Search

The origin and evolutionary history of the European bison Bison bonasus (wisent) have become clearer after several morphological, genomic, and paleogenomic studies in the last few years, but these paleogenomic studies have raised new questions about the evolution of the species. Here, we present additional information about the population diversity of the species based on the analysis of new subfossil Holocene remains from the Balkan Peninsula. Seven ancient samples excavated from caves in Western Stara Planina in Bulgaria were investigated by mitochondrial D‐loop (HVR1) sequence analysis. The samples were dated to 3,800 years BP by radiocarbon analysis. Additionally, a phylogenetic analysis was performed to investigate the genetic relationship among the investigated samples and all mitochondrial DNA sequences from the genus Bison available in GenBank. The results clustered with the sequences from the extinct Holocene South‐Eastern (Balkan) wisent to the fossil Alpine population from France, Austria, and Switzerland, but not with those from the recent Central European (North Sea) one and the now extinct Caucasian population. In conclusion, these data indicate that the Balkan wisent that existed in historical time represented a relict and probably an isolated population of the Late Pleistocene‐Holocene South‐Western mountainous population of the wisent., Bison bonasus remains from Ponor part of Western Stara Planina Mountain (Bulgaria). The data indicate that the Balkan wisent that existed in historical time represented a relict and probably isolated population of a distinct South European mountainous population of the wisent.

In order to investigate the detailed zoogeographical history of brown bears (Ursus arctos) in Bulgaria and their relationships with populations in neighboring regions of Europe and Asia, the amplified product length polymorphism (APLP) method for the mitochondrial DNA (mtDNA) haplogrouping was applied to ancient bone remains. The results showed that haplogroups for 12 of 31 samples (38.7%) were successfully classified using the APLP method, although partial sequences of the mtDNA control region were determined with PCR product-direct sequencing for only 6 samples (19.4%). Even among 25 samples, of which nucleotide sequences could not be determined, the APLP method successfully classified haplogroups of 6 samples (6/25, 24.0%), indicating an advantage of the APLP method. In Bulgaria, although both the Balkan/Italian lineage of mtDNA (clade 1b) and the Eastern European lineage (clade 3a1) have been identified from modern bears as reported in other studies, all bear remains examined in the present study had APLP haplogroup W. The mtDNA phylogenetic analysis showed that the ancient Bulgarian brown bears had clade 1b. This indicates that clade 1b was originally distributed in Bulgaria, whereas those with clade 3a1 could have entered from Romania to Bulgaria. In addition, the APLP and phylogenetic analysis of recent skin samples from Turkey showed that they have mtDNA of the Middle Eastern/Turkish lineage (referred to clade 7a). Therefore, Bulgaria on the Balkan Peninsula might have been located between two distribution borders: one is between clades 1b and 3a1, and the other is between clades 1b and 7.

With advances in molecular phylogeny, the Ursidae affinity of Ailuropoda is no longer controversial. However, the early evolution of Ailuropoda and its close relatives (the tribe Ailuropodini) is still unclear. In this study, we describe a new fossil discovery from Bulgaria, which represents a new taxon of Ailuropodini, ?Agriarctos nikolovi. The materials of Ailurarctos are restudied and the evolution and dispersal of Ailuropodini are discussed. Early Ailuropodini split into two lineages, one in Europe as Agriarctos (three species, whose assignment to the same genus is not certain), and one in southeastern Asia as Ailurarctos and later Ailuropoda. Ailurarctos is a paraphyletic group, with both known species as successive direct ancestors to Ailuropoda. Subtribe Ailuropodina is proposed here to include Ailurarctos and Ailuropoda. Turolian European Agriarctos paralleled with Ailuropodina in many aspects, which reflects similar adaptation towards a specific herbivorous diet.

International audience

The article presents new data on the status of the jungle cat (Felis chaus) in Azerbaijan, a species from the Red Data Book of Azerbaijan.

