Before the Pliocene, only ochotonids such as Eurolagus, Amphilagus, or Prolagus are known from Austria (Angelone and Veitschegger, 2015 and Prieto et al., 2012 and references therein; Prieto et al., 2014). The first appearance of modern leporids in Europe was in the Late Miocene (Flynn et al., 2014 and references therein), and the first known leporid from Austria is Hypolagus petenyii (“H. beremendensis beremendensis”) from Middle Pliocene fissure fillings (Deutsch-Altenburg 9, 20, 21; MN 16; ca. 3.0–2.5 Ma old) (Čermák, 2009, Döppes and Rabeder, 1997 and Fladerer and Reiner, 1996a). This species evolved into, or was replaced by, the Early Pleistocene Hypolagus brachygnathus, which survived until the beginning of the Middle Pleistocene (Čermák, 2009 and Fladerer and Reiner, 1996a). The first species of the genus Lepus may have appeared in the Middle Pleistocene in Austria. A Lepus sp. (“L. terraerubrae”) inhabited the Early to Middle Pleistocene steppes of eastern Austria. This species, however, has not yet been properly described (Döppes and Rabeder, 1997 and Fladerer, 1984).

Finds of Lepus timidus occur rather frequently in Late Pleistocene sediments in eastern Austria (Fig. 1) and the oldest dated evidence of this species are from the Salzofenhöhle (Styria), about 54,000–31,200 a BP, and from the Peggauerwandhöhle (Styria), about 42,400–22,600 a BP (Döppes and Rabeder, 1997, Fladerer, 1994 and Pittioni, 1980). So far, the oldest occurrence of L. timidus in Lower Austria around the Vienna Basin is from the Late Palaeolithic locality Alberndorf 1 with radiocarbon dates of around 28,500 and 26,900 a BP (Jöris et al., 2010). The most recent evidences for fossil mountain hares in Austria so far are from the Merkensteinhöhle (Lower Austria), with an age of 13,000–10,000 a BP, and from the Teufelslucke near Eggenburg (Lower Austria), with an age of 13,000–7,000 a BP. The relative age of both caves, however, is based on the micromammal assemblages (e.g., Dicrostonyx) (Fig. 1) (Döppes and Rabeder, 1997).

So far the most reliable oldest evidence of the European hare, Lepus europaeus, in Austria is from the Knochenhöhle near Kapellen (Styria), where one specimen of L. europaeus was found among several hundred of L. timidus. A ¹⁴C age of a mountain hare from the same layer dated back to about 14,000 a BP (Fladerer and Reiner, 1996b).

Oryctolagus cuniculus is endemic on the Iberian Peninsula, as well as southern France, and was introduced to Austria in the late Middle Ages around the 15^th century (Spitzenberger, 2001). Newer evidence from eastern Austria (canabae legionis, Vindobona) suggests an earlier introduction, within the Roman period, but the temporal origin of these specimens still needs to be verified (Czeika, 2005).

The faunal material was excavated between 2004 and 2011 by members of the Paläontologischer Forschungsverein Lindabrunn (J. and M. Neitz, J. and M. Zierhofer), under the direction of Gerhard Trnka, Institute of Prehistoric and Historical Archaeology of the University of Vienna, and Michael Brandl, Austrian Academy of Sciences, Institute of Oriental and European Archaeology.

The leporid teeth were measured using the method of von den Driesch (1976), using a digital calliper to the closest 0.1 mm. Measurements of the upper and lower incisors from different localities (F.A. Fladerer, personal database) were added for species distinction. The excavations at the Smrcka Lorenz-Abris yielded 577 remains of leporids (Fig. 2). The majority of the material could not be identified to species level and therefore 386 remains are attributed to Lepus sp. and 65 to Leporidae indet. The remaining 126 remains belong to one of the three preserved leporid species: Lepus europaeus, L. timidus and Oryctolagus cuniculus.

Five bones from the total faunal sample (four large mammals, one leporid) were selected for ¹⁴C dating by the VERA-laboratory at the Faculty of Physics, University of Vienna (Tab. 1). The faunal material is stored in the Department of Palaeontology at the University of Vienna (PIUW) under the collective inventory number 7000/LB. The archaeological material is stored in the Institute of Prehistoric and Historical Archaeology of the University of Vienna (IUHA).

