, bowed medially, fl attened on the dorsal side of its tubercle, and faintly tapering laterally. The processus an-conaeus attachment is distinct, slightly passing the cranial margin of the olecranon. Below the olecranon, a smooth trochlea notch is located; it is tapered in the center and of semicircular shape in lateral view. Indistinct lateral and medial coronoid processus are present in the ventral margin and on the lateral and medial side of the olecranon notch. Between these processus, a radius notch with an articular area is located on the cranio-lateral side of the ulna. Body of the ulna is fl attened medio-laterally, faintly bowed, with a caudal ridge and a tuberosity on the cranial side. The distal end of the ulna consists of small a small head, which continues on the caudo-lateral side to join the semicircular fl attened styloid process with a small articular area.

Similarly, the right radius (fi g. 7: 3a, 3b, pl. 5, fi gs 5, 6) is also complete whereas only the distal part of the left radius body is preserved. The proximal end of the radius has on its dorsal side an undivided hole-like articular area. Below this articular area, a transverse articular area is present on the caudal side. The lateral ligament tubercle located on the lateral side of the head is damaged. The angular radius body is faintly bowed cranially, with sharp caudal, lateral, and medial margins. The distal end of the radius consists of a damaged carpal articular area and short caudal ridge, located dorsal to the articular area. Ventral to this caudal ridge, a circular depression is present. The styloid process is blunt and rounded on the medial side, almost spike-like on the latero-cranial side.

Table 6: Measurements of the ulnae from Ivanovce and Hajnáčka

Lutrini sp. Ivanovce Z 26396

L. cf. bravardi,Z26374/20 Z26374/19

maximum height - 67.4 - width of the proximal part - 13.0 12.9 width of the distal part - 8.9 - minimum body width - 5.8 - height of trochlea notch 10.9 9.5 9.8 width of trochlea notch in the middle 5.0 4.4 4.4

áHajn ka

Measurements of the radii from Hajnáčka: maximum height is 49.6 mm (rigth), width of the proximal bone part is 12.9 mm (right), width of the distal bone part is 9.0 mm (right) – 9.6 mm (left), latero-medial width of middle shaft is 5.1 mm (right) – 5.0 mm (left), and medio-lateral width of shaft below the head) is 5.8 mm (right).

Right scapholunatum (pl. 5, fi g. 13) is almost undam-aged. Its dorsal part is arch-like. The ventral part is divided by a distinct crest to form two large articular areas, lateral of which is further divided by an indistinct longitudinal ridge. A small process is present on the dorso-medial mar-gin. This process is tapers ventrally grading into a sulcus on the lateral side of the bone.

Measurements of the scapholunatum from Hajnáčka: maximum length is 9.1 mm, maximum width is 10.4 mm.

Of metacarpals, four right ones and one left were found (pl. 5, fi gs 9-12). The right Mc I is short, with wide dorso-plantar fl attened proximal surface, very short in the shaft, and robust distal body. The medially expanded proximal head consists of massive articulate area, extending espe-cially on the proximal and dorsal parts of the head. Hereby, its dorsal part is distinctly separated from proximal one. A robust trochlea forms the distal part. The trochlea is divided by shallow dorso-plantar area to larger medial and lesser lateral part. A distinct crest forms the medial margin of this area. On the dorsal side, a distinct tuberosity is present

below trochlea. This tuberosity almost expands towards a lesser articulate area on the medial side of the distal part, and which is distinctly separated from trochlea. The right Mc II (pl. 5, fi g. 11) is more robust developed with wider distal part. The proximal part consists of elongated articulate area, which is divided by indistinct ridge to lesser lateral and larger medial part. The area plantarly passes to beak-like shape. On the lateral side of the proximal part, one small oval limited area is still situated. On the medial side of the proximal part, a distinct tuberosity is developed below the main articulate area. The bone body is fl at, wide, dorso-plantarly compressed and faintly dorsally bowed. Distally, the body passes to trochlea, which is separated from it by distinct sulcus. Distinct cusp-like tuberosity


29

limits the trochlea along its medial and lateral side. Ad-ditionally, the trochlea plantarly expands and it is divided by distinct crest to lesser lateral and larger medial part.

The right Mc III (pl. 5, fi g. 12) is long slender bone with distinctly developed distal part (proximal part is broken off in right and left metacarpus). The body is long and slender, faintly dorso-plantarly compressed. The distal trochlea is separated from bone body by distinct sulcus, running from lateral and dorsal side to the medial. Below trochlea, massive tuberosity is present on the both lateral and medial sides. From the distal view, the trochlea is arch-like in the shape, expanding plantarly, and divided by distinct ridge (crista sagittalis) to two approximately same large parts.

The right Mc V (pl. 5, fi g. 9) is short fl at bone, which is larger than Mc I. Its proximal part is faintly extended. A shallow articulate area dominates in this bone part. The area dorso-plantarly expands and is separated by blunt marginal crests from two small articulate areas on the lateral and medial side of the proximal part. On the medial side, one small circular articular area is present below main articular surface. The body is short, fl at, wide, dorso-plantarly compressed, distally passing to trochlea, which is separated from it by shallow notch. The trochlea is relatively low and wide, expanding plantarly and divided by distinct crest to two almost same large parts.

Table 7: Measurements of metacarpals from Hajnáčka

Z 26374/15 Z 26374/11-14

right MCI right MCII right MC3 left MC3 left MC5 maximum length 10.4 20.3 - - 13.8 maximum width of the proximal part 5.7 4.6 - - 4.0 maximum width of the distal part 3.3 5.1 5.2 5.1 4.0

Discussion: The records of dentary fragments of fossil lutrine show some differences from the dentaries of the recent European otter (Lutra lutra LINNÉ, 1758). Dentaries of the fossil otters from Hajnáčka are more robust than these of recent species. On the other hand, the measure-ments of extant European otter p4 and m1 are larger, and teeth of the recent otters are more robust. Together, the anterior margin of the masseteric fossa in fossil otters is more widespread below m2. The mandible foramen of the fossil lutrine remains is located nearer to m2 than it is in L. lutra. Thus, the lutrine records from Hajnáčka do not belong with the species Lutra lutra, but they represent other species of the genus Lutra which might belong with high probability to Lutra bravardi from the Late Pliocene from Perrier-Les Étouaires. Also, the measurements of the teeth and dentary fragments of the fossil indicate the pos-sibility of sexual dimorphism or of individual variability in the frame of one species. Contrary to Lutra sp. 1 and 2 from Ivanovce a small accessory cusp is developed at the posterior ridge of the protoconid in Lutra lutra, too. Our comparisons with the extant species of Lutra sumatrana GRAY, 1862 (fig. 1: 3 and fig. 2: 9) indicate relationship for the Pliocene Lutra bravardi as its possible ancestor.