Many ideas have been proposed to explain the origin of bipedalism in hominins and suspension in great apes (hominids); however, fossil evidence has been lacking. It has been suggested that bipedalism in hominins evolved from an ancestor that was a palmigrade quadruped (which would have moved similarly to living monkeys), or from a more suspensory quadruped (most similar to extant chimpanzees)1. Here we describe the fossil ape Danuvius guggenmosi (from the Allgau region of Bavaria) for which complete limb bones are preserved, which provides evidence of a newly identified form of positional behaviour-extended limb clambering. The 11.62-million-year-old Danuvius is a great ape that is dentally most similar to Dryopithecus and other European late Miocene apes. With a broad thorax, long lumbar spine and extended hips and knees, as in bipeds, and elongated and fully extended forelimbs, as in all apes (hominoids), Danuvius combines the adaptations of bipeds and suspensory apes, and provides a model for the common ancestor of great apes and humans.

Extant colobine monkeys are specialized leaf eaters. But during the late Miocene, western Eurasia was home to colobines that were less efficient at chewing leaves than they were at breaking seed shells. To understand the link between folivory and granivory in this lineage, the dietary niche of Mesopithecus delsoni and Mesopithecus pentelicus was investigated in southeastern Europe, where a major environmental change occurred during the late Miocene. We combined dental topographic estimates of chewing efficiency with dental microwear texture analysis of enamel wear facets. Mesopithecus delsoni was more efficient at chewing leaves than M. pentelicus, the dental topography of which matches an opportunistic seed eater. Concurrently, microwear complexity increases in M. pentelicus, especially in the northernmost localities corresponding to present-day Bulgaria. This is interpreted as a dietary shift toward hard foods such as seeds or tubers, which is consistent with the savanna and open mixed forest biomes that covered Bulgaria during the Tortonian. The fact that M. delsoni was better adapted to folivory and consumed a lower amount of hard foods than M. pentelicus suggests that colobines either adapted to folivory before their dispersal to Europe or evolved adaptations to leaf consumption in multiple occurrences.

International audience; The late Miocene locality Hadjidimovo in Southwestern Bulgaria has yielded a huge collection of mammalian fossils, including a complete skull of Machairodus Kaup, 1833, first described (in Bul- garian) by Kovachev (2002). We re-describe it here, compare it with other Machairodus, and review the evolution of the genus. We conclude that the transition from M. aphanistus (Kaup, 1832) to M. giganteus (Wagner, 1848) is gradual and mosaic, that assigning these species to different genera fails to reflect this relationship, and that the Hadjidimovo skull represents a rather late evolutionary stage of this lineage.; Un crâne de Machairodus Kaup, 1833 (Felidae, Mammalia) du Miocène supérieur de Hadjidimovo (Bulgarie), et sa place dans l’évolution du genre.La localité de Hadjidimovo dans le Sud-Ouest de la Bulgarie a livré une énorme collection de mam- mifères fossiles, dont un crâne complet de Machairodus Kaup, 1833, décrit (en Bulgare) par Kovachev (2002). Il est ici re-décrit, et comparé avec les autres Machairodus, genre dont l’évolution est revue. Nous concluons que la transition entre M. aphanistus (Kaup, 1832) et M. giganteus (Wagner, 1848) est graduelle et mosaïque, que de rapporter ces espèces à des genres distincts ne reflète pas cette rela- tion, et que le crâne de Hadjidimovo illustre un stade avancé de cette lignée.

The fallow deer (Dama dama Linnaeus, 1758) has a long history of interaction with prehistoric humans. Beginning in the Neolithic, humans introduced fallow deer to several areas of the eastern Mediterranean and mainland Europe, with later additional importing happening in the Bronze and Iron Ages. However, in some parts of southeastern Europe, autochthonous populations of extant fallow deer may have survived through the end of the Pleistocene and into the early Holocene, making them available for exploitation by Neolithic and Chalcolithic communities. Climatic and vegetational regimes favorable to fallow deer covered nearly all of Bulgaria during this period, yet the heavy use of the species by human communities was restricted to a very small area around southeastern Bulgaria. Eventually, climate deterioration, habitat change, and overhunting led to the decline of fallow deer in later prehistory in Bulgaria. This paper offers a discussion of the environmental and cultural background of human‐fallow deer interactions from the 6th‐4th millennium BCE in Bulgaria.

The marbled polecat, Vormela peregusna, is a small marten-like mustelid distributed from southeastern Europe to western China. Since the individual numbers of this species have declined, it has bee...

