Southern Balkans Research Articles

The last amphisbaenian (Squamata) from continental Eastern Europe

The fossil record of amphisbaenians in Europe has recently been augmented through the description of new finds across the continent. It has been suggested that the post-Miocene range of amphisbaenians followed a progressive southward constriction that ultimately lead to their extant disjunct distribution. We herein describe amphisbaenian vertebral material from the Pliocene of northern Greece, which demonstrates that these reptiles were still present in the late Neogene of the southern Balkans. The new find represents the youngest occurrence of amphisbaenians in continental Eastern Europe and further highlights the role of the Greek area as a biogeographic “refugium” for certain reptile groups during the late Neogene and Quaternary.

Little Ice Age glaciers and climate in the Mediterranean mountains: a new analysis

Glaciers were common across the Mediterranean mountains during the Little Ice Age. In parts of Turkey some glaciers were several kilometres longer than they are today, whilst in the Pyrenees glaciers were up to several hundred metres longer. In the wettest Mediterranean mountains, such as the Dinaric Alps, many small glaciers and perennial snow patches would have been present. Even in driest and most southerly mountains, such as the High Atlas, small glaciers and perennial snowfields were present. This paper examines the evidence from these two contrasting regions (the western and southern Balkans and the High Atlas) and the climatic significance of glaciers in these areas during the Little Ice Age. Particular focus is given on the climatological controls on glacier mass balance in different climatic conditions. Glaciers in cold and dry climates exhibit different sensitivity to regional climate change compared with glaciers in cold and wet climates. In addition, the factors controlling ablation of glaciers in different climatic regimes can differ considerably, especially the relative contributions and effects of melting and sublimation. All Mediterranean mountain glaciers were strongly controlled by local topoclimatic factors. Avalanche-fed glaciers have proven to be the most resilient to climate change and dramatically increased accumulation from avalanching snow explains the surviving glaciers in the Dinaric Alps and the semi-perennial snow fields of the High Atlas. In addition, geology as well as landscape morphology inherited from Pleistocene glaciations plays a role in explaining the patterns of Little Ice Age glacier distribution and especially the patterns of retreat and survival of these glaciers. The resilience of some of the last remaining Mediterranean glaciers, in the face of warming climate, presents a contradiction and comparisons between glaciers gone and those that remain provides important insight into the future of similar glaciers globally.

Scrub communities along a climatic gradient in the southern Balkans: maquis, pseudomaquis and shibljak

The research deals with three scrub communities/formations in the southern Balkans, maquis, pseudomaquis and shibljak, and their changes along the main (macro)ecological gradient. We performed DCA analysis and, since the first axis correlates with climatic data, the projection of relevés on this axis was accepted as proxy for the (macro)ecological gradient, and the species turnover along this gradient was elaborated. Linear regression was used to test the turnover of life forms, chorotypes and the life strategy of species (syntaxonomical affiliation). It was found that life forms, except hemicryptophytes and nanophanerophytes, do not change significantly along the gradient change; most chorotypes change significantly; among species, life strategies (syntaxonomical affiliation), the proportion of species of deciduous and evergreen forests and scrub, as well as species of dry grassland, change significantly. The classification established three clusters, representing three formations and we tested differences among them. It was established that stenomediterranean and Eurasian species and species of perennial grasslands differentiate all three clusters and some groups only two of them. We found that discontinuous formations can be established along a continuous gradient. Communities were also classified within the present synsystematic system.

Extra-Mediterranean glacial refuges in barred and common grass snakes (Natrix helvetica, N. natrix)

Extra-Mediterranean glacial refugia of thermophilic biota, in particular in northern latitudes, are controversial. In the present study we provide genetic evidence for extra-Mediterranean refugia in two species of grass snake. The refuge of a widely distributed western European lineage of the barred grass snake (Natrix helvetica) was most likely located in southern France, outside the classical refuges in the southern European peninsulas. One genetic lineage of the common grass snake (N. natrix), distributed in Scandinavia, Central Europe and the Balkan Peninsula, had two distinct glacial refuges. We show that one was located in the southern Balkan Peninsula. However, Central Europe and Scandinavia were not colonized from there, but from a second refuge in Central Europe. This refuge was located in between the northern ice sheet and the Alpine glaciers of the last glaciation and most likely in a permafrost region. Another co-distributed genetic lineage of N. natrix, now massively hybridizing with the aforementioned lineage, survived the last glaciation in a structured refuge in the southern Balkan Peninsula, according to the idea of ‘refugia-within-refugia’. It reached Central Europe only very recently. This study reports for the first time the glacial survival of a thermophilic egg-laying reptile species in Central Europe.