Comparisons of the fossil Lutra bravardi from Per-rier-Étouaires (based on the cast of the Type) and Lutra cf. bravardi from Hajnáčka with the extant European otter Lutra lutra show following differences:

Table 8: Tabellar comparison of the features of lower dentition

features Lutra bravardi (PE) Lutra cf. bravardi (Ha) Lutra lutra with acc. basal cusps without acc. basal cusp

posterior cristae of p3, p4 more developed, run parallel along depression weak, divergent, no post. depression cingulum of p3, p4 laterally interrupted, weak, continuous,

m1 similar shape as Ex, morphology, basin of talonid deep basin of talonid shallower

buccal cingulum of m1 similar shape, + outer cementum (HA) no outer cementum ! lingual cingulum of m1 no developed along paraconid only

nterior crista of p3, p4 a


30

Lutra sp. I.(pl. 6, fi gs 1–3, table 4)

1985 Lutra cf. bravardi POMEL. – FEJFAR & HEINRICH: 221.

Material : incomplete right mandible with c, p2 – p4, and m1 in situ (SNM-MNH, Z 26392) from Ivanovce.

Description: incomplete fragment of right dentary with unabraded teeth: canine, p2–p4, and m1. The canine is slightly fl exed with an indistinct crest on the dorsal and lingual side, wrinkled on the surface near the crown base with a distinct basal cingulum. The canine has a character-istic S-like shape, in lateral view its root is partly exposed. The second lower premolar is the smallest, with an oval occlusal outline. The dominant protoconid is situated in the anterior part of the crown. A weak cingulum forms the crown posterior margin. Similar as in the lower jaw from Hajnacka, the premolar is oriented oblique to the tooth row because of little space, ousted to the lingual side in a “coulisse position”. The p3 is larger than p2, with crown slightly expanding toward the posterior side. The protoconid of p3 is located medial to the midline, the cusp is blunt in the top with weak anterior and posterior crests, which descend from the protoconid apex till to the crown base. A cingulum is well developed on both the mesial and lingual sides. The lingual side of both the p2 and p3 are slightly wrinkled on the surface. The p4 is the largest premolar. The posterior part of its oval crown is wider than anterior part. The protoconid dominant is the cusp, located more centrally not as anterior as in ront premolars. It has two weak blunt crests (one anterior and one posterior) descending from cusp apex toward the crown base. A cingulum is well developed around the base of the crown base.

In the fi rst lower molar a small paraconid is present anteriorly. The paraconid is connected with the protoconid by weakly abraded crest on the buccal side of the tooth. Protoconid (its top is broken off); it was connected with the metaconid by a crest on the lingual side. The metaconid is of approximately the same shape as the paraconid. All three cusps of the trigonid limit the trigonid lingual depression. The hypoconid is low separated from the protoconid by a small notch. Posterior to the hypoconid, an indistinct abraded hypoconulid is located at the posterobuccal margin of the crown. The entoconid and entoconulid are not de-veloped, being a part of the narrow crest-like wall forming the lingual and posterior margin of the talonid. The talonid is a shallow semilunate depression. A weak cingulum is present on the buccal side, the mesial margin and on the anterior part of the lingual side of the tooth. A broken root of m2 is preserved in a damaged alveolus. The root shows an oblique orientation of this molar in the tooth row.

A fragment of the dentary is preserved, with front margin of the external temporal fossa.

Discuss ion : In comparison with Lutra cf. bravardi from Hajnáčka, the dentary of Lutra sp. 1 from Ivanovce is longer, the tooth row is less compressed, but teeth are lesser robust, especially m1 is smaller and more narrow. However, the canine of Lutra sp. 1 is more robust than in the Hajnáčka specimen. Also, accessory cusp at the posterior ridge of the p4 protoconid is missing.

Lutra sp. II.(pl. 6, fi gs 4–6, table 4)

Mater ia l : incomplete left dentary with p4 – m2 (SNM-MNH, Z 26393) from Ivanovce.

Table 9: Tabellar comparison of the features of upper dentition

features Lutra bravardi (PE) Lutra cf. bravardi (Ha) Lutra lutra

M1

paracon and metacon low, in one line, less “cusp-like”, not separated→ buccal outline straight; wear of pa + me →lingitudinal facet (Ha); anterior crest of protocone with protoconulus and parconulus (Ha) - in PE lesser indicated (juvenil!)

paracon and metacon well differenciated and separated→buccal outline nicked

talon and cingulum in

M1

talon less expanded, in lingual cingulum low hypocon on the postero-lingual edge, small metaconulus expands the distal border, anterior sharp cingular arm up to the mesial protoconulus (Ha). Buccal cingulum increase at paracon→small depression ! (Ha,PE)

talon more expanded, hypocon (=postero-lingual cingulum) splitted in cuspules; metaconulus sometimes indicated; lingual arched cingulum fine denticulated (juv.!). Buccal cingulum less indicated

P4: triangular outline; low paracon-metacon not lingually inclined→carnassial edge short, less effecive. Protocone low indicated (PE). Low buccal cinglum, well indicated at parastyl PE), with outer cementum (Ha).

linguiform talon expanded, circular outline with cingular rim; high paracon-metacon distinctly inclined lingually→longer and effective carnassial edge. Protocone invisible. Buccal cingulum missing, trace only at low parastyl.

Note: Ha = Hajnáčka, PE = Perrier-Étouaires, the Type, Ex = extant Lutra lutra


31

Descr ip t ion: The anterior part of the left dentary is broken. The unabraded teeth: p4, m1 and m2 are present. Other teeth are either lost (p3) or preserved as fragments of their roots (c and p2).

The occlusal outline of p4 is oval in the shape, wider in the posterior part. A dominant character of the crown is the protoconid located anterior to the center. From the top of the protoconid, three crests are developed: distinct anterior crest descends toward a small accessory cusp on the anterior cingulum at the mesial crown margin; a lingual protoconid crest descends posteriorly to the distal cingu-lum. Two roots are present, the posterior root is probably more robust than the anterior root.

The m1 is the largest tooth. A small paraconid is situ-ated in the anterior trigonid. On the buccal side of the paraconid, a carnassial edge descends posteriorly toward the protoconid where it is interrupted by a notch. All three trigonid cusps limit the trigonid depression, which is open on the lingual side of the tooth. A distinct but blunt crest descends oblique-medially from the protoconid apex to the metaconid where it is interrupted, by a notch. The meta-conid is a small cone shaped cusp lower than paraconid. A posterior crest from the metaconid descends toward the low lingual margin of the talonid. The talonid consists of only a hypoconid and hypoconulid, it is a shallow semi-lunate depression. Blunt posterior crest descends from the apex of the protoconid steeply toward a notch on the buccal side of the tooth, separating the protoconid from the low hypoconid on the buccal margin of the crown; the anterior and posterior crest is separated from the proto-conid and small hypoconulid by shallow notches. A weak cingulum is developed on the both buccal-mesial base of the paraconid. The small m2 has one root, it is orientated oblique relative to tooth row. The occlusal outline of m2 is semicircular in shape with reduced morphology: only both low protoconid and larger metaconid are developed, connected by low crest in the center.

