Short Communication
Molecular phylogeny of the extinct cave lion Panthera leo spelaea

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Abstract

To reconstruct the phylogenetic position of the extinct cave lion (Panthera leo spelaea), we sequenced 1 kb of the mitochondrial cytochrome b gene from two Pleistocene cave lion DNA samples (47 and 32 ky B.P.). Phylogenetic analysis shows that the ancient sequences form a clade that is most closely related to the extant lions from Africa and Asia; at the same time, cave lions appear to be highly distinct from their living relatives. Our data show that these cave lion sequences represent lineages that were isolated from lions in Africa and Asia since their dispersal over Europe about 600 ky B.P., as they are not found among our sample of extant populations. The cave lion lineages presented here went extinct without mitochondrial descendants on other continents. The high sequence divergence in the cytochrome b gene between cave and modern lions is notable.

Introduction

The cave lion [Panthera leo spelaea (Goldfuss, 1810)] was one of the most important carnivorous competitors of early man in Europe, from the early Middle Pleistocene onwards. It was an object of Palaeolithic art, such as the magnificent colour paintings in the Chauvet Cave (Ardèche, France) (Lorblanchet, 1995) or the impressive ivory sculptures from the Vogelherd cave (Swabian Alb, Germany) (Fig. 1). The first comprehensive morphological studies of cave lion remains, in the 19th and the beginning of the 20th century, showed a relationship to modern lions. Subsequent reinterpretations either linked cave lions to modern tigers or declared them a separate species. Osteological revisions have, however, always indicated a relationship to modern lions (Hemmer, 1974), although lately a new case has been made for a relationship to tigers based on brain endocasts (Groiss, 1996). Comparative morphological analysis of Pleistocene and Holocene lions on the level of geographic populations resulted in the description of two basic evolutionary lines: the spelaea group of the Holarctic Pleistocene and the leo group of Africa and southern Asia (Hemmer, 1974). Most authors favour the taxonomic combination of these groups within the species Panthera leo (Hemmer, 1974; Kurtén, 1968; Turner and Antón, 1997), but some prefer a taxonomic separation at the species level, into Panthera spelaea and Panthera leo (Baryshnikov and Boeskorov, 2001). Here, we report the mtDNA analysis of two Upper Pleistocene cave lions, one (Si) 47,180 + 1190/−1040 year B.P. and one (Ku) 31,890 ± 300 year B.P. old. Our results are consistent with the taxonomy of pantherine cats presented in Table 1.

Section snippets

Fossil bone samples

One specimen (Si), an almost complete skeleton of a cave lion embedded in a greyish silty clay, was excavated in 1985 (Rosendahl and Darga, 2004) at Siegsdorf in southeastern Bavaria, Germany. For preservation the bone surfaces were treated with a silica gel resin. Samples were taken for radiocarbon dating and DNA analysis from the interior compact bone of the right femur. The high collagen yield (19.4 wt% bone) suggests that the bones were not significantly altered diagenetically and that the

Diagenetic measurements (Table 3)

To test the general biomolecular state of preservation of the specimens before starting with the extensive ancient DNA analysis, bone samples were subjected to three separate diagenetic screening techniques: elemental analysis, Fourier-transform infrared spectroscopy, and SAXS. Very little mineral alteration is evident in either archaeological sample, as compared to the modern values for both SF and C/P. All crystallites measured using SAXS were determined to be plate-like (data not shown),

Acknowledgments

We thank Robert Darga, Naturkunde- und Mammutmuseum Siegsdorf, and Hugo Oberkofler, Museum Kufstein for providing cave lion samples. We are indebted to Sabine Hilsberg and Bert Geyer from the Zoo in Frankfurt/Main for providing blood samples, and Sabine Moeller-Rieker and Ursula Arndt for assistance. J.H. gratefully acknowledges funding from the Wellcome Trust Programme in Bioarchaeology. We thank Prof. Nicholas Conard for providing Fig. 1.

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