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Predrag Radović
  • Kraljevo, Serbia
  • Paleoanthropologist and vertebrate paleontologist My research focuses on the hominin fossil record (particularly the ... moreedit
Finds of Pleistocene rhinoceros are rare in Serbia, and only one species (the woolly rhinoceros Coelodonta antiquitatis Blumenbach, 1799) has been reported so far. The current paper presents the dental material of an extinct so-called... more
Finds of Pleistocene rhinoceros are rare in Serbia, and only one species (the woolly rhinoceros Coelodonta antiquitatis Blumenbach, 1799) has been reported so far. The current paper presents the dental material of an extinct so-called Hundsheim rhinoceros, Stephanorhinus hundsheimensis Toula, 1902 from Bogovina Cave (Eastern Serbia). Both the morphological and metric characteristics of the teeth are consistent with the attribution to S. hundsheimensis. Unfortunately, the rhinoceros material originated from an uncertain geological context, so there is no firm basis for dating. However, a potential association with a caballoid horse tooth could hint at an age of 600 ka or less for the assemblage, which would indicate a relatively late date for the Hundsheim rhinoceros from Bogovina Cave.
The Central Balkans represents a significant geographical gap in the human fossil record of Eurasia. Here we present two new human fossils from Pešturina Cave, Serbia: a partial atlas vertebra (C1) and a fragment of radial diaphysis. The... more
The Central Balkans represents a significant geographical gap in the human fossil record of Eurasia. Here we present two new human fossils from Pešturina Cave, Serbia: a partial atlas vertebra (C1) and a fragment of radial diaphysis. The atlas (Pes-1) derives from the lower portion of Layer 2 and conforms to modern human morphology. This layer is characterized by Gravettian industry despite uncertainties caused by bioturbation and difficulties in separating Layers 2 and 3. The radial fragment (Pes-2) was recovered from the contact zone between Layers 3 and 4, both of which represent Mousterian industries, and is tentatively assessed as Neanderthal based on morphology. With the recent publication of a Neanderthal molar (Pes-3) from the same site, Pešturina currently stands as the only Neanderthal fossil-bearing site in Serbia. These additional finds make Pešturina Cave the only site in the Central Balkans which preserves both modern human and Neanderthal remains with associated lithic industries and highlights the importance of Pešturina in the current discourse on hominin dispersals and migrations in the Balkans.
The paper provides an overview of the Neogene fossil mammals recorded in the vicinity of Kraljevo in Central Serbia. Large mammals are represented exclusively by the accidental finds of proboscidean dental specimens, deposited both in the... more
The paper provides an overview of the Neogene fossil mammals recorded in the vicinity of Kraljevo in Central Serbia. Large mammals are represented exclusively by the accidental finds of proboscidean dental specimens, deposited both in the Čačak-Kraljevo and Gruža basins. We provide re-descriptions and comparative analyses of the proboscidean material. The following taxa are recorded: Deinotherium giganteum at Ravanica and Adrani; Gomphotherium angustidens at Bogutovac; Tetralophodon longirostris at Godačica and at an unidentified sand pit (majdan) near Kraljevo; Anancus sp. at Miločaj; Proboscidea indet. at Oplanići. Small fossil mammals are recorded at three sites. The Progorelica assemblage is currently attributed to the MN6 zone of the Middle Miocene (Langhian), based on the finds of Eulipotyphla sp., Eomyops sp., Cricetodon sp., Megacricetodon sp. and Alloptox sp. – one of the first finds of this Asian genus in the Balkan Peninsula. At Tavnik, more than 120 remains of small mammals were attributed to the MN9 zone of the Late Miocene (Tortonian); the site yielded remains of Eulipotyphla indet., Prolagus sp., Megacricetodon similis and Miodyromys sp. Two fossil teeth of Talpidae sp. were also recorded at Oplanići.