International audience; This study represents the first extensive systematic investigation of the Miocene mammalian faunas of the Republic of Macedonia (FYROM), stored in the Macedonian Museum of Natural History, Skopje. They range in age from perhaps the early Miocene to the early Ruscinian, but the bulk of the fossils represent middle Turolian mammals. At least 57 taxa have been identified, from 25 different paleontological sites, mostly from the Vardar and Strumitsa river basins, but also from the Morievo and Delchevo regions. The richest localities are the middle Turolian localities of Karaslari (with 22 species) and Kiro Kuchuk (17 species). The rich fossil material greatly improves our knowledge of the Turolian Hipparion faunas of the Balkan-Iranian zoogeographic paleo-province, whose westernmost part was mostly documented in Greece and Bulgaria. The fauna displays the typical faunal features of the Balkan Pikermian biome, with dominance of hipparions (especially H. brachypus, but our revision does not confirm the presence of Hipparion verae in the Turolian faunas) and bovids such as Gazella, Tragoportax, and spiral-horned antelopes. Other forms usually found in the area, such as Microstonyx erymanthius, Dihoplus pikermiensis, chalicotheres, Choerolophodon pentelici, Mesopithecus pentelicus, or Adcrocuta eximia are also common. Several new forms have been identified among the carnivores, the giraffids and the bovids. The Macedonian material contributes to reconstructing the history of several taxa such as Simocyon, Metailurus, several hipparion species, Propotamochoerus, Bohlinia, Sivatherium. The most noticeable features of these Turolian faunas are: the abundance of spiral-horned antelopes, and rarity of antelopes of the Protoryx-Pachytragus group, as in Bulgaria, the co-existence of chalicotheriins and schizotheriins, the frequency of Dihoplus compared to Ceratotherium, the presence of Chilotherium, which reaches its westernmost longitude, and the presence of Anancus sp. in some localities, considered here as post-Pikermian.

The discovery of rich, well preserved skull material in the latest Middle Pleistocene deposits of Mishin Kamik cave (N-W Bulgaria) presents an opportunity for a new analysis of the taxonomy and the phylogeny of the so called “small cave bears”. Not all the small cave bears known would have necessarily had a common origin, the size decrease could be related in a number of cases to a parallelism. The bear from Mishin Kamik is identical with “Ursus rossicus” Borissiak from Krasnodar (S. Russia). Both these samples must be referred to U. savini Andrews from Bacton near Cromer (England). The specimens from Krasnodar and Mishin Kamik could be classified as U. sa. rossicus, a late form of the species. This species may have affinities with some Middle Pleistocene Siberian forms. U. savini is a small but very robust spelaeoid bear which is more advanced in a number of features than U. deningeri. In several aspects it attained the evolutional adaptations of the cave bears of the U. spelaeus-U. ingressus group and represents an independent lineage of spelaeoid bears. The Mishin Kamik population was adapted to a mosaic landscape of forests and open areas in hilly terrain. This bear had well developed grazing adaptations but had weak motor abilities.

New Machairodontinae material from the late Miocene localities of Hezheng (China) and Hadjidimovo (Bulgaria) represents a new species of Paramachaerodus Pilgrim. Both localities are similar in age and suggest that the new species had a very large geographic range extending from northwestern China adjacent to the Tibetan Plateau (Gansu Province) to southeastern Europe or probably to all of southern Europe. The new species—Paramachaerodus transasiaticus sp. nov is characterized by a combination of features of “Promegantereon” and Paramachaerodus. This specific morphology, as well as the age of the Hezheng and Hadjidimovo (early Turolian, after the European Land Mammal Ages) put the new species in intermediary position between “Promegantereon” and Paramachaerodus. The new felid material give grounds to discuss and revise in a new light the systematic and evolution of the “Promegantereon”–Paramachaerodus lineage, which should represent successive stages of one and the same genus: Paramachaerodus Pilgrim.

