The 10,000-year biocultural history of fallow deer and its implications for conservation policy.

Over the last 10,000 y, humans have manipulated fallow deer populations with varying outcomes. Persian fallow deer (Dama mesopotamica) are now endangered. European fallow deer (Dama dama) are globally widespread and are simultaneously considered wild, domestic, endangered, invasive and are even the national animal of Barbuda and Antigua. Despite their close association with people, there is no consensus regarding their natural ranges or the timing and circumstances of their human-mediated translocations and extirpations. Our mitochondrial analyses of modern and archaeological specimens revealed two distinct clades of European fallow deer present in Anatolia and the Balkans. Zooarchaeological evidence suggests these regions were their sole glacial refugia. By combining biomolecular analyses with archaeological and textual evidence, we chart the declining distribution of Persian fallow deer and demonstrate that humans repeatedly translocated European fallow deer, sourced from the most geographically distant populations. Deer taken to Neolithic Chios and Rhodes derived not from nearby Anatolia, but from the Balkans. Though fallow deer were translocated throughout the Mediterranean as part of their association with the Greco-Roman goddesses Artemis and Diana, deer taken to Roman Mallorca were not locally available Dama dama, but Dama mesopotamica. Romans also initially introduced fallow deer to Northern Europe but the species became extinct and was reintroduced in the medieval period, this time from Anatolia. European colonial powers then transported deer populations across the globe. The biocultural histories of fallow deer challenge preconceptions about the divisions between wild and domestic species and provide information that should underpin modern management strategies.

A new Miocene ape and locomotion in the ancestor of great apes and humans.

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 Allgäu 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.

Terrestriality as reflected in the humerus of Mesopithecus delsoni (Cercopithecidae, Colobinae) from Hadjidimovo, Bulgaria.

The fossil colobine genus Mesopithecus is the oldest European monkey, ranging from the Late Miocene to the earliest Pleistocene. It is one of the most successful genera of Old World monkeys since the late Neogene. Its ecology, as an indicator of Late Miocene environments, is of particular interest. Several investigations have clarified the locomotor adaptations of the middle and late Turolian Balkan Mesopithecus pentelicus, but such studies are virtually absent for the earliest known taxon, the early Turolian Mesopithecus delsoni, due to the lack of fossil material. However, a large collection of postcranial material of M. delsoni from the Bulgarian Early Turolian locality of Hadjidimovo provides the first opportunity for such an analysis. The present study examines the functional morphology of the fossil humeri of M. delsoni from Hadjidimovo, Bulgaria, and of M. pentelicus from Bulgarian and Greek fossil localities. We provide detailed comparative qualitative descriptions and use univariate and multivariate quantitative analyses of one angular and 12 linear measurements in comparison with 149 extant Cercopithecidae, representing 14 genera and 34 species. Our analyses demonstrate that the humeral elements from Hadjidimovo show important morphological differences from those of M. pentelicus from Pikermi, Kalimantsi, and Gorna Sushitsa, suggesting strong terrestrial tendencies for M. delsoni. This finding, when considered together with the paleobiologial inference of semiterrestriality for the early cercopithecoid Victoriapithecidae, might indicate that the first colobines (still unknown) were also semiterrestrial. Finally, the morphological features related to terrestriality in M. delsoni, which differ from those of the later M. pentelicus, provide additional data in support of the idea that the older taxon represents a separate species.

Status and Numbers of the Brown Bear (Ursus arctos L.) in Bulgaria.

Until recently, the Bulgarian bear population (Ursus arctos L.) was considered one of the significant ones in Europe and one of the few with more than 500 bears. While the numbers of some neighboring populations may be increasing, the Bulgarian population has been on a downward trend since the early 1990s. The probable numbers of the species at the end of the 1980s was about 700-750 individuals. Calculations based on field data from national monitoring and statistical analysis show probable numbers in Bulgaria in 2020 of about 500 individuals (data for the autumn state). This decline is mostly related to poaching due to weaker control activity, the reduction of forest areas and habitat fragmentation. The preservation of the Bulgarian population, which, together with the other Balkan populations and the Apennine bear, has a unique gene pool, is particularly important from the point of view of preserving the biodiversity of the species in Europe.

Diversity of mitochondrial D-loop haplotypes from ancient Thracian horses in Bulgaria.

The domestication of the horse began possibly more than 5000 years ago in the western part of the Eurasian steppe, and according to the leading hypothesis, horses first spread from the Steppe toward the region of the Thracian culture, starting in the second half of the 2nd millennium BCE and flourished from the fifth to first centuries BCE, mainly located in present-day Bulgaria. We analyzed 17 horse bone remains excavated from Thracian archaeological sites (fourth to first centuries BCE) in Bulgaria and successfully identified 17 sequences representing 14 different haplotypes of the mitochondrial D-loop. Compared with the mtDNA haplotypes of modern horses around the world, ancient Thracian horses in Bulgaria are thought to be more closely related to modern horses of Southern Europe and less related to those of Central Asia. In addition, the haplotypes we obtained represented 11 previously reported modern horse mtDNA haplogroups: A, B, D, E, G, H, I, L, N, P, and Q. All the haplogroups contain modern and regionally predominant haplotypes occurring in Europe, the Middle East, and Central Asia. Our results indicate that Thracian horses in Bulgaria have had relatively high genetic diversity and are closely related to modern horse breeds.