The cercopithecid fossil record of the Balkan Peninsula extends from the Late Miocene to the Early Pleistocene, but to date no fossils of non-human primates have been identified in Serbia. Here we report the identification of two primate... more
The cercopithecid fossil record of the Balkan Peninsula extends from the Late Miocene to the Early Pleistocene, but to date no fossils of non-human primates have been identified in Serbia. Here we report the identification of two primate teeth from Ridjake, a rich paleontological site in western Serbia. NHMBEO 042501 is an upper third molar with heavy occlusal wear and taphonomic weathering. NHMBEO 042502 is a well-preserved lower third molar with only minor damage to the cusps and root apices. We performed an analysis of non-metric traits and made bivariate comparisons of crown linear measurements in order to assess the taxonomic affinity of the molars. Both show typical papionin occlusal patterns and relatively large overall sizes. In combination with the early Villafranchian (MN16) age of the site, we attribute both Ridjake primate fossils to cf. Paradolichopithecus sp. This represents the first identification of a non-human primate in Serbia, and the first identification of any primate in the Neogene period of Serbia. Along with recent hominin discoveries, the Ridjake fossils contribute to the growing primate fossil record in Serbia, and indicate the need for increased research into fossil primates in the country.
Neanderthals were the only human group in Europe throughout the Late Pleistocene until the arrival of modern humans, and while their presence has been confirmed in the surrounding regions, no Neanderthal fossils are known to date from the... more
Neanderthals were the only human group in Europe throughout the Late Pleistocene until the arrival of modern humans, and while their presence has been confirmed in the surrounding regions, no Neanderthal fossils are known to date from the Central Balkans. Systematic excavations of Pešturina Cave (Serbia) resulted in the discovery of a permanent right M1 (Pes-3). The specimen was recovered from stratigraphic Layer 4b with an estimated age of 102.4 ± 3.2 ka, associated with Mousterian artifacts. The exceptional state of preservation and minimal wear of the molar enabled a detailed description and comparative analysis of the inner and outer dental structure, including non-metric dental traits and morphometric features of the crown, roots, and dental tissues. The results of this study strongly support the identification of Pes-3 as Neanderthal. Non-metric traits of the occlusal surface of the crown, enamel-dentine junction, and roots are consistent with Neanderthal morphology. The crown shows morphometric features typical for Neanderthal M1, such as a buccolingually skewed crown shape, internally compressed cusps, and a relatively large hypocone. The specimen also shows Neanderthal-like dental tissue proportions, characterized by relatively thin enamel and large coronal dentine and coronal pulp volumes. The discovery of the Pes-3 molar therefore confirms the presence of Neanderthals in the territory of Serbia and the Central Balkans at the end of Marine Isotope Stage (MIS) 5c.
The hominin mandible BH-1 from the Middle Pleistocene cave of Mala Balanica suggested the possibility that human populations in this part of the continent were not subject to the process of Neanderthali-zation observed in the west. We... more
The hominin mandible BH-1 from the Middle Pleistocene cave of Mala Balanica suggested the possibility that human populations in this part of the continent were not subject to the process of Neanderthali-zation observed in the west. We review the paleoanthropological evidence from the Central Balkans in the context of the Eastern Mediterranean geographic entity. The current hominin fossil record of the early Middle Pleistocene in the region suggests that Europe was inhabited by two different populations: a population in the west of the continent with derived Neanderthal morphology; and a more variable population in the east characterized by a combination of plesiomorphous and synapomorphous traits. We suggest that e in order to continue using the nomenclature of Homo heidelbergensis e the current hypodigm needs to be revised to include only the specimens from the latter group.
Elephantoid cheek teeth from the late Early and Middle Miocene of Europe frequently display mixtures of bunodont and zygodont features, making their taxonomical attributions difficult and subjective. Today, these teeth with “intermediate”... more
Elephantoid cheek teeth from the late Early and Middle Miocene of Europe frequently display mixtures of bunodont and zygodont features, making their taxonomical attributions difficult and subjective. Today, these teeth with “intermediate” morphologies are attributed either to the mammutid species Zygolophodon turicensis, or to the gomphothere genus Gomphotherium - as a variation of G. angustidens or as a separate species G. subtapiroideum. This paper presents one such specimen, a fragmented lower third molar which originated from Miocene sandstones within the Kruševac Basin (possibly from the village of Bela Voda) in Central Serbia. We described the fossil and examined its metric properties in comparative context. Furthermore, we applied the semiquantitative method of Wang et al. (2016) in order to reduce subjectivity in our assessment of the degree of specimen’s zygodonty. Our results suggested that the specimen resembles Z. turicensis more closely than either G. angustidens and G. subtapiroideum, both in terms of the metrics and the degree of zygodonty. However, we were not able to make a firm taxonomical attribution, due to the fact that the specimen represents an isolated and incomplete fossil.