In this study, we investigated molecular phylogenetic status of the marbled polecat (Vormela peregusna) from Bulgaria, using sequences of two Y-chromosomal genes (SRY and ZFY). The phylogenetic tree inferred using combined sequences of both genes indicated that the marbled polecat was split from genera Lutra, Neovison and Mustela after genus Martes was diverged in family Mustelidae. In addition, we analyzed molecular phylogeography of the Bulgarian population of the marbled polecat, using cytochrome b and control region sequences of mitochondrial DNA (mtDNA). The phylogenetic tree of cytochrome b indicated that the haplotypes of the Bulgarian population comprised two haplogroups, which were the most ancestral clades. Additionally, the control region phylogeny showed that the haplotypes of Bulgaria formed two haplogroups: one was the most ancestral clade, and the other was the derivative clade. One individual with the most ancestral cytochrome b clade had a control region haplotype of the derivative clade. Thus, this study revealed that the most ancestral lineages of the marbled polecat are included in the population of Bulgaria. The Bulgarian population could be a remnant lineage from a basal for the species, which in Pleistocene occupied a relatively large area related to the Balkan-Caucasian.

Many ideas have been proposed to explain the origin of bipedalism in hominins and suspension in great apes (hominids); however, fossil evidence has been lacking. It has been suggested that bipedalism in hominins evolved from an ancestor that was a palmigrade quadruped (which would have moved similarly to living monkeys), or from a more suspensory quadruped (most similar to extant chimpanzees)

ObjectivesThe history of dog domestication is still under debate, but doubtlessly, it is a process of an ancient partnership between dogs (Canis familiaris) and humans. Although data on ancient DNA dog diversity are scarce, it is clear that several regional dog populations had been formed in Eurasia up to the Holocene. During the Neolithic Revolution and the transition from hunter-gatherer to farmer societies, followed by civilization changes in the Antiquity period, the dog population structure also changed. This process was due to replacement with newly formed dog breeds.MethodsIn this study we have presented for the first time mitochondrial data about South-Eastern Europe (the Balkans) ancient dog remains from the Early Neolithic (8 000 years BP) to the Late Antiquity ages (up to 3th century AD). A total of 25 samples were analyzed using the mitochondrial D-loop region (HVR1).ResultsThe results have shown the presence of A (70%) and B (25%) clades throughout the whole investigated period. In order to clarify the position of our results within the ancient dog population in Eneolithic Eurasia, we performed phylogenetic analysis with the available genetic data sets. This data revealed a similarity of the Bulgarian dogs’ structure to that of ancient Italian dogs (A, B, and C clades), which suggests a new prehistoric and historic Mediterranean dog population. A clear border can be seen between South-European genetic dog structure, on the one hand, and, on the other hand, Central-West (clade C), East (clade D) and North Europe (clades A and C). This corresponds to genetic data for European humans during the same period without admixture between dog populations.ConclusionsOur data have shown for the first time the presence of clade B in ancient Eurasia. This is not unexpected as the B haplogroup is widely distributed in extant Balkan dogs and wolves. The presence of this clade both in dogs and in wolves on the Balkans may be explained with hybridization events before the Neolithic period. The spreading of this clade across Europe together with the A clade is related to the possible dissemination of newly formed dog breeds from Ancient Greece, Thrace and the Roman Empire.

Two trenches excavated at Magura Cave, north-west Bulgaria, have provided Late Pleistocene lithic artefacts as well as environmental evidence in the form of large and small mammals, herpetofauna and pollen recovered from Crocuta coprolites. One of the trenches also has a visible tephra layer which has been confirmed as representing the major Campanian Ignimbrite eruption and is accurately dated at the source area to 39,280 ± 55 yrs and radiocarbon determinations have added to chronological resolution at the site. The palaeoenvironment of the region during the Late Pleistocene is discussed in the context of hominin presence and shows a mosaic landscape in a region considered a crucial refugium for both plants and mammals, including hominins.