On Mesopithecus habitat: Insights from late Miocene fossil vertebrate localities of Bulgaria.

The aim of this study is to describe the environments where the cercopithecid Mesopithecus was found during latest Miocene in Europe. For this purpose, we investigate the paleoecology of the herbivorous ungulate mesofauna of three very rich late Miocene fossil localities from southwestern Bulgaria: Hadjidimovo, Kalimantsi and Strumyani. While Mesopithecus has been found in the two first localities, no primate remains have yet been identified in Strumyani. Comparison between localities with and without primates using the herbivore mesofauna allows the cross-corroboration of paleoenvironmental conditions where this primate did and did not live. A multi-parameter statistical approach involving 117 equid and 345 bovid fossil dental and postcranial (phalanges, metapodia, astragali) remains from these three localities provides species to generic-level diet and locomotor habit information in order to characterize the environment in which Mesopithecus evolved. The analysis of dental mesowear indicates that the bovids were mainly mixed feeders, while coeval equids were more engaged in grazing. Meanwhile, postcranial remains show that the ungulate species from Hadjidimovo and Kalimantsi evolved in dry environments with a continuum of habitats ranging from slightly wooded areas to relatively open landscapes, whereas the Mesopithecus-free Strumyani locality was in comparison reflecting a rather contrasted mosaic of environments with predominant open and some more closed and wet areas. Environments in which Mesopithecus is known during the late Miocene were not contrasted landscapes combining open grassy areas and dense forested patches, but instead rather restricted to slightly wooded and homogeneous landscapes including a developed grassy herbaceous layer.

New data on the evolutionary history of the European bison (Bison bonasus) based on subfossil remains from Southeastern Europe.

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.

From leaves to seeds? The dietary shift in late Miocene colobine monkeys of southeastern Europe.

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.

Folivory or fruit/seed predation for Mesopithecus, an earliest colobine from the late Miocene of Eurasia?

Here we compare dental microwear textures from specimens of the fossil genus Mesopithecus (Cercopithecidae, Colobinae) from the late Miocene of Eastern Europe with dental microwear textures from four extant primate species with known dietary differences. Results indicate that the dental microwear textures of Mesopithecus differ from those of extant leaf eaters Alouatta palliata and Trachypithecus cristatus and instead resemble more closely those of the occasional hard-object feeders Cebus apella and Lophocebus albigena. Microwear texture data presented here in combination with results from previous analyses suggest that Mesopithecus was a widespread, opportunistic feeder that often consumed hard seeds. These data are consistent with the hypothesis that early colobines may have preferred hard seeds to leaves.

Dispersal history of the golden jackal (Canis aureus moreoticus Geoffroy, 1835) in Europe and possible causes of its recent population explosion.

BACKGROUND: Data on the historical distribution of the golden jackal in Europe and its primary habitats are scarce. There are many new data on the population explosion and the rapid spread of the in Europe. However, the main factors for this expansion, the core population and its routes of dispersal, remain controversial or insufficiently studied. NEW INFORMATION: This study provides a profound analysis of the history of the jackal's (Canis aureus moreoticus Geoffroy, 1835) occurrence in Europe, the factors limiting or those triggering its expansion on the continent. The analysis shows that the timing of the species appearance in Europe still remains unclear. Historical data show that the species is a typical inhabitant of South-Eastern Europe, with some pulsations within its core area, as well as extensions to the north and west of it in favourable periods. Nowadays, the increase of the species range in Europe is the largest documented population explosion on the continent. We argue that this expansion originates from only three core populations, the Peri-Strandja area and the Dalmatian coast in the Balkans and the east parts of Western Transcaucasia in the Caucasus. This population explosion is largely due to a unique combination of factors of an anthropogenic nature.

Evidence for Early European Neolithic Dog Dispersal: New Data on Southeastern European Subfossil Dogs from the Prehistoric and Antiquity Ages.

The history of dog domestication is still under debate, but it is doubtless the process of an ancient partnership between dogs (Canis familiaris) and humans. Although data on ancient DNA for dog diversity are still incomplete, it is clear that several regional dog populations had 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 populations. In this study, we present for the first time mitochondrial data of ancient dog remains from the Early Neolithic (8000 years before present (BP)) to Late Antiquity (up to 3th century AD) from southeastern Europe (the Balkans). A total of 16 samples were analyzed, using the mitochondrial D-loop region (HVR1). The results show the presence of A (70%) and B (25%) clades throughout the Early and Late Neolithic 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 showed a similarity of the ancient Bulgarian dogs to Italian (A, B, and C clades) and Iberian (clades A and C) dogs' populations. A clear border can be seen between southern European genetic dog structure, on the one hand, and on the other hand, central-western (clade C), eastern (clade D) and northern Europe (clades A and C). This corresponds to genetic data for European humans during the same period, without admixture between dog populations. Also, our data have shown 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.