A fragmented right branch of a ruminant mandible from Rusce (Pčinja basin, Serbia) was originally published in the first half the twentieth century as Micromeryx flourensianus, a small ruminant common in the middle Miocene of Europe.... more
A fragmented right branch of a ruminant mandible from Rusce (Pčinja basin, Serbia) was originally published in the first half the twentieth century as Micromeryx flourensianus, a small ruminant common in the middle Miocene of Europe. Based on this determination, sedimentary filling of the Pčinja basin was considered to be of late Miocene age. However, later paleobotanical and micromammalian studies pointed to a late Eocene age for these deposits. The redescription and discussion of the ruminant fossil mandible from Rusce led to the conclusion that the specimen may belong to a small species of Bachitheriidae, probably to Bachitherium thraciensis. This ruminant was originally only known from late Eocene strata in Bulgaria. The peculiar late Eocene faunal composition from the Balkans (e.g., rodents, perissodactyls, and ruminants) confirms that the “Balkanian High” was a distinct paleobiogeographical province from that of Western Europe until the Bachitherium dispersal event, which occurred during the early Oligocene ca. 31 Mya.
There was a fauna of baleen whales (Cetacea: Mysticeti) in the Central Paratethys, a Miocene water body which covered the area of present-day Central Europe. Most of the fossil baleen whales from the Central Paratethys have been found in... more
There was a fauna of baleen whales (Cetacea: Mysticeti) in the Central Paratethys, a Miocene water body which covered the area of present-day Central Europe. Most of the fossil baleen whales from the Central Paratethys have been found in strata assigned to the regional Badenian age and comprise endemic forms, known only from this region. Here a new description is provided for a fragmentary skeleton of a whale found in Belgrade, Serbia, and its age context and biological aspects are discussed. This specimen, consisting of a fragmentary horizontal ramus of the mandible and eight caudal vertebrae, is tentatively identified as ?Parietobalaena sp., a globally known Miocene taxon, with related forms having been recorded earlier from the Eastern Paratethys. An early Sarmatian age (i.e., about 12.7-12.4 Ma) is proposed for the specimen. Based on epiphyseal fusion of caudal vertebrae, the specimen's age is identified as a subadult, with a body length of around 2.5 m and an estimated adult body length around 3-3.5 m, which is extremely small for baleen whales. This find represents one of the latest records of baleen whales in the Central Paratethys and one of the latest published records of Parietobalaena-like taxa in the world.
The paper presents the description of six isolated fossil teeth discovered in 2011 at Sibnica 4 locality (Central Serbia). Based on the morphological characteristics, the material is identified as belonging to a small fossil moschid... more
The paper presents the description of six isolated fossil teeth discovered in 2011 at Sibnica 4 locality (Central Serbia). Based on the morphological characteristics, the material is identified as belonging to a small fossil moschid Micromeryx sp., which makes the first occurrence of this genus in Serbia. Moreover, the associated small mammalian fauna suggests MN4 age for the fossils from Sibnica, constituting the earliest record of this genus discovered so far.
The palaeoecology of fossil molluscs of Serbia is insufficiently studied. Drilling predation indicates interactive relationships among Middle Miocene Badenian molluscs from the Rakovica sands. These sands are present in the Rakovica... more
The palaeoecology of fossil molluscs of Serbia is insufficiently studied. Drilling predation indicates interactive relationships among Middle Miocene Badenian molluscs from the Rakovica sands. These sands are present in the Rakovica stream valley (southern part of Belgrade), and well-known data about the fauna has existed since the end of the XIX century. The collected material (one bulk sample) from the section was used for studies that included 2301 shells of molluscs (1617 of gastropods and 684 of bivalves). Among them were specimens with predatory drill holes. Determining the value of drilling frequency (DF) and prey effectiveness (PE) facilitates identification of the relationship between shell-drilling predators and their prey. Based on analysis of the overall association, carnivorous gastropods were dominant in the sample, comprising more than fifty percent (52.32%) of all gastropod specimens. In addition, they constitute a large percentage of the total sample (43.19%). The largest number of victims was observed among bivalves, which are indicated by a DF of 16.96%. The results were compared with other localities of Miocene age in Paratethys and with studies on the Miocene sediments from the Boreal and the Atlantic Provinces. These comparisons show that PE and DF are very similar to results from the Central Paratethys, but that they are significantly higher than in the other are as mentioned.