Anterior part of the mandible is broken, but the poste-rior part is preserved almost undamaged. Coronoid process is massively developed, triangle-like in the shape with a relatively narrow top. The shallow masseteric fossa is fi nely limited, especially on the anterior and ventral side. The cylindrical shaped condylar process tapers obliquely to the buccal side, the angular process is broken. The mandibular foramen is small, circular, situated below the central part of coronoid process base. Only one mental foramen is preserved.

Discussion: In comparison with the Lutra sp. 1 from Ivanovce, p4 of Lutra sp. 2 is less robust and narrower, m1 is smaller, and the mandible body is not bowed, but is more straight. Compared to the taxon Lutra cf. bravardi from Hajnáčka, p4 of Lutra sp. 2 has a different shape and its protoconid is located more anteriorly, without an accessory cusp at the posterior ridge of the protoconid, m1 is narrower and longer; otherwise is together with m2 much less robust, and the tooth row was probably not as compressed.

Lutrini gen. et spec. indet.(table 4, 6)

Mater ia l : left C (SNM-MNH, Z 26394), sin C (SNM-MNH, Z 26395), fragment of right edentulous dentary (SNM-MNH, Z 26398), fragment of left ulna (SNM-MNH, Z 26396), and right calcaneus (SNM-MNH, Z 26397) from Ivanovce (fi ssure 6512).

Descr ipt ion: Both left upper canines have a distinct longitudinal crest on the antero-mesial side and one to two ridges on the posterior side or on the lingual side. A cingulum is distinctly developed on the mesio-lingual side of the crown. The root is straight, faintly tapered in the base. The canine Z 26395 is more slender and longer than canine Z 26394.

The edentulous right mandible consists of part of the central body and posterior segment. Anterior part of the deep masseteric fossa is preserved, the fossa extended below m2, of which only small alveolus is preserved. The alveolus is oriented out of the main tooth row. Also, partly exposed mandible foramen is present on the lingual side of the fragment.

Fragment of the left ulna consists of proximal and central part of the shaft. The olecranon is damaged and broken. The anconeus process of the ulna is distinct, run-ning laterally. Below this process, the smooth trochlear notch is preserved, it is slightly narrower in the central part and appears semicircular to arch-like from the lateral side. On its distal margin, a damaged lateral and larger medial coronoid process is preserved. An indistinct shallow notch with distinct articular area is located on the cranio-lateral side of the ulna between both coronoid processes. Below the medial process, a shallow longitudinal depression is preserved. The body of the ulna was probably fl attened medio-laterally, with a distinct caudal ridge.

The right calcaneus is gently fl attened transversally. The coracoid process is small and blunt. On the medial side, the semicircular sustentaculum tali has one articulate area on the dorsal side with a wide sulcus for muscles below it. On the dorsal side of the calcaneus, minimally three articulate areas are located, which tent to merge into one articular area. These dorsal articular areas are sepa-rated from the articular area on the sustentaculum tali by a distinct sulcus. In the ventral part of the lateral side, one sulcus is preserved, it separates a blunt longitudinal process on the plantar side of which an articular area is preserved. The plantar margin is relatively wide.

The measurements of the calcaneus are as follows: maximum height is 26.9 mm, maximum dorso-plantar width is 10.6 mm, and maximum medio-lateral width is 15.0 mm.

Subfamily Mustel inae FISCHER DE WALDHEIM, 1817Genus Mustela LINNEUS, 1758


32

Mustela sp.

Mater ia l : fragment of left edentulous dentary (SNM-MNH, Z 26375) from Hajnáčka I (probe 8/56).

Description: damaged toothless fragment of left dentary consists of part of body and part of the posterior part, which includes base of distinct masseteric fossa. The mandibular foramen is preserved on the lingual side only. The suban-gular process was not preserved. Only a relatively large m1 alveolus and a small m2 alveolus are present on preserved surface of the mandible body. The large m1 alveolus has anterior and posterior root. The small m2 alveolus is situ-ated higher in tooth row than the large alveolus, almost level with the anterior margin of the coronoid process base. The dentary depth below m1 is 4.5 mm.

Mustelidae gen. et spec. indet.

Material: fragment of left maxilla with P2 and P3 (SNM-MNH, Z 26391) from Ivanovce.

Descr ipt ion: A canine alveolus is partially preserved. Both preserved premolars are separated by diastema; their crowns are intact and unabraded.

The occlusal outline of P2 is oval, slightly wider anteri-orly than posterior one. A paracone is well developed with anterior and posterior crests descending from relatively blunt apex toward small accessory cusps, which are located on the anterior and posterior cingulum. The cingulum is continuous but it is better developed on the mesial, lingual and distal sides. Two roots are present.

Measurements of P2 are: length is 5.6 mm, width is 3.0 mm, height of paracone is 3.6 mm, and paracone length is 5.0 mm.

The P3 is larger than P2. The occlusal outline is oval, but wider posteriorly. A small paracone is located anteriorly, with a shorter less distinct anterior crest and longer postero-buc-cal crest. The both crests descend from sharp apex toward small accessory cusps, located on the anterior and posterior cingulum. Postero-buccal base of the paracone is wrinkled. A cingulum circles the crown base, better developed on the mesial, lingual, and distal side. Two roots are present.

Measurements of P3 are: length is 6.0 mm, width is 3.8 mm, height of the paracone is 3.8 mm, and paracone length is 4.2 mm.

Family Procyonidae GRAY, 1825Subfamily Ailur inae GRAY, 1843Genus Parailurus SCHLOSSER, 1899

Parailurus hungaricus KORMOS, 1934(fi g. 5, 1–9, 12, 13, fi g. 7: 1–6, pls 7, 8)

1917 Parailurus n. sp. KORMOS. – KORMOS: 578, 581.

1917 Parailurus hungaricus KORMOS. – KORMOS: 137 – 138 (nomen nudum)

1934 Parailurus hungaricus KORMOS: 17–18, 26, pl. I fi g. 8, 9.1961 Parailurus hungaricus KORMOS. – FEJFAR: 262.1964 Parailurus hungaricus KORMOS. – FEJFAR: 57 - 59.1985 Parailurus hungaricus KORMOS. – FEJFAR & HEINRICH: 221.1990 Parailurus hungaricus KORMOS. – FEJFAR, HEINRICH & HEINTZ: 52.2001 Parailurus anglicus (DAWKINS). – MORLO & KUNDRÁT: 173–184,

fi g. 15–30.