Neolithic/Chalcolithic livestock domestication is an important issue for understanding the mode of life and economics of ancient human communities. The Balkans appears to be a crucial point for clarifying the socio-economical interrelations between the Oldest Middle Eastern/Anatolian and newly formed cultures in Europe. Two main hypotheses regarding the early history of cattle domestication, from their ancestor - the aurochs, have been discussed: multipoint domestication centers or single point origin and subsequent worldwide dissemination. In this study, we provide molecular data about the Balkan aurochs for the first time as well as additional information for the Neolithic/Chalcolithic cattle populations in this geographic location. A total of seventeen samples from different ancient settlements were analyzed according to D-loop control region. The results did not show different genetic profile of wild and domestic populations. All haplotypes were found to belong to the basic macro-haplogroup T. The majority of specimens (n = 14) were defined to form a new Balkan-specific T6 haplogroup. Only two of the ancient samples analyzed were assigned to the T3 haplotype predominating in Europe. We attempt to throw new light on the earliest cattle domestication events in Europe, thus, the results presented are discussed in two directions: (a) The possibility of local independent domestication processes in Neolithic South-Eastern Europe; (b) The single point domestication in the Middle East and subsequent cattle dissemination in Europe. Our data does not exclude the possibility for independent domestication events followed by a second wave of parallel dissemination of cattle herds via the Mediterranean route.

International audience; We describe the oldest known European ruminant, Bachitherium thraciensis sp. nov., from late Eocene (latest Bartonian or early Priabonian) strata in Bulgaria. The new specimen, which possesses the most primitive dental morphology known in the Bachitheriidae family, predates its western European relatives by at least 4.5 Myr. The discovery suggests that ruminants dispersed into Europe in a series of east to west waves, preceding and following the Eocene–Oligocene boundary. Probably originating from Asia, the oldest ruminant (reported here) migrated into southeastern Europe. During a second wave, comprising the Grand Coupure dispersal event, ruminants moved into western Europe at the Eocene–Oligocene boundary. A third wave composed by Bachitheriidae and Tragulidae arrived in western Europe, 2 Myr later, in the earliest Oligocene. The origin of this east to west diachronism was probably facilitated by orogenesis that, episodically, allowed dispersal of ruminants from Asia, through southeastern Europe, and into western Europe, in what we herein term the “Bachitherium dispersal event”. However, none of the abundant perissodactyl fauna from southeastern Europe migrated to western Europe during this time, because, like their relatives in central Asia, they may not survive to terminal Eocene cooling and aridity.

A hominid tooth found in a river terrace layer from Azmaka (Bulgaria) has been biochronologically dated to ~7 Ma. This age is younger than expected and indicates that hominids became extinct in Europe later than previously thought. In this study, authigenic 10 Be/ 9 Be is applied to potentially determine the absolute age of the Miocene river terrace layers containing the hominid tooth. Modern fluvial channel bed deposits provide the initial starting value for the geochronological clock. Homogeneity of modern fluvial channel bed deposits and closed-system behavior of the authigenic minerals in the river terrace layers are required. 10 Be/ 9 Be ratios analyzed for modern fluvial channel beds vary by a factor of two. The initial starting value for the geochronological clock is not well-constrained and age calculations uncertain. Furthermore, the 10 Be/ 9 Be ratios analyzed for different cycles of the terrace layers are variable within different layers and do not decrease with formation depth. The mean authigenic age based on the mean 10 Be/ 9 Be ratio from six terrace layers and using the mean initial ratio from measured modern fluvial channel bed samples is 5.80 ± 0.70 Ma. An authigenic 10 Be/ 9 Be age of 6.20 ± 0.41 Ma is calculated for the river terrace layer associated with the hominid tooth. Despite these first promising ages, the method of authigenic minerals to date river terrace sediments needs further and more detailed investigations.

The split of our own clade from the Panini is undocumented in the fossil record. To fill this gap we investigated the dentognathic morphology of Graecopithecus freybergi from Pyrgos Vassilissis (Greece) and cf. Graecopithecus sp. from Azmaka (Bulgaria), using new μCT and 3D reconstructions of the two known specimens. Pyrgos Vassilissis and Azmaka are currently dated to the early Messinian at 7.175 Ma and 7.24 Ma. Mainly based on its external preservation and the previously vague dating, Graecopithecus is often referred to as nomen dubium. The examination of its previously unknown dental root and pulp canal morphology confirms the taxonomic distinction from the significantly older northern Greek hominine Ouranopithecus. Furthermore, it shows features that point to a possible phylogenetic affinity with hominins. G. freybergi uniquely shares p4 partial root fusion and a possible canine root reduction with this tribe and therefore, provides intriguing evidence of what could be the oldest known hominin.