Order LAGOMORPHA BRANDT, 1855

Family LEPORIDAE FISCHER, 1817

Genus Lepus LINNÉ, 1758

Lepus europaeus PALLAS, 1778

Material: thirty-five bones and teeth are preserved. Among the 16 teeth are lower incisors (four left, seven right), upper incisors (three left, one right), and a lower third premolar (one right). Furthermore, a premaxillary bone with incisor (one right), lower jaws (five left, five right), and ulnae (three left, five right) are preserved. Seven right lower incisors are represented, which shows that at least seven individuals are represented.

Lepus timidus LINNÉ, 1758

Material: thirty-one bones and teeth of the leporid assemblage could be assigned to the mountain hare: lower incisors (three left, nine right), upper incisors (three left, two right), lower third premolars (eight left, three right), one complete lower jaw and ulnae (two left).

Taxonomy:

The measurements of the incisor teeth (Fig. 3 and Fig. 4) show a rather clear distinction between Lepus timidus and L. europaeus. The mountain hare has generally more quadratic incisors than the European hare (e.g., Koby, 1959 and Morel and Müller, 1997). Additionally, the lower third premolar can be used to separate these two species. The lower third premolars of both species share the same basic morphology with, for example, two different sized anteroconids (Callou, 1997). In L. timidus, however, this tooth is more elongated and rectangular compared to the more roundish outline the one of L. europaeus (Koby, 1959). Hauser (1921) also presented diagnostic features to distinguish between postcranial elements of L. timidus and L. europaeus. These characters, however, have proven to be not feasible and therefore dental characters are used for species distinction (Koby, 1959). Nonetheless, in the present study, we found that the olecranon of the ulna was useful for species distinction. They represent two morphotypes that were only found in layers with the one or the other species. In the mountain hare, the olecranon of the ulna is more slender and smaller than in the European hare (Fig. 5).

Genus Oryctolagus LILLJEBORG, 1873

Oryctolagus cuniculus (LINNÉ, 1758)

Material: sixty specimens of bones and teeth could be identified: one skull, two fragments of the skull, pre-/molars (two left, one right), upper jaws (three left), lower jaws (two left, four right), radii (two right), ulnae (two left), coxae (four left, four right), femora (two right), tibiae (two left, three right), calcanei (two left, three right), metacarpals (five left), and metatarsals (six left, ten right). Inferred from the right mandibles, a minimum number of four individuals could be identified.

Taxonomy:

There are several diagnostic characters known to distinguish Oryctolagus from Lepus, apart from a generally smaller size. These have been extensively described by Callou (1997) and are only briefly presented here. Among other features, the skull of the European rabbit has, in general, a small sized choana, an os interparietale, a reduced protuberance occipital externa, a concave medio-caudal boarder of the parietal bone, a sphenoid crest, a slender and long maxillary palatine processus, a long processus temporalis. The foramen mentale in the lower jaw is rather big and the diastema is longer than in the hare. The upper incisors have a more roundish mesial lobe and, in the lower third premolar, the two anteroconids are about the same size and shape. There is a long processus hamatus on the acromion of the scapula and the tuberositas deltoidea of the humerus has an abrupt end. The radius is curved and the diaphysis of the ulna is continuous in thickness in cranial view. The foramen nutricium on the ilium is small. Also, the collum ossis femoralis is quite short in cranial view and the two tubercles which are forming the eminentia intercondylaris of the tibia are blunt.

The well-documented stratigraphy of the excavation at the Smrcka Lorenz-Abris as well as the well-preserved remains reflect the dynamics between concurring species at the end of the Pleistocene and within the Holocene. The European rabbit, the mountain hare, as well as the European hare are preserved in one locality and in a stratigraphic order. Lepus europaeus and L. timidus do not overlap in their stratigraphic range. This site has yielded the most recent fossil record so far of the mountain hare in the Vienna Basin, with an age of 14,180–13,830 cal a BP. The European hare became the dominant species in this area around 7,000 a BP. The record of Oryctolagus cuniculus, however, is not well resolved. Still, it is only found in the uppermost parts of the profile.

It seems that in warmer conditions, L. timidus is more likely to vanish from areas where L. europaeus is present due to interspecific competition, disease, hybridisation, and due to the impact of humans (Spitzenberger, 2001 and Thulin, 2003). In general, the Austrian fossil and historical record of its three extant leporid species is patchy. The succession found during the excavation at Smrcka Lorenz-Abris is an important contribution to fill in the gaps in the record. Only further excavations will increase our understanding and knowledge of the complex picture of extinction and habitat occupation at the end of the Pleistocene and within the Holocene.