The paper presents the first detailed description and the first analysis of the fossil colobine monkey Mesopithecus housed in the collections of the Natural History Museum in Belgrade, Serbia. The material is... more
The  paper  presents  the  first  detailed  description  and  the  first  analysis  of  the  fossil  colobine  monkey  Mesopithecus  housed  in  the  collections  of  the  Natural  History Museum in Belgrade, Serbia. The material is described using the standard methodology  and  analyzed  in  a  comparative  context  of  specimens  from  the  published literature (n=45), by using bivariate and multivariate statistical methods (principal  component  analysis).  We  determined  several  species:  M.  pentelicus, M. delsoni, M. cf.  pentelicus, M. cf.  delsoni.  The  associated  fauna  suggests  MN11  to  MN12 (Turolian) age, and a relatively open landscape – bushy or sparsely wooded savannah. However, further research into the associated collection will allow us to discuss biostratigraphic and paleoecological characteristics in more detail.
tA fragmented human calotte was discovered during the early 1950snear Baˇcki Petrovac (Serbia), in association with Palaeolithic stonetools. After its initial publication, the fossil specimen remainedlargely unknown outside of the Serbian... more
tA fragmented human calotte was discovered during the early 1950snear Baˇcki Petrovac (Serbia), in association with Palaeolithic stonetools. After its initial publication, the fossil specimen remainedlargely unknown outside of the Serbian academe and no detailedcomparative study has ever been carried out. Since the whereaboutsof the fossil itself are currently unknown, and given its potentialsignificance for the Pleistocene human evolution, we re-examinethe data published byˇZivanovi´c (1966, 1975). Using the originalmeasurements, mostly taken on the frontal bone, and a wide com-parative sample of 68 fossil specimens, the fossil was compared andanalyzed by statistical multivariate methods. We also conducted avisual examination of the morphology based on the available pho-tographic material. Our analysis reveals phenetic similarity withMiddle Pleistocene archaic Homo from Africa and anatomicallymodern Homo sapiens. However, the absence of primitive cranialtraits in Baˇcki Petrovac indicates a clear modern Homo sapiens des-ignation. Although lost at the moment, there is a chance for there-discovery of the fossil in the years to come. This would give usan opportunity to acquire absolute dates and to study the specimenin a more detailed manner.
With the exact nature of the interactions between Neanderthals and anatomically modern humans, and the identity of the of the bearers of early Upper Paleolithic technology still open questions essential to expand the human fossil data... more
With the exact nature of the interactions between Neanderthals and anatomically modern humans,
and the identity of the of the bearers of early Upper Paleolithic technology still open questions essential
to expand the human fossil data of Southeast Europe. In our attempt to do so, we investigated a small
collection of six previously unpublished human cranial fragments from Serbia, housed at the Natural History
Museum in Belgrade and the National Museum in Kraljevo. Tenuous contextual evidence suggested a
possible Pleistocene age for the specimens. We conducted a macro-morphological analysis and accelerator
mass spectrometry radiocarbon dating in order to assess taxonomical positions and absolute dates for the
specimens. Thorough prescreening and chemical characterization of bone samples were used to ensure
high reliability of 14C dates. Although the results showed all specimens to be Holocene-aged anatomically
modern humans, this should not discourage future research. On the contrary, if indeed we want to understand
the early presence of modern humans in the Central Balkans, more research is needed. This includes
further checking of old museum collections, but emphasis will need to be placed on new excavations of
Pleistocene sites in the region.
Research Interests:
Research Interests:
An unusual anatomical variation was observed in the skull excavated at Lanište (Raška district, Serbia), a Roman necropolis dating to the second half of the 4th century AD. The skull of an adult male showed a remarkably large... more
An unusual anatomical variation was observed in the skull excavated at Lanište (Raška district, Serbia), a Roman necropolis dating to the second half of the 4th century AD. The skull of an adult male showed a remarkably large supernumerary bone at bregma co-occurring with a continuous persistent metopic suture. Few similar cases have been reported in scientific literature. We describe the case and discuss possible mechanisms and underlying causes, including pathological conditions.
