Middle Pleistocene

Musil, Rudolf, Sabol, Martin, Ivanov, Martin & Doláková, Nela, 2014, Middle Pleistocene Stratigraphy Of The Deposits In Za Hájovnou Cave (Javoříčko Karst, Northern Moravia, Czech Republic), Acta Musei Nationalis Pragae Series B 70 (1 - 2), pp. 107-119 : 111-112

Middle Pleistocene palaeobiological record from studied sections

Findings of lions

The findings of the lions do not show an uniformity in metric and morphological characters and all the finds can be divided into two groups. 1) The lion remains attributed to Panthera fossilis (REICHENAU, 1906) on the basis of morphological and metric data, demonstrate relatively big individuals of lion-like felids close to these from other European Middle Pleistocene sites. They were found in layers dated to MIS 9 and/or older (MIS 11–13). Typical individuals of the species are known from the MIS 11–13 period.

2) The individuals of P. fossilis from layers dated to MIS 9 and/or younger exhibit characteristics similar to so called “intermediate forms” (towards Panthera spelaea ) from the Saalian Complex (MIS 6–8). These forms are preliminarily named by some authors (e.g. Argant 2010 or Marcziszak et al. 2014) as “ P. intermedia ” (nomen nudum).

According to one of co-authors ( Sabol 2014), P. spelaea has not to be a descendant of European palaeopopulations of P. fossilis . It is not excluded that P. fossilis evolved autochthonously in Europe and it gained some characteristics convergent with those of the Late Pleistocene P. spelaea at the end of the Middle Pleistocene (so called “intermediate forms” from MIS 6–8). P. spelaea probably evolved in Asia and penetrated Europe during the Late Pleistocene (or during the Middle/Late Pleistocene transmission resp.), when replaced autochthonous palaeopopulations of P. fossilis . From the biostratigraphic point of view, all records of lion-like felids from the European Middle Pleistocene are so far attributed to P. fossilis with stratigraphical range from MIS 17 to MIS 8 (6), representing probably two “subspecies” ( P. fossilis fossilis from MIS 17 to MIS 9 and P. fossilis “ intermedia ” from MIS (9) 8 to MIS 6) or two “chronospecies” ( P. fossilis from MIS 17 to MIS 9 and P. aff. fossilis (= P. “ intermedia ”) from MIS (9)8 to MIS 6) with interfragmentary boundary at MIS 9. Based on that, the Za Hájovnou Cave could play an important role in our knowledge of the P. fossilis evolution and the MIS 9 horizon could be a crucial. Unfortunately, limited quantity of suitable fossils as well as suitable characteristics do not enable to specify our conclusions and the abovementioned scenario is so far only hypothetical.

From the relationship viewpoint of cave lion ( P. spelaea ) to modern lion ( P. leo ), some authors (predominantly of the western provenience) regard cave lion as only an extinct subspecies of modern lion, falling back on results of ancient mitochondrial DNA analysis. On the other hand, some Russian scientists (such as Baryshnikov and Boeskorov 2001 or Sotnikova and Nikolskyi 2005), however, regard cave lion as a separate species on the basis of cranial and dental morphometrics. In addition, the results of phylogenetic analysis ( Burger et al. 2004, Barnett et al. 2009) rather shows that cave lion forms a distinct clade that is close related to extant lions from Africa and Asia.

Voles and lemmings

Fauna of small mammals, especially voles (arvicolids), is of particular importance for stratigraphic classification of the layers within studied sections. Remains of voles and lemmings were described only from two sections including ZH P-2 and ZH P-8b ( Ivanov and Vöröš 2014):

Velikonoční jeskyně (= Easter Corridor), ZH P-2, layers 6a, 6ab, 6b ( Microtus gregalis , M. arvalis /agrestis, M. oeconomus , Clethrionomys glareolus , Lemmus lemmus , Dicrostonyx cf. torquatus , Arvicola cantiana ). Layer 7a–f ( Microtus gregalis , M. arvalis /agrestis, M. oeconomus , Clethrionomys glareolus , Dicrostonyx cf. torquatus ).