Material : right M1 (SNM-MNH, Z 26381; pit 9/56), left m1 (SNM-MNH, Z 26380; pit 8/56), fragment of right M1 (SNM-MNH, Z 26382; pit 9/56), fragment of trochanter major and distal part of right femur (SNM-MNH, Z 26383/2; pit 3/56), right tibia (SNM-MNH, Z 26383/1; pit 3/56), distal part of right fi bula (SNM-MNH, Z 26383/5; pit 3/56), right astragalus (SNM-MNH, Z 26383/3; pit 3/56), right calcaneus (SNM-MNH, Z 26383/4; pit 3/56), left cal-caneus (SNM-MNH, Z 26384; pit 8/56) from Hajnáčka.

Type local i ty: Hajnáčka, Slovakia

Age: Early Villányian, Late Pliocene (MN 16a)

Descript ion: The record of the lesser panda during the excavations 1955–1959 of the Geological Survey in Prague were introduced by FEJFAR (1964: 57–58). The isolated teeth belong to several individuals; the bones of the right hindlimb probably belong to one individual.

Dent i t ion: The occlusal outline of the right M 1 (fi g. 5: 1–4, 9) is a rounded triangular- trapezoid; its shape is slightly assymmetrical, the protocone shifted mesially. All major cusps are deeply worn; however the valleys between cusps are still preserved. The paracone and metacone are symmetrical. The accessory buccal styles are connected by fi ne crests, well visible in buccal view (fi g. 5: 1). The largest stylar cusp is the mesostyl separated by shallow grooves from the paracone and metacone; two other styles are present but the distal metastyl is larger. The protocone and hypocone are divided by a shallow groove. A lingual shallow serrated cingulum, distally notably limited by a posterior bold ridge, surrounds the protocone.

The second right M1 (fi g. 5: 12, 13) is incomplete, represented by the lingual fragment of an deep worn large and massive tooth with a broad lingual root, plus partial hypocone and protocone. In occlusal view the protocone is bordered by a shallow lingual cingular groove terminated posteriorly in a depression surounded by small cuspules; it corresponds to the worn groove between hypocone and protocone in the fi rst M1 from Hajnáčka.

The m1 of another individual (pl. 7, fi gs 1–3, 7, 9) has a broad talonid caused by massive hypoconid, and a narrow trigonid with a lingual gap between paraconid and metaconid. The protoconid is the largest and tallest cusp of the trigonid. The paraconid is the smallest cusp of the trigonid and is displaced lingually. The trigonid has a mesio-buccal short cingulum shown in the buccal and anterior view (pl. 8, fi gs 1, 9). Above the cingulum extends


33

the oblique carnassial cutting adge with a central notch pro-duced by the opposite ridges of procotonid (anteriorly) and paraconid (posteriorly). A similar lower notch is located between paraconid and metaconid opening the trigonal basin lingually. The posterior crests of both protoconid and metaconid are symmetrically doubled; the medial opposing crests meet in a small notch closing posteriorly the troginid basin, the distal crests fall obliquely into the groove divid-ing trigonid from talonid. This groove is lingually barred by a blunt central metaconulid and by the postero-lingual

crest of metaconid. The talonid is dominated by the volu-minous hypoconid. Its buccal circular-conic steep wall is symmetrically surrounded by a cingular groove visible in the distal view (pl. 7, fi g. 7); its narrowest point is on its buccal edge. The less steep lingual wall of the hypoconid is ends in the assymmetrical talonid V-shaped basin well connected with the cingular groove. Three cusps form the lingual border of the talonid: the anterior entoconulid, the middle entoconid and the posterior hypoconulid separated by trenches oriented radially toward the hypoconid.

Table 10.: Tabellar comparison of the features of M1 of Parailurus

Features Parailurus hungaricus Parailurus anglicus (Wölfersheim) buccal styli

(in buccal view) more stressed (mesostyl !) less developed

postprotocrista interrupting the ling. cingulum distally terminates in the cingular rim

lingual cingulum low depression distinct, wrinkled sharp groove flanking completely the protoconid

protocone encreased, shifted mesially → assymmetry of the occlusal outline

less encreased and shifted mesially → moderate outline assymmetry

áHajn ka) )

Table 11: Tabellar comparison of the features of m1 of Parailurus

Features Parailurus hungaricus Parilurus anglicus (Wölfersheim) lingual outline ragged straight

trigonid (in occlusal + mesial views) broad, massive narrow, moderate

paraconid larger + longer → short, smaller →→ carnassial blade → longer (expanded), mesially bowed, → shorter (reduced), straight

talonid broad, massive, outline shifted buccaly narrow, moderate hypoconid distinctly massive, “inflated” moderate

entoconid-hypoconulid (in lingual + distal views) entoconid larger than hypoconulid entoconid equals hypoconulid

talonid lingual wall (in distal view !) hihger and steeper lower and rounded

talonid cingulum regular flank around hypoconid in the buccalmost area interrupted

áHajn ka) )

Table 12: Measurements of the molars m1 and M1 of the extant Ailurus fulgens

teeth

features

m1 NMP 105�

m1 NMP10646�

m1 ZMB 3256�

m1 NMP082�

M1NMP 105�

M1NMP10646�

M1ZMB 3256�

M1NMP082�

length 11,5 11,9 11,3 11,9 9,2 9,2 9,5 9,6 width 6,0 6,3 6,2 6,0 11.0 11,8 11,0 11,2

angle A / / / / 85o 72o 70o 77o

angle B / / / / 73o 77o 72o 67o

( + = male; × = female)

(H = Hanáčka; BK = Barót Köpecz, Ru., Bo = Boyton GBr., Wö = Wölfersheim, FRG)

Features Parailurus hungaricus Parailurus anglicus (Wölfersheim) length 15,4 (H) 13,8 - 13,3 (BK) 14,5 (Bo) 14,25 (Wö) width 15,6 (H) 14,0 - 14,2 (BK) 14,5 (Bo) 14,25 (Wö)

angle A 83o 74o

angle B 61o 67o

Table 13: Measurements of the upper molars M1 of Parailurus


34

The postcranial skeleton: The right femur preserves part of the greater trochanter together with a longitudinal deep trochanteric fossa and the complete distal epiphysis (pl. 8, fi gs 7, 8, 10, 11). In the ventral view, the distal fe-mur is less anteroposteriorly deep than transversally wide. The both condyles show a gentle assymmetry: the lateral condyle is somewhat wider than the medial one and in

Table 14: Measurements of the lower molars m1 of Parailurus

Features Parailurus hungaricus Parailurus anglicus (Barót Köpecz) length 20,5 15,0 17,4 15,8 width 11,6 8,7 8,8 8,1

features P. hungaricus Ailurus fulgensmaximum width of the distal epiphysis 22,7 20,2 20,2 maximum width of intercondylar fossa 6,9 4,0 4,0 maximum width of patellar trochlea 9,4 9,4 9,8

Table 15: The measurements of distal femur

Table 16: The measurements of tibia

features P. hungaricus Ailurus fulgensmaximum length 106,8 105,0 105,0 107,0 width of proximal head 26.8 21,0 21,4 26,8 width of the shaft in the middel 7.6 5,8 5,5 8,0 width of distal end 19.7 16,0 16,0 20,0

Table 17: The measurements of fi bula

features P. hungaricus Ailurus fulgensmaximim width of the distal end 14,0 10,0 9,8 16,5 diameter of the distal end / 6,5 6,5 8,0

caudal view they are separated by a broad intercondylar fossa. The patellar groove is shallow with low trochlear ridges, the medial ridg is slightly higher and sharper than the lower and rounded lateral one. This form resemble conditions known in climbing carnivores, e.g., the viverrid Arctictis TEMMINCK, 1824.