Spiralled patchwork in pottery manufacture and the introduction of farming to Southern Europe—ERRATUM

Pottery-manufacturing sequences can act as proxies for human migration and interaction. A good example is provided by the �spiralled patchwork technology� (SPT) identified at two key early farming sites in the Ligurian-Provencal Arc in the north-west of the Italian peninsula. SPT is distinct from the ceramic technology used by early farmer communities in south-east Italy that shows technical continuity with the southern Balkans. Macroscopic analysis and micro-computed tomography suggests the presence of two communities of practice, and thus two distinct social groups in the northern Mediterranean: one of southern Balkan tradition, the other (associated with SPT) of as yet unknown origin. The identification of SPT opens up the exciting possibility of tracing the origins and migrations of a second distinct group of early farmers into Southern Europe

A MIS 9/MIS 8 speleothem record of hydrological variability from Macedonia (F.Y.R.O.M.)

The period corresponding to Marine Isotope Stages 9 (MIS 9) offers the opportunity to study orbital and sub-orbital scale climate variability under boundary conditions different from those of better studied intervals such as the Holocene and the Last Interglacial. Yet, it is poorly represented in independently-dated continental archives around the Mediterranean Region. Here, we present a speleothem stable isotope record (δ18O and δ13C) from the Former Yugoslavian Republic of Macedonia (F.Y.R.O.M., southern Balkans), which consists of two periods of growth broadly covering the ca. 332 to 292 ka and the ca. 264 to 248 ka intervals (MIS 9e-b and late MIS 8). We interpret the speleothem δ18O as mostly related to regional hydrology, with variations that can be interpreted as due to changes in rainfall amount, with higher/lower values associated to drier/wetter condition. This interpretation is corroborated by a change in mineralogical composition between aragonite and calcite at ca. 328 ka, which marks increasing precipitation at the onset of MIS 9 and occurs within a trend of decreasing δ18O values. Also the comparison with the multiproxy climate record available from the nearby Lake Ohrid seems to support the proposed interpretation. The MIS 9e interglacial appears to be characterized by wettest conditions between ca. 326 and 321 ka, i.e. lasting ca. 5 kyr. Decreasing precipitation and enhanced millennial scale variability matches the glacial inception (MIS9 d to b), with drier events at ca. 319 ka (ca. 2 kyr long) and 310 ka (ca. 1 kyr long), and a major rainfall reduction between 306 and 298 ka. The latter is followed by a prominent wetter period between 298 and 295 ka, for which carbon data values suggest high infiltration rate. Rainfall decreases again after 295 ka, and remain low until the growth interruption at ca. 292 ka. Resumption of the growth and progressive soil development, expressed by the carbon isotope record, occurred during the late part of MIS 8. Despite the rather high temporal uncertainty (average 6 ka), the speleothem hydrological record complements the environmental information provided by the Lake Ohrid record and also fits well to the framework of regional and extra-regional variability, showing similarities with pollen records from southern and western Europe, both at orbital and at sub-orbital time scale.

Reconstructing the genetic history of Italians: new insights from a male (Y-chromosome) perspective

Background: Due to its central and strategic position in Europe and in the Mediterranean Basin, the Italian Peninsula played a pivotal role in the first peopling of the European continent and has been a crossroad of peoples and cultures since then.Aim: This study aims to gain more information on the genetic structure of modern Italian populations and to shed light on the migration/expansion events that led to their formation.Subjects and methods: High resolution Y-chromosome variation analysis in 817 unrelated males from 10 informative areas of Italy was performed. Haplogroup frequencies and microsatellite haplotypes were used, together with available data from the literature, to evaluate Mediterranean and European inputs and date their arrivals.Results: Fifty-three distinct Y-chromosome lineages were identified. Their distribution is in general agreement with geography, southern populations being more differentiated than northern ones.Conclusions: A complex genetic structure reflecting the multifaceted peopling pattern of the Peninsula emerged: southern populations show high similarity with those from the Middle East and Southern Balkans, while those from Northern Italy are close to populations of North-Western Europe and the Northern Balkans. Interestingly, the population of Volterra, an ancient town of Etruscan origin in Tuscany, displays a unique Y-chromosomal genetic structure.