The cat has long been important to human societies as a pest-control agent, object of symbolic value and companion animal, but little is known about its domestication process and early anthropogenic dispersal. Here we show, using ancient DNA analysis of geographically and temporally widespread archaeological cat remains, that both the Near Eastern and Egyptian populations of Felis silvestris lybica contributed to the gene pool of the domestic cat at different historical times. While the cat's worldwide conquest began during the Neolithic period in the Near East, its dispersal gained momentum during the Classical period, when the Egyptian cat successfully spread throughout the Old World. The expansion patterns and ranges suggest dispersal along human maritime and terrestrial routes of trade and connectivity. A coat-colour variant was found at high frequency only after the Middle Ages, suggesting that directed breeding of cats occurred later than with most other domesticated animals.

A well-preserved felid skull from the Turolian (late Miocene) of Karaslari (FY Republic of Macedonia) is described and assigned to a new taxon, Yoshi garevskii, gen. et sp. nov. The new genus Yoshi is characterized by a round, short, and deep skull, and short canines that are somewhat transversely compressed but are not serrated. We assign to the same genus the poorly preserved skulls from the Aegean region described as Metailurus parvulus (Hensel, 1862), the Chinese type of “Metailurus” minor Zdansky, 1924, as well as some other skulls from China. The latter name thus becomes Y. minor (Zdansky, 1924), but the former name is a nomen dubium. The genus Metailurus Zdansky, 1924, is represented by the type-species M. major Zdansky, 1924; it considerably differs from the forms here referred to the new genus, and is closer to the Machairodontinae. Although definitely more plesiomorphic in dental features, Yoshi, especially Y. garevskii, reaches a high specialization in its skull shape, parallel to that of the cheetah, suggesting that it represents the first attempt towards the morpho-functional model of this modern felid.

International audience; The upper Miocene sites of Hadjidimovo and Kalimantsi in South-Western Bulgaria have yielded a few fossils of the ailurid Simocyon Wagner, 1858, a skull and associated mandible of Promeles Zittel, 1890 (a relatively well-known member of the Mustelidae), and several skulls of Promephitis Gaudry, 1861, an Eurasian member of the Mephitidae, a family represented today by three American and one South-Asian genera, to which were also ascribed Palaeomephitis Jäger, 1839, Miomephitis Dehm, 1950, and Proputorius Filhol, 1890, from the Middle Miocene of Europe, as well as several American Clarendonian to Hemphillian genera. We show that the mephitid status of Palaeomephitis, Miomephitis, and Proputorius is doubtful, and that the late Miocene Promephitis is at a lower grade of encephalization than modern Mephitidae and could be the sister taxon of Pleistocene and modern forms, which can be called Mephitinae. Within this subfamily, the American Pleistocene and modern taxa form the sister group of the Asian Mydaus Cuvier, 1821, which displays numerous autapomorphies. A major, previously unnoticed characteristic of the Mephitinae and Promephitis is that their P4 lingual cusp is a hypocone, unlike that of the Clarendonian American forms, in which it is a protocone. If the latter taxa really belong to the Mephitidae, they must have split early in its history.

The present study aimed to compare mitochondrial diversity among Balkan Neolithic/Chalcolithic cattle and present day Shorthorn Rhodopean cattle (Busha) to throw a new insight into European cattle domestication. The results showed that both ancient- and present-day samples belonged to the macrohaplogroup T. From the 28 sequences (8 ancient and 20 modern), the T1 and T2 haplogroup represent about 3.6% (1/28; 1/28). The T3 haplogroup was with the highest frequency – 57% (16/28). Based on the SNPs on 16057A and 16133C, the new T6 haplogroup was proposed. This haplogroup represents 75% from the ancient and 20% from the present day Bulgarian brachicerous cattle population. The survey in GenBank data base did not find a similar motif, except for the recent Serbian Busha cattle. Overall, these results showed that: (i) The newly named T6 haplogroup is Balkan specific; (ii) The T6 haplogroup survives in present day Bulgarian rhodopean cattle; (iii) The Balkan brachicerous cattle is the oldest European catt...