The right tibia (pl. 8, fi gs 1–6, 9): is almost straight and relatively robust in anterior view showing a gentle torsion. The proximal half of the shaft displays anteriorly a tibial crista shifted medially and seen then in medial view

as a distinct crest joining the medial malleolus. The both condyles are each other separated by the eminentia inter-condylica with lesser lateral and larger medial intercondy-lar tubercle. The almost fl at lateral and medial condyles

are asymmetrical, the lateral one being larger and higher. Both display similar shallow depressions in the central part. In caudal view a distinct notch (incisura poplitea) separates both condyles of proximal head. The relatively wide tibial tuberosity limits the cranial and partly also lateral margin of the proximal head. Whereas the lateral shaft of the tibia is smooth, the medial area is distinctly

rough. The distal articular cochlea is anteroposteriorly low, and transversally much wider. On the medial side, the distal end is delimited by massive malleolus with a sulcus on the dorso-caudal side. The less distinct lateral malleolus has a small articular facet the for the fi bula on the lateral margin.

Only the distal part of the the right fi bula is preserved (pl. 8, fi gs 12, 13). It consists of lateral malleolus with a narrow and shallow sulcus on the lateral side and a more distinct sulcus on the medio-caudal side. Craniodistally

from the medio-caudal sulcus, one shallow malleolus sulcus is present. The fi bula base consists of an articular facet on the cranio-lateral side, which is divided into two parts by a well-marked transversal crest.


35

The right astragalus (fi g. 7: 5, 6; pl. 7, fi gs 20–23) body is broad with a moderately grooved robust trochlea; the lateral ridge is taller and more massive. Articular facets are developed on the latelar, distal and plantar sides, respec-tively. The head of the astragalus bears slender neck with

Table 18: The measurements of the astragalus

features P. hungaricus Ailurus fulgenswidth of proximal trochlea 14.0 13,0 16,3 maximum length 22,8 18,0 23,2 maximum height 12,9 8,2 11,5

Table 19: The measurements of the calcaneus

f e a t u r e s P. hungaricus right left

Ailurus fulgens

maximum length 31,8 33,9 27,0 32,0 maximum width 16,9 19,5 14,0 18,0

a distal sharp limited circular-rounded navicular facet. The calcaneal facets seen in caudal view are separated by deep groove: the lateral one is wide concave, the sustentacular facet is convex and is linked distally with the navicular facet by a narrow band.

Two calcanei (fi g. 7: pl. 7, fi gs 12– 19) of two individu-als are of relatively robust “bear-like” shape. The right calcaneus is smaller. The calcaneus tubercle is a shallow depression, divided into two approximately equal areas dorsally; the medial of these areas is taller. The body of the calcanei is narrowed distally and then it is faintly ex-

tending proximally. The plantar surface is wide and faintly concave, whereas the dorsal surface is relatively narrow to crest-like. On the dorsal side, two to three articular areas are present. The largest of the articular areas is located on the blunt beak-like coracoid process. The second largest articular area is oval to circular, located on the sustentacu-lum tali. Both articular areas are separated by a narrow sulcus. Lateral to this sulcus, the smallest articular area is located. The semicircular sustentaculum tali terminates medially, and is separated from main calcaneus body by a shallow sulcus on the plantar side. On the lateral side of the calcanei, two short crest-like ridges situated near their base constrain a narrow sulcus. A triangular articular area is present near the base on the medial side. The base is separated from a larger oval basal articularte area by a small oval depression.

Discussion: MORLO AND KUNDRÁT (2001) found after comparison of the accessible fossil material of Parailu-rus SCHLOSSER, 1899 no reason to consider Parailurus hungaricus KORMOS, 1934 as valid taxon. Therefore they “propose P. hungaricus to be a junior subjective synonym

of P. anglicus”. This conclusion was corroborated by the detailed analysis of the numerous fi nds of Parailurus in the late Pliocene layers of Wölfersheim, FRG.

To discuss this statement and to compare again the material of Parailurus from the locality Hajnáčka and

Ivanovce, Slovakia and from Wölfersheim, FRG, we have fi gured anew the important teeth from both sites in one table and in one scale using the coating with amonium chloride to reach maximum objectivity (pl. 7, 8). Here, we summerize our conclusions:

The teeth M1 and m1 from Hajnacka are in comparison with those from Wölfersheim similar in general occlusal pattern. However, the changes in morphology of the record from Hajnacka give impression of a further step in adap-tation by increase of the the size, massiveness (all main cusps, in particular: protocon, hyoconid, paraconid) and,

too, the thickness of enamel. In addition, slight increas-ing of the height of crown might be visible in distal view as to the more steep talonid lingual wall (pl. 7, fi gs 7, 8). We consider this summary of features as evidence of the validity of a taxon different from Parailurus anglicus for which the name Parailurus hungaricus KORMOS, 1934 is in our opinion still justifi ed. The difference between both species should be explained either by stratigraphy – since the level of Hajnacka is slightly younger than that of Wölfersheim (LINDSAY, OPDYKE & FEJFAR 1997, FEJFAR & REPENNING 1998), either by a related progressive con-temporaneous taxon due to the provincialism in a forested country during the warm and humid interval of the early Villanyian time (MN 15–16).

Parailurus cf. anglicus (Dawkins).(fi g. 7: 7–9, pl. 7, fi g. 11)

1961 Parailurus sp. – FEJFAR: 263.1985 Parailurus sp. – FEJFAR & HEINRICH: 221.


36

Material: distal fragment of left m2 (SNM-MNH, Z 26390) from the horizontal fi lling Nr. 6513 from Ivanovce.

Descr ipt ion: The distal part of left m2 reperesents a complete talonid broken transversally in the junction with trigonid. In a relatively small and low crown the cone of the moderately abraded hypoconid is the most prominent cusp; it is separated from other cusps of the crown by a shallow lingually shifted sagittal groove of the talonid basin. Several small cusps separated by fi ne notches (the hypoconulid and the anterior accessory cusps with the slightly abraded entoconid in the middle) are located in the posterior and lingual margin. Short lingual cingulum is indicated between hypoconid and hypoconulid. A laterally fl at compressed and buccally bent root is placed along the whole distal part of the m2.

The measurement: The width of the tooth at the hy-poconid is 7.6 mm.