Spiralled patchwork in pottery manufacture and the introduction of farming to Southern Europe

Abstract

First thermochronological constraints on the Cenozoic extension along the Balkan fold-thrust belt (Central Stara Planina Mountains, Bulgaria)

The Balkan fold-thrust belt, exposed in Bulgaria and north-east Serbia, is part of the north-east vergent segment of the bi-vergent Eastern Mediterranean Alpine orogen. It was formed during two distinct compressional stages; the first one lasted from the Middle Jurassic to the Early Cretaceous and the second from Late Cretaceous to the Paleogene. Although the compressional tectonic evolution of the Balkan fold-thrust belt since the Middle Jurassic and during most of the Mesozoic is relatively well studied, the final exhumation of the rocks of the belt during the Cenozoic has remained poorly understood. Here, we present the first thermochronological constraints, based on fission-track and [U-Th-(Sm)]/He analysis, showing that along the central part of the belt syn- to post-orogenic extension could have started as early as the middle Eocene. Low-temperature thermochronological analysis of samples collected from three areas reveals at least two phases of increased cooling and exhumation during the Cenozoic. The first exhumation phase took place between ~ 44 and 30 Ma and appears to be related to the syn- to post-orogenic collapse coeval with the earliest Cenozoic extensional stage observed across the southern Balkan Peninsula. A period of relative quiescence (between ~ 30 and ~ 25 Ma) is followed by the next cooling stage, between 25 and 20 Ma, which appears to be related to late Oligocene to early Miocene crustal extension across the Balkan Peninsula. Extension accommodated by the late Miocene to Recent age Sub-Balkan Graben System does not appear to have produced exhumation of rocks from beneath 2–4 km depth, as it was not detected by the low-temperature thermochronological methods applied in this study.

Brown hares ( Lepus europaeus Pallas, 1778) from the Balkans: a refined phylogeographic model

The contemporary geographical distribution and genetic structure of temperate species have been strongly influenced by the climatic oscillations during the Late Quaternary. As spatial genetic reconstructions are often markedly affected by geographically meaningful sample distributions, we focused in our study on the analyses of mtDNA control region sequences of brown hares from different regions in northern, central and south-central Balkans that have so far not been covered, with the aim to delineate the most likely glacial refugia wherefrom the postglacial northward expansion into central Europe has originated. Three major haplogroups (“Anatolia/Middle East”, “the Balkans”, and “central Europe”) were revealed with apparent south-north gradual decrease in molecular diversity indices. Moreover, phylogenetic and demographic history analyses identified the southeastern central Balkans as the putative origin for most populations from the southern and northern Balkans, while populations from central and northwestern Europe have originated from the northern Balkans. Click here to access supplementary files .

Vulture mortality resulting from illegal poisoning in the southern Balkan Peninsula.

A study was carried out to determine the incidents of illegal poisoning of griffon vulture (Gyps fulvus), Egyptian vulture (Neophron percnopterus), black vulture (Aegypius monachus), and bearded vulture (Gypaetus barbatus) in the southern Balkan Peninsula between 1982 and 2017. A total of 38 poisoning cases affecting 224 vultures were analyzed to identify their causes and the primary target species for poisoning. Nine different compounds were used in these incidents and the most frequently applied were strychnine, carbamate, and organophosphoros compounds. The poison used to kill gray wolf had the most significant collateral damage to the vulture populations in comparison to the other investigated reasons. It was the primary cause of 60% of all registered vulture poisoning events in the southern Balkan Peninsula during the last 36years. Establishing permanent feeding sites for vultures in areas with wolves appears to be an effective way to minimize the risk of poisoning. There is a pressing need for the development of an appropriate conservation practice taking into consideration relationships among the main and casual target species for poisoning as an essential element in conjunction with the human activities.