Spojovací chodba, ZH P-8b, layer 1c. This layer is affected by gravitational redeposition of sediments of various ages. ( Microtus gregalis , M.arvalis /agrestis, M. aff. “ coronensis ”), layers 2a, 2b ( Microtus gregalis , M. arvalis /agrestis, M. oeconomus , Clethrionomys glareolus , Arvicola terrestris ).

As regards ZH P-2 section remains of voles and lemmings indicate that the assemblage is most probably of Saalian age. Layer 7(a–f) probably deposited during the MIS 8 and MIS 7, whereas layer 6 is also of the Saalian age but it originated later, probably within MIS 6 ( Ivanov and Vöröš 2014). Fossil remains of other groups are very scarce in ZH P-2 and there is no radiometric dating from this section. Therefore, the stratigraphic classification is not widely discussed.

The Middle Pleistocene vole assemblage from ZH P-8b, namely that of the layers 2a and 2b which are both situated in the Vykopaná chodba, as well as in the Narozeninová chodba, appears to be older than assemblages from ZH P-2 ( Ivanov and Vöröš 2014).

Palynospectra

Number of palynomorphs was very different in individual layers of studied sections. Some layers were rich in pollen and spores; other contained only sparse grains without substantial relevance. Other studied samples were sterile ( Doláková 2014). Short description of studied sections is as follows:

Velikonoční jeskyně, ZH P-2: Layers 6 (sterile) and 7 were studied. Overestimation of small Asteroideae , pollen of other herbs ( Chrysosplenium / R. trichophylus , Poaceae , Galium , Ranunculaceae ) was observed in the layer 7. No pollen of trees has been observed. This oryctocoenosis gives evidence for mechanical selection during transport with sediment flow. However, according to the study of cave sediments (Balcarka, Doláková 2004; Pod Hradem, Doláková, personal data) such palynospectra are typically found in sediments from the steppe environment of colder climatic stages.

Vykopaná chodba, ZH P-5: layer 1b – only several grains ( Pinus , Betula , Asteraceae ). Layers 2a and 2b were rich in palynological content (similar, but impoverished as ZH P-8b – see below). Layer 3 only single findings ( Picea , Fagus , Buxus ). Layers 5, 6 and 7 sterile

Narozeninová chodba, ZH P-8b: Layer 1 only several pollen grains ( Pinus , Asteraceae ). Layers 2a and 2b rich in palynomorphs. Frequent tree elements elements ( Pinus up to 40%, general - Carpinus , Tilia , Quercus , sporadic – Acer , Hedera ). Single pollen of Pterocarya , Ilex , Buxus , or Celtis / Juglans were found. Layer 4 lower part (debris cone) has different palynological content ( Corylus and herbs prevail, Carpinus and Tilia occur) if compared to layer 4 upper part from ZH P-10 and 11.

Narozeninová chodba, ZH P-9: Layer 5 in underlayer of a debris cone. Different pollen spectra, including variability in total content of pollen grains, were observed in individual depths of this macroscopically uniform layer. In the 9.5 m – trees prevailed over herbs (67:33%), representing vegetation of deciduous woodland with only a small admixture of conifers (5%). Corylus was most abundant (over 40%), Carpinus , Tilia , Quercus , Juglans and Alnus (reached about 15%). In the overlying samples there was an increased proportion of Pinus , Asteroideae and other herbs and a decrease in Corylus , Alnus and Tilia until they became absent at the 7.5– 6.5 m level. From the 3 m upwards the pollenspectra were approaching to the ones from debris cone (layer 4 lower part) and layers 2a, 2b (from ZH P-5 and ZH P-8b) without pre-Quaternary elements.

Narozeninová chodba, ZH P-10: Layers 2 and 3 sterile. Layer 4 upper part (debris cone) high amount of pollen and spores – Pinaceae highly prevailing, without Corylus , Carpinus and Tilia .

Narozeninová chodba, ZH P-11: Layer 1 only several single grains ( Pinus , Asteraceae , Poaceae ). Layers 3b and 4 (debris cone) were rich in palynomorphs. Palynological pictures of 3b were similar to 2b of ZH P-8b. Layer 4 was analogical to layer 4 of ZH P-10.