Discussion: this record of Parailurus from Ivanovce is of smaller size and it probably represents the smaller taxon P. anglicus (Dawkins) which might be supported also by the higher age of the karst fi ssures of Ivanovce: the Ruscinian MN zone MN 15. The comparison with the extant taxon Ailurus fulgens in the fi g. 8 and fi g. 9, 7–9, pl. 8, fi gure 11 shows molars comparble with our material: m1, m2 and M1: The teeth of Ailurus fulgens: 1. are much smaller, 2. the smaller and accessory cusps are better expressed as higher cones and therefore nearly in one occlusal level, 3. the paraconid is shorter and steeper. – Thus, the extant Ailurus fulgens which exhibits morpho-logically more conservative only slightly modifi ed level is more similar to Parailurus anglicus than the more derived taxon P. hungaricus.

Undeterminable carnivoran fi nds from Ivanovce

In the sample from Ivanovce some no-measured tooth fragments of undeterminable carnivoran taxa are present: the paracone of M1 dex., belonging to larger, probably ursid carnivore (Ursus sp. indet. ?); a fragment of left dP4 (SNM-MNH, Z 26399/1), a fragment of right M1-2 (SNM-MNH, Z 26399/2), and a fragment of premolar (SNM-MNH, Z 26399/3) (all probably from the fam-ily Viverridae?); the lower right incisor (SNM-MNH, Z 26407/1), right 2 C (SNM-MNH, Z 26407/3, Z 26408/2), right c (SNM-MNH, Z 26407/4) (all probably from the family Mustelidae?); a fragment of the lower left incisor (SNM-MNH, Z 26408/1), and 3 fragments of carnivoran premolars.

Results and discussion

The record of carnivores from the late Pliocene sites Hajnáčka and Ivanovce comprise, i.a., fi ve species of four

families: Lutra cf. bravardi, Pliohyaena perrieri, two spe-cies of Parailurus (the larger Parailurus hungaricus and the smaller Parailurus cf. anglicus) and Hesperoviverra carpathorum. The less represented taxa as Megantereon sp. and undeterminable species of Lutra and Mustelidae are of minor importance. The remains of the lutrine from Hajnacka provide a good support for the affi nity to the late Pliocene species Lutra cf. bravardi from Perrier-Les Ètouaires as well as to the extant Lutra sumatrana which might represent a descendant relic in the (sub-)tropical environment similar to the warm conditions in Europe in the span of MN 15–17 of the mammalian age Ruscin-ian and Villaynian. Thanks to the excellent monograph of QIU ZHAN-XIANG (1987) we have now a more detailed picture on the evolution and wide palearctic distribution of Pliohyaena. The small and relatively “gracile” species Pliohyaena perrieri is part of a well documented lineage in which the initial smaller species (P. pyrenaica and per-rieri) change in time in the more robust P. brevirostris. Also Pliohyaena has a relic species in the early Pleistocene assemblage of the Chinese Choukoutien (P. brevirostris sinensis). The remains of the genus Parailurus represent another typical but rare element of the late Pliocene. The new fi ndings from Hajnáčka provide support for an own large species Parailurus hungaricus, introduced from Hajnáčka site already by KORMOS (1934). We consider here Parailurus hungaricus either as a slightly younger descendant member of the Parailurus lineage, either as an own “parallel” more robust species in the highly pro-vincial mosaic of the warm and humid conditions. The elder level of the MN 15 fi ssures from Ivanovce provided a less represented record of the smaller species Parailurus anglicus better recorded in the European late Pliocene. The large viverrid species Hesperoviverra carpathorum from the oldest level from Ivanovce (the horizontal fi lling 6523 containing an early MN 15 assemblage) belongs to the most interesting form. The Turolian “Hipparion-fau-nas” with prevailing species of the genus Ictitherium do not yet include the large viverrids. It seems that the short occurrence of the large viverrids fi lled the “gap” just since the dissappearance of ictitheriids and just before the appearance of the “true” Canids. Thus, the presence of Hesperoviverra in Ivanovce represent an exceptional record of a short term eastern migration event of a rare but successful predator which preceded the main wave of the Canidae. Within the same interval, Hesperoviverra reached north Italy (Hesperoviverra (?) apennina in the MN 16a level of Arondelli-Triversa, and east Africa (the record of Hesperoviverra (?) leakeyi in Laetoli).

Acknowledgements

The authors are deeply indebted: to Prof. Dr. Miklós Kretzoi for valuable discussions and support since the start of the research in both sites Hajnáčka and Ivanovce. To Dr. Germaine Petter (Musée Histoire Naturelle, Paris)


37

for who kindly provided the valuable cast of the Type of Lutra bravardi and measurements of Hesperoviverra (?) leakeyi from Laetoli and Hesperoviverra intertuber-culata from Yassiören. Dr. Gerhard Storch (Senckenberg Museum, Frankfurt) for the access to the material of Parailurus; Dr. Petr Benda (National Museum, Prag) and Dr. Wolf-Dieter Heinrich (Museum für Naturkunde der Humboldt-Universität, Berlin) who provided access to

the extant species of Lutra and Ailurus; to Prof. Everett H. Lindsay for support during the preparation of the paper and improving the English. The authors are indebted to the Grant Agency for Science, Slovakia, for fi nancial support (project No. 1/7304/20), and to the sponsors: The North Bohemian Browncoal Mining Company in Chomutov, Czech republik, and Tauris Co. in Rimavská Sobota for fi nancial and material support.

Fig. 1: Upper P4 and M1 of fossil and extant species of Lutra BRUNNICH, 1772. 1: Lutra cf. bravardi POMEL, 1843; right fragment of upper jaw from Hajnáčka (SNM-MNH, Z 26374/3). 2: Lutra bravardi POMEL, 1843, the Type; rigth P4 and M1, Perrier, Auvergne (MNP, Paris). 3: Lutra sumatrana GRAY, 1862; P4 – M1, Telok Betong, Sumatra, original (SNHMB, 30683 m. F. 4). 4: Lutra lutra LINNÉ, 1758; rigth P4 – M1. – 1a,2a, 3, 4: occlusal, 1b,2b: buccal views.


38

Fig. 2: The mandibles and dentition of fossil and extant species of Lutra BRUNNICH, 1772. 1–2, 3–4: Lutra cf. bravardi POMEL, 1843 from Hajnáčka. 1–2: right mandible with c, p3–4, and m1–2, (SNM-MNH, Z 26374/1). 3–4: left upper canine, Hajnáčka (SNM-MNH, Z 26374/10). 5–8: Lutra lutra LINNÉ, 1758; p4 – m2 (SNHMB, 306750 (5,7); SNHMB, 30675 (6,8, male)). 9: Lutra sumatrana GRAY, 1862; right mandible with I 1–3, c, p2–4, and m1–2, (SNHMB, 30683 m. F. 4). – 1, 4, 5, 6: buccal, 2, 7–9: oc-clusal, and 3: lingual views.