Nomenclatural notes and typifications in Campanula versicolor (Campanulaceae) and related names

Campanula versicolor is a member of the Campanula pyramidalis complex. It is distributed in the southern Balkan Peninsula, with a small disjunct range in SE Italy (Puglia and Basilicata administrative regions). Due to its high morphological variability, 17 taxa have been described (at specific and infraspecific level). However, the taxonomic status of these taxa is not clear. In modern floristic literature and checklists they are considered as synonyms within broadly defined C. versicolor. Considering the fact that misinterpretations of their taxonomy in floras and checklists might be caused by unresolved nomenclatural issues, after studying the original material from relevant herbarium collections, we designated lectotypes or epitypes for the following names: C. corymbosa, C. planiflora, C. plasonii, C. rosanii, C. tenorei, C. versicolor, C. versicolor f. mrkvickana, C. versicolor subsp. thessala subvar. lancifolia, C. versicolor var. rosanii, C. versicolor var. thessala, C. versicolor var. thessala f. tomentella, C. versicolor var. tomentella and C. willdenowiana. Note on the already typified C. mrkvickana is also provided.

Hybridization patterns in two contact zones of grass snakes reveal a new Central European snake species

Recent studies found major conflicts between traditional taxonomy and genetic differentiation of grass snakes and identified previously unknown secondary contact zones. Until now, little is known about gene flow across these contact zones. Using two mitochondrial markers and 13 microsatellite loci, we examined two contact zones. One, largely corresponding to the Rhine region, involves the western subspecies Natrix natrix helvetica and the eastern subspecies N. n. natrix, whereas in the other, more easterly, contact zone two lineages meet that are currently identified with N. n. natrix and N. n. persa. This second contact zone runs across Central Europe to the southern Balkans. Our analyses reveal that the western contact zone is narrow, with parapatrically distributed mitochondrial lineages and limited, largely unidirectional nuclear gene flow. In contrast, the eastern contact zone is very wide, with massive nuclear admixture and broadly overlapping mitochondrial lineages. In combination with additional lines of evidence (morphology, phylogeny, divergence times), we conclude that these differences reflect different stages in the speciation process and that Natrix helvetica should be regarded as a distinct species. We suggest a nomenclatural framework for presently recognized grass snake taxa and highlight the need for reconciling the conflicts between genetics and taxonomy.

Microevolution of the noble crayfish (Astacus astacus) in the Southern Balkan Peninsula

BackgroundThe noble crayfish (Astacus astacus) displays a complex historical and contemporary genetic status in Europe. The species divergence has been shaped by geological events (i.e. Pleistocene glaciations) and humanly induced impacts (i.e. translocations, pollution, etc.) on its populations due to species commercial value and its niche degradation. Until now, limited genetic information has been procured for the Balkan area and especially for the southernmost distribution of this species (i.e. Greece). It is well known that the rich habitat diversity of the Balkan Peninsula offers suitable conditions for genetically diversified populations. Thus, the present manuscript revisits the phylogenetic relationships of the noble crayfish in Europe and identifies the genetic make-up and the biogeographical patterns of the species in its southern range limit.ResultsMitochondrial markers (i.e. COI and 16S) were used in order to elucidate the genetic structure and diversity of the noble crayfish in Europe. Two of the six European haplotypic lineages, were found exclusively in Greece. These two lineages exhibited greater haplotypic richness when compared with the rest four (of “Central European” origin) while they showed high genetic diversity. Divergence time analysis identified that the majority of this divergence was captured through Pleistocene, suggesting a southern glacial refugium (Greece, southern Balkans). Furthermore, six microsatellite markers were used in order to define the factors affecting the genetic structure and demographic history of the species in Greece. The population structure analysis revealed six to nine genetic clusters and eight putative genetic barriers. Evidence of bottleneck effects in the last ~5000 years (due to climatic and geological events and human activities) is also afforded. Findings from several other research fields (e.g. life sciences, geology or even archaeology) have been utilized to perceive the genetic make-up of the noble crayfish.ConclusionsThe southernmost part of Balkans has played a major role as a glacial refugium for A. astacus. Such refugia have served as centres of expansion to northern regions. Recent history of the noble crayfish in southern Balkans reveals the influence of environmental (climate, geology and/or topology) and anthropogenic factors.