Genetic turnovers and northern survival during the last glacial maximum in European brown bears.

The current phylogeographic pattern of European brown bears (Ursus arctos) has commonly been explained by postglacial recolonization out of geographically distinct refugia in southern Europe, a pattern well in accordance with the expansion/contraction model. Studies of ancient DNA from brown bear remains have questioned this pattern, but have failed to explain the glacial distribution of mitochondrial brown bear clades and their subsequent expansion across the European continent. We here present 136 new mitochondrial sequences generated from 346 remains from Europe, ranging in age between the Late Pleistocene and historical times. The genetic data show a high Late Pleistocene diversity across the continent and challenge the strict confinement of bears to traditional southern refugia during the last glacial maximum (LGM). The mitochondrial data further suggest a genetic turnover just before this time, as well as a steep demographic decline starting in the mid-Holocene. Levels of stable nitrogen isotopes from the remains confirm a previously proposed shift toward increasing herbivory around the LGM in Europe. Overall, these results suggest that in addition to climate, anthropogenic impact and inter-specific competition may have had more important effects on the brown bear's ecology, demography, and genetic structure than previously thought.

Balkan brachicerous cattle - the first domesticated cattle in Europe.

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 cattle breed.

Potential hominin affinities of Graecopithecus from the Late Miocene of Europe.

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.

An independent event of Neolithic cattle domestication on the South-eastern Balkans: evidence from prehistoric aurochs and cattle populations.

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.

Messinian age and savannah environment of the possible hominin Graecopithecus from Europe.

Dating fossil hominids and reconstructing their environments is critically important for understanding human evolution. Here we date the potentially oldest hominin, Graecopithecus freybergi from Europe and constrain the environmental conditions under which it thrived. For the Graecopithecus-bearing Pikermi Formation of Attica/Greece, a saline aeolian dust deposit of North African (Sahara) provenance, we obtain an age of 7.37-7.11 Ma, which is coeval with a dramatic cooling in the Mediterranean region at the Tortonian-Messinian transition. Palaeobotanic proxies demonstrate C4-grass dominated wooded grassland-to-woodland habitats of a savannah biome for the Pikermi Formation. Faunal turnover at the Tortonian-Messinian transition led to the spread of new mammalian taxa along with Graecopithecus into Europe. The type mandible of G. freybergi from Pyrgos (7.175 Ma) and the single tooth (7.24 Ma) from Azmaka (Bulgaria) represent the first hominids of Messinian age from continental Europe. Our results suggest that major splits in the hominid family occurred outside Africa.

PESI - a taxonomic backbone for Europe.

BACKGROUND: Reliable taxonomy underpins communication in all of biology, not least nature conservation and sustainable use of ecosystem resources. The flexibility of taxonomic interpretations, however, presents a serious challenge for end-users of taxonomic concepts. Users need standardised and continuously harmonised taxonomic reference systems, as well as high-quality and complete taxonomic data sets, but these are generally lacking for non-specialists. The solution is in dynamic, expertly curated web-based taxonomic tools. The Pan-European Species-directories Infrastructure (PESI) worked to solve this key issue by providing a taxonomic e-infrastructure for Europe. It strengthened the relevant social (expertise) and information (standards, data and technical) capacities of five major community networks on taxonomic indexing in Europe, which is essential for proper biodiversity assessment and monitoring activities. The key objectives of PESI were: 1) standardisation in taxonomic reference systems, 2) enhancement of the quality and completeness of taxonomic data sets and 3) creation of integrated access to taxonomic information. NEW INFORMATION: This paper describes the results of PESI and its future prospects, including the involvement in major European biodiversity informatics initiatives and programs.

Dolichopithecus balcanicus sp. nov., a new Colobinae (Primates, Cercopithecidae) from the early Pliocene of southeastern Europe, with a discussion on the taxonomy of the genus.

We describe a new species of Dolichopithecus, D. balcanicus sp. nov., from the Pliocene (late Ruscinian) of the Balkans. Although known by mandibular remains only, it is readily distinguished from the common European species, D. ruscinensis, by its smaller size, much shallower mandibular corpus, and crowding of the premolars. Ukrainian finds of "Adelopithecus" can be accommodated within D. ruscinensis as a separate, smaller subspecies, D. ruscinensis hypsilophus, but the specimens reported from Asia are more distinct, and they probably represent a separate subgenus of Dolichopithecus, the valid name of which is D. (Parapresbytis) Kalmykov and Mashchenko.