39

Fig. 3: The comparison of left humeri of Lutra cf. bravardi POMEL, 1843 from Hajnáčka (SNM-MNH, Z 26374/17) (1a, 1b) and Lutra lutra LINNÉ, 1758 (2a, 2b); caudal (1a, 2a) and medial (1b, 2b) views.

Fig. 4: The comparison of right ulnae and radii of Lutra cf. bravardi POMEL, 1843 from Hajnáčka (SNM-MNH, Z 26374/19) (1a, 1b; 3a, 3b) and Lutra lutra LINNÉ, 1758 (2a, 2b; 4a, 4b). – Anterior (1a, 2a, 3a, 4a) and lateral (1b, 2b, 3b, 4b) views.


40

Fig. 5: The upper M1 of the genus Parailurus SCHLOSSER, 1899 from the European late Pliocene. 1–4, 9, 12–13: Parailurus hungari-cus KORMOS, 1934 from Hajnáčka; 1–4, 9: SNM-MNH, Z 26381., 12–13: ingual fragment of right M1 (SNM-MNH, Z 26382). 5–8, 10, 11: Parailurus anglicus (DAWKINS, 1888) from Wölfersheim; 5–8, 10: SMF 2000/235, 11: lingual fragment of right M1 (SMF 2000/23x). – 1, 5 – buccal, 2, 6, 11, 12 – occlusal, 3, 7 – anterior, 4, 8, 13 – lingual, and 9, 10 – poterior views.

Fig. 6: The lower molars m 1 (upper line) and m 2 (middle line) upper M 1 (lower line) of the extant lesser panda Ailurus fulgens CUVIER, 1825 from the collections of the NMP (column 1, 3, 4) and ZMB (column 2). Upper line: left m1; middle line: m 2 sin.; lower line: rigth M1. – The molars, m1-2 and M1 in one collumn belong to one individual, are drawn as right-sided. – 1 – female, NMP, 10646; 2 – male, ZMB, 3256,; 3 – ?, NMP, C-082; 4 – male, NMP, C-0105.


41

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FEJFAR, O. & REPENNING, CH. A. (1998): The ancestors of the lemmings (Lemmini, Arvicolinae, Cricetidae, Rodentia) in the early Pliocene of Woelfersheim near Frankfurt am Main; Germany. – Senckenbergiana lethaea, 77: 161-194.

FEJFAR, O., HEINRICH. W.-D., PEVZNER, M.A. & VANGENGEIM, E.A. (1997): Late Cenozoic sequences of mammalian sites in Eurasia: an updated correlation. – Palaeogegraphy, Palaeoclimatology, Palaeoecology, 133: 259-288.

FEJFAR, O., HEINRICH, W.-D. & LINDSAY, E.H. (1998): Updating the Neo-gene Rodent biochronology in Europe. – In: KOLFSCHOTEN, van Th. & GIBBARD, P.L. (Eds): Mededelingen Nederlands Instituut voor Toegepaste Geowetenschappen TNO. The Dawn of the Quaternary. – Proceedings of the SEQS-EuroMam Symposium, 1996: 533-553, Haarlem.

FICARELLI, G. & TORRE, D. (1970): Remarks on the taxonomy of hyaenids.

Fig. 7: Tarsal bones of the genus Parailurus Schlosser, 1899 from Ivanovce and Hajnáčka. 1–6: Parailurus hungaricus KORMOS, 1934 from Hajnáčka. 1–2: left calcaneus (SNM-MNH, Z 26384). 3–4: right calcaneus (reversed) (SNM-MNH, Z 26383/4). 5–6: right astragalus (SNM-MNH, Z 26383/3). 7–9: Parailurus sp.; distal part of left m2 from Ivanovce (reversed; SNM-MNH, Z 26390). 1,3,6 – dorsal, 2,4,5 – plantar, 7 – lingual, 8 – occlusal, and 9 – buccal views.


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– Palaeontographica Italiana, 66, N.S. 36, 1-33.GROMOVA, I.M. & BARANOVOJ, G. I. (1981): Katalog mlekopitajuscich

SSSR. – Pliocen – sovremennost. – 456 p.; Leningrad.HELLER, F. (1936): Eine oberpliozäne Wirbeltierfauna aus Rheinhessen.

– Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 76(B): 99-160.

HOWELL, F.C. & PETTER, G. (1980): The Pachycrocuta and Hyaena line-ages (Plio-Pleistocene and extant species of Henidae). Their relationship with Miocene Ictitheres: Palhyaena and Hyaenic-titherium. – Géobios, 13(4): 579-623.

JÁNOSSY, D. (1986): Pleistocene vertebrate fauna of Hungary. – 208 p.; Amsterdam, Oxford, New York, Tokyo (Elsevier).

KORMOS, T. (1917): Die pliozänen Schichten von Ajnacskö und ihre Fauna. – Jahresbericht ungarische Geologische Reichsanstalt, (1915): 564-582.

KORMOS, T. (1934): Beiträge zur Kenntnis der Gattung Parailurus. – Mit-teilungen Jahrbuch königlich ungarische Anstalt, 30(2): 1-40.

KRETZOI, M. (1938): Die Raubtiere von Gombaszög nebst einer Übersicht der Gesamtfauna. – Annales Musei Nationalis Hungarici, 31 (1937–1938) 88-157.

KRETZOI, M. (1954): Bericht über die calabrische (villafranchische) Fauna von Kisláng. – Mitteilungen Jahrbuch königlich ungarische An-stalt (1953): 213-265.

KRETZOI, M. & FEJFAR, O. (1982): Viverriden (Carnivora, Mammalia) im europäischen Altpleistozän. – Zeitschrift geologische Wis-senschaften, 10(7): 979-995.

LEHMANN, U. (1953): Eine Villafranchiano-Fauna von der Erpfinger Höhle (Schwäbische Alb). – Neues Jahrbuch für Geologie und Paläontologie, ML., 437-464, Stuttgart.

LINDSAY, E.H., OPDYKE, N.D. & FEJFAR, O. (1997): Correlation of selected late Cenozoic European mammalian faunas with the magnetic

polarity time scale. – Palaeogegraphy, Palaeoclimatology, Palaeoecology, 133: 205-226.

LEHMANN, U. (1957): Weitere Fossilfunde aus dem ältesten Pleistozän der Erpfi nger Höhle (Schwäbische Alb). – Mitteilungen geolo-gisches Staatsinstitut Hamburg, 26: 60-69.

MAZZA, P. & RUSTIONI, M. (1994): On the phylogeny of Eurasian bears. – Palaeontographica, A, 230, 1/3: 1-38.

MCKENNA, M.C. & BELL, S.K. (1997): Classification of Mammals. – 632 p.; New York.

MORLO, M. & KUNDRÁT, M. (2001): The first carnivoran fauna from the Ruscinium (Early Pliocene, MN 15) of Germany. – Paläontolo-gische Zeitschrift, 75(2): 163–187.

PEI, W.-Ch. (1934): On the Carnivora from locality 1 of Choukoutien. – Paleontologica Sinica, 8(1): 1–166.