Lithic production strategies in the Middle Paleolithic of the southern Balkans

One of the major challenges in Paleolithic research is disentangling various factors that contributed to variation in Neandertal technological behavior. As a result of variable abundance of the Paleolithic record across Eurasia, our understanding of regional and temporal patters of Middle Paleolithic industries differs from one region to another. Southeastern Europe is one of the areas that stands in contrast to rich and thoroughly investigated regions of other parts of Europe. A few sites in the region, nevertheless, offer an opportunity for the study of technological behavior of Middle Paleolithic hominins. The rockshelter sites of Crvena stijena and Bioče in Montenegro, located on the eastern Adriatic coast in the Mediterranean zone, have relatively dense remains of hominin occupations compared to other sites in the Balkan peninsula. Previous research identified several characteristics of the eastern Adriatic industries. Firstly, they have been characterized as Micromousterian due to the small average size of the artifacts, a feature that potentially results from the size of the local raw material. On typological grounds, some assemblages have been said to show typological similarity to Charentian, and some researchers have emphasized frequent denticulate tools as a late Mousterian feature. In this study we aim to examine lithic production strategies at the two sites based on samples from Crvena stijena deposits that tentatively range from late MIS 5 to MIS 3 and from the still undated upper sequence of Bioče. We will evaluate variability in the industries in the Adriatic region by looking at both the process of blank production and further tool production and consumption to provide a clearer picture of lithic production strategies during the late Middle Paleolithic. We will address the issue of how raw material affects the structure of these assemblages, artifact size, and particularly diachronic patterns of stone tool production strategies.

Assessment of transboundary river basins for potential hydro-political tensions

Abstract This paper presents a systematic, global assessment of transboundary watersheds that identifies regions more likely to experience hydro-political tensions over the next decade and beyond based upon environmental, political, and economic indicators. The development of new water infrastructure in transboundary basins can strain relationships among fellow riparians as the impacts of new dams and diversions are felt across borders. Formal arrangements governing transboundary river basins, such as international water treaties and river basin organizations, provide a framework for dialogue and negotiation, thus contributing to assuaging potential disputes. Our study examines these two issues in tandem − the stresses inherent in development and the mitigating impact of institutions − and maps the risk of potential hydro-political tensions that exist where basins may be ill-equipped to deal with transboundary disputes triggered by the construction of new dams and diversions. We also consider several factors that could exacerbate those hydropolitical tensions in the near future, including changes in terrestrial water storage, projected changes in water variability, per capita gross national income, domestic and international armed conflicts, and recent history of disputes over transboundary waters. The study points to the vulnerability of several basins in Southeast Asia, South Asia, Central America, the northern part of the South American continent, the southern Balkans as well as in different parts of Africa, where new water infrastructure is being built or planned, but formal transboundary arrangements are absent. Moreover, in some of these regions there is a concomitance of several political, environmental and socioeconomic factors that could exacerbate hydropolitical tensions. This study contributes to the understanding of how the recent proliferation of development accompanied with unfavourable socio-economic and environmental indicators may influence global hydropolitical resilience.

Ancient and recent admixture layers in Sicily and Southern Italy trace multiple migration routes along the Mediterranean

The Mediterranean shores stretching between Sicily, Southern Italy and the Southern Balkans witnessed a long series of migration processes and cultural exchanges. Accordingly, present-day population diversity is composed by multiple genetic layers, which make the deciphering of different ancestral and historical contributes particularly challenging. We address this issue by genotyping 511 samples from 23 populations of Sicily, Southern Italy, Greece and Albania with the Illumina GenoChip Array, also including new samples from Albanian- and Greek-speaking ethno-linguistic minorities of Southern Italy. Our results reveal a shared Mediterranean genetic continuity, extending from Sicily to Cyprus, where Southern Italian populations appear genetically closer to Greek-speaking islands than to continental Greece. Besides a predominant Neolithic background, we identify traces of Post-Neolithic Levantine- and Caucasus-related ancestries, compatible with maritime Bronze-Age migrations. We argue that these results may have important implications in the cultural history of Europe, such as in the diffusion of some Indo-European languages. Instead, recent historical expansions from North-Eastern Europe account for the observed differentiation of present-day continental Southern Balkan groups. Patterns of IBD-sharing directly reconnect Albanian-speaking Arbereshe with a recent Balkan-source origin, while Greek-speaking communities of Southern Italy cluster with their Italian-speaking neighbours suggesting a long-term history of presence in Southern Italy.