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A relatively small number of Lower and Middle Palaeolithic open-air sites have been registered in southeast Europe. There are only two regions where sites from this period have been systematically surveyed, and where they have been... more
A relatively small number of Lower and Middle Palaeolithic open-air sites have been registered
in southeast Europe. There are only two regions where sites from this period have been systematically
surveyed, and where they have been encountered in considerable numbers: the valley of the Pineios river
in Greece and northern Bosnia. Not a single site with significant concentration of finds has been found
in Serbia and isolated artifacts ascribed to the Middle Palaeolithic were found at few sites. All this has
changed dramatically in recent years when a large number of sites with finds from the Lower and Middle
Palaeolithic have been recorded in the zone from Čačak to Kraljevo in the West Morava valley. The site
surveying in the area has opened an entirely new perspective in the investigation of the Palaeolithic in this
part of the Balkans.
Research Interests:
Research Interests:
Pešturina cave is located in Southern Serbia on a tributary of the Nišava River, in the vicinity of Sićevo Gorge (43°10′ N, 21°54′ E). Together with Mala and Velika Balanica, the deposits cover the last 500 ka of likely continuous human... more
Pešturina cave is located in Southern Serbia on a tributary of the Nišava River, in the vicinity of Sićevo Gorge (43°10′ N, 21°54′ E). Together with Mala and Velika Balanica, the deposits cover the last 500 ka of likely continuous human occupation in the region. We obtained a series of dates by combined ESR and 14C methods that constrain the three distinct archaeological layers between 108 ka and 29 ka. From the top to bottom of the sequence layers 2, 3 and 4 represent Pleistocene accumulations with Gravettian, Denticulate Mousterian, and Charentian Mousterian, respectively. Radiocarbon dates on faunal bones show that materials have been mixed between lithostratigraphic layers. 14C dates of faunal bones with cut marks provide two secure ranges during which human occupation occurred: between 33-29 ka cal BP, associated with the Gravettian, and 45-43 ka cal BP, associated with Mousterian; layer 4 was dated to between app. 83 and 108 ka by ESR. Hominin remains are distributed unevenly through the layers: A maxillary P3 crown of a juvenile individual was excavated from the layer 2; however, its attribution to the Pleistocene is not confirmed as the layer had some Holocene intrusions. In addition, the layer 2 also produced a fragment of a human cervical vertebra. All other hominin fossil material comes from layer 4: a shaft fragment of a juvenile human radius, an adult femoral head, and a maxillary M1 of an adult individual. We compare the morphology of these specimens to the published data on Neanderthals and modern humans of comparable geological age. The maxillary premolar is consistent with modern human morphology while the maxillary molar is consistent with Neanderthals. In addition, we note the robusticity of all postcranial remains. The addition of this material to our understanding of Neanderthal presence in the region is important. Further excavations as well as aDNA analyses (currently underway) will shed more light on the relationship of this material to adjoining regions which are increasingly recognized as key to our understanding of migrations in Europe.
A fragmented human calotte was discovered during the early 1950s near Bački Petrovac (Serbia), in association with Palaeolithic stone tools. After its initial publication, the fossil specimen remained largely unknown outside of the... more
A fragmented human calotte was discovered during the early 1950s near Bački Petrovac (Serbia), in association with Palaeolithic stone tools. After its initial publication, the fossil specimen remained largely unknown outside of the Serbian academe and no detailed comparative study has ever been carried out. Since the whereabouts of the fossil itself are currently unknown, and given its potential significance for the Pleistocene human evolution, we re-examine the data published by Živanović (1966, 1975). Using the original measurements, mostly taken on the frontal bone, and a wide comparative sample of 68 fossil specimens, the fossil was compared and analyzed by statistical multivariate methods. We also conducted a visual examination of the morphology based on the available photographic material. Our analysis reveals phenetic similarity with Middle Pleistocene archaic Homo from Africa and anatomically modern Homo sapiens. However, the absence of primitive cranial traits in Bački Petrovac indicates a clear modern Homo sapiens designation. Although lost at the moment, there is a chance for the re-discovery of the fossil in the years to come. This would give us an opportunity to acquire absolute dates and to study the specimen in a more detailed manner.
Research Interests:
The text describes the development of beekeeping in Serbia from ancient times to the present day, with special reference to the improvement of beekeeping in Kraljevo and its surroundings.