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Manuskript submitted 2003 – 6 – 11Manuskript accepted 2003 – 8 – 06

Plate 1

Fossil records of Carnivores from Hajnáčka

Figs 1–11: Hyaena perrieri CROIZET & JOBERT, 1828:

Figs 1–3: fragment of the right mandible with c and p2-4 (SNM-MNH, Z 26379).

Figs 4–6: detail view of lower premolars p2-4 of the right mandible (SNM-MNH, Z 26379).

Figs 7–9: upper left juvenile canine (SNM-MNH, Z 26378). 10-11: left calcaneus (SNM-MNH, Z 26377).

Figs 12–13: Megantereon sp.: the fi rst right metacarpus (SNM-MNH, Z 26376). – 1, 4, 7 – lingual, 2, 5 – buccal, 3, 6 – occlusal, 8 – distal, 9 – labial, 10 – dorsal, 11 – medial, 12 – plantar, and 13 – dorsal views. – Scale A for fi gs 1–3, scale B for fi gs 4–9, scale C for fi gs 10–11, and scale D for fi gs 12–13.

Cour. Forsch.-Inst. Senckenberg, 246, 2004 Plate 1


44

Plate 2

Fossil records of viverrids from Ivanovce

Figs 1–8: Hesperoviverra carpathorum (KRETZOI & FEJFAR, 1982).

Figs 1–2: the Holotype; fragment of the right mandible with p4 and m1 (SNM-MNH, Z 26386).

Figs 3–5: detail view to preserved lower teeth of the right mandible (SNM-MNH, Z 26386).

Figs 6–8: the Paratype; P4 dex. (SNM-MNH, Z 26387).

Figs 9–11: Viverrinae gen. et spec. indet.; fragment of the left mandible (inv.) (SNM-MNH, Z 26389). – 1, 4, 6, 10 – buccal, 2, 5, 8, 11 – lingual, and 3, 7, 9 – occlusal views. – Scale A for fi gs 1–2, scale B for fi gs 3–8, and scale C for fi gs 9–11.



46

Plate 3

Fossil records of carnivores from Hajnáčka

Figs 1–11: Lutra cf. bravardi POMEL, 1843; 4–6: Ursidae gen. et spec. indet.

Figs 1–3: right mandible with c, p3–4, and m1–2 (SNM-MNH, Z 26374/1).

Figs 4–6: anterior premolar (left P1 or right p3) (GMM, B-4077).

Figs 7–8: left upper canine (SNM-MNH, Z 26374/10).

Figs 9–11: fragment of the left mandible branch with p4 and m1 (reversed) (GMM, B-3598).

1, 5, 7, 10 – buccal, 2, 6, 9 – occlusal, and 3, 4, 8, 11 – lingual views.Scale A for fi gs 1–3, 9–11, and scale B for fi gs 4–8.


48

Plate 4

Upper P4 and M1 of fossil and extant species of Lutra BRUNNICH, 1772.

Figs 1–6: Lutra cf. bravardi POMEL, 1843.

Figs 1–3: from Hajnáčka (SNM-MNH, Z 26374/3).

Figs 4–6: Cast of the Type (MHN, Paris) from Perrier-Les Étouaires, Auvergne, France.

Figs 7–9: Lutra lutra LINNÉ, 1758, occlusal views of the right P4 and M1 of three extant species.

Figs 1, 4 – buccal, 2, 5, 7–9 – occlusal, and 3, 6 – lingual views.



Plate 5

The bones of the forelimb of Lutra cf. bravardi POMEL, 1843 from Hajnáčka.

Figs 1–4: right humerus (SNM-MNH, Z 26374/18).

Figs 5, 6: right radius (SNM-MNH, Z 26374/21)

Figs 7–8: right ulna (SNM-MNH, Z 26374/19); 1, 5, 7 – anterior, 2 – caudal, 3 – lateral, and 4, 6, 8 – medial views.

Figs 9–12: dorsal (left) and plantar (right) views of the metapodials: 9 – right mc5. (SNM-MNH, Z 26374/14), 10: right mc1 (SNM-MNH, Z 26374/16), 11: right mc2 (SNM-MNH, Z 26374/11), 12: right mc3 (SNM-MNH, Z 26374/12) – 13: right scapholunatum (SNM-MNH, Z 26374/15) dorsal (left) and plantar (right) views. – Scale A for fi gs 1–4, scale B for fi gs 5–8, and scale C for fi gs 9–13.

49

50

Plate 6

Fossil record of Lutra BRUNNICH, 1772 from Ivanovce.

Figs 1–3: Lutra sp. I.; right mandible with c and p2 – p4 (SNM-MNH, Z 26392).

Figs 4–6: Lutra sp. II.; left mandible with p4 – m2 (reversed) (SNM-MNH, Z 26393). – 1, 4 – buccal, 2, 5 – occlusal, and 3, 6 – lingual views.

Plate 7

Figs 1–11: The lower m1 of the genus Parailurus SCHLOSSER, 1899 from the European late Pliocene (scale A).

Figs 1–3, 7, 9: Parailurus hungaricus KORMOS, 1934; left m1 (reversed), Hajnáčka (SNM-MNH, Z 26380.

Figs 4–6, 8, 10: Parailurus anglicus (DAWKINS, 1888); right m1, Wölfersheim (SMF 2000/233.

Fig. 11: Parailurus sp.; fragment of left m2, Ivanovce (SNM-MNH, Z 26390; occlusal view). – 1, 4 – buc-cal, 2, 5 – occlusal, 3 6 – lingual, 7, 8, – posterior and 9, 10 – anterior views.

Figs 12–23: The tarsal bones of Parailurus hungaricus KORMOS, 1934 from Hajnáčka (scale B). 12, 14, 16, 18: left calcaneus (SNM-MNH, Z 263849).

Figs 13, 15, 17, 19: right calcaneus (reversed) (SNM-MNH, Z 26383/40).

Figs 20–23: right astragalus (SNM-MNH, Z 26383/39). – 12, 13, 20– dorsal, 14, 15, 21 – plantar, 16, 17, 22– me-dial, and 18, 19, 23 – lateral views.




52

Plate 8

The bones of the hindlimb of Parailurus hungaricus KORMOS, 1934 from Hajnáčka

Figs 1–6, 9: right tibia (SNM-MNH, Z 26383/1); 1 – anterior, 2 – caudal, and 3 – medial views (scale A). Scale B: 4 – dorsal, 5 – anterior and 6 – caudal views of the proximal head; 9 – plantar view of the tibial cochlea.

Figs 7, 8, 10, 11: distal part of right femur (SNM-MNH, Z 26383/2); 7 – ventral, 8 – anterior, 10 – caudal and 11 – medial views.

Figs 12–13: distal end of right fi bula (SNM-MNH, Z 26383/5); 12 – lateral, and 13 – medial view.


Date post:	12-Apr-2021
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