The distribution of the invasive pest, rice water weevil Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae), is expanding in Europe: First record in the Balkans, confirmed by CO1 DNA barcoding

This is the first report of the invasive rice pest, rice water weevil (Lissorhoptrus oryzophilus Kuschel) in the Balkans. Adult specimens were collected in the rice field region of Central Macedonia, Greece during July–August, 2016. Morphological identification was verified by DNA barcoding, and the first COI sequence of L. oryzophilus from Greece was deposited in the GenBank database. This semi-aquatic insect is considered one of the most important rice pests globally, and its presence in the region may result in important yield and economic losses in rice production. Considering the highly invasive nature of this species, it is possible that it has spread to neighboring rice wetlands of the Southern Balkans, and additional field investigations are essential to accurately determine its geographic distribution and to help contain its spread.

Crvena Stijena revisited: The Late Mousterian assemblages

Crvena Stijena represents one of the key Middle Paleolithic sites in southeastern Europe. In the course of earlier investigations, the upper part of the Mousterian sequence was excavated on two occasions: in 1956 and 1958 by M. Brodar and from 1961 to 1964 by Đ. Basler. Materials from these campaigns were published separately and interpreted differently. After re-analysis of all these materials, it has been established that the artifact assemblages can not be directly assigned to any of the traditional facies (Denticulate Mousterian, Micromousterian, etc.), but rather to an undifferentiated (non-Levallois) facies of Typical Mousterian, which is characterized by pronounced expedient technology based on the exploitation of local raw materials and the application of ad hoc methods of core reduction and the preparation of tools. This re-analysis pointed out especially that in the final Mousterian at Crvena Stijena, as at several other sites in the Adriatic-Ionian region, blade/bladelet technology and Uluzzian elements (flaking of blades and bladelets from unipolar or bipolar cores on flakes, splintered pieces, backed tools) are present. This seems to indicate continuity and connection among cultural phenomena over the entire Adriatic-Ionian region during MIS3 and suggests the possibility that the Uluzzian might have spread from the southern Balkans towards Italy along the southern rim of the “Great Adriatic Plain,” all of which could have belonged to a single “social territory.”

The rapid spread of early farming from the Aegean into the Balkans via the Sub-Mediterranean-Aegean Vegetation Zone

Close examination of the geographic position of Early Neolithic settlements in SE-Europe shows that the oldest sites are almost exclusively situated in some very specific biogeographic areas. These earliest Neolithic settlements are all concentrated in a region that Pavle Cikovac calls the Sub-Mediterranean-Aegean (SMA) biogeographic region. It covers the northern and north-western edge of the Aegean, including Thessaly, Greek Macedonia and Greek Thrace, and extends further into the Balkans, but only along the valleys of the Vardar (Axios), Struma (Strymon) and Mesta (Nestos) rivers. Surprisingly, although Thrace is the closest landscape to Anatolia, it does not contain any sites of the earliest phase of the Neolithic at all beyond the narrow zone of the northern Marmara. In the present paper we explain this remarkable situation in terms of the natural environment in this particular region of the Southern Balkans. To begin, we propose that the lack of oldest Early Neolithic settlements in Thrace is related to the extreme microclimate of this region. As shown by modern vegetation analogues, Thracian oriental hornbeam-downy oak forests are exposed to stronger continental influence with frosts in the winter and average temperatures during the coldest months that are ca. 2–3 °C lower than those in the Central Balkans that have Sub-Mediterranean vegetation. In general terms, what we may expect is that the earliest Neolithic groups would first appear in regions with similarly mild conditions, on a yearly average, to those in the Mediterranean. Such mild conditions are indeed present in the SMA biogeographic region. On the other hand, before moving further to the north along the north-south oriented river systems of the Central Balkans, the Neolithic economy based on agriculture and stockbreeding would first have to be adapted to the relatively harsh winters in the Balkans. In consequence, it would have been possible to apply the new Neolithic lifestyle in the neighbouring areas of Thrace, Walachia, Dobrudzha and the Carpathian Basin only after a certain period of adaptation. Available 14C-data show that the adaptation period is identical to the time-span of Rapid Climate Change (RCC: 6550-6050 calBC) as defined in previous studies.