Southern Balkans Research Articles

Discovery of Anthophora onosmarum Morawitz, 1876 in Europe (Hymenoptera, Apidae)

Anthophora onosmarum is a poorly known bee species described from the Caucasus (Georgia). New fieldwork has revealed that the species is present in Bulgaria and Greece, and is also widespread across Turkey, predominantly on the Anatolian plateau. New behavioural data shows that A. onosmarum is likely narrowly oligolectic on the plant genus Onosma (Boraginaceae), and is capable of buzzing flowers to release pollen. This finding illustrates the growing trend of overlooked bee species with more Caucasian, Anatolian, or even Middle Eastern distributions occurring in under-surveyed mountainous parts of the Southern Balkans.

Ottoman Tobacco and the Moral Economy of State-Led Development (c. 1908–12)

Abstract In the Second Constitutional Era (1908–18), proponents of various models for national development debated the Ottoman future under the Committee for Union and Progress (cup). Public discussions of cup policies related to the tobacco sector were central to these discussions in the southern Balkans. As such, they provide a window into the political and moral economy of the region and some of the public critiques of international finance after the 1908 Constitutional Revolution. By 1908, tobacco had been at the crossroads of regional economic development and European financial domination for nearly three decades. Since the domestic tobacco monopoly was set to expire in 1913, the region was primed for debates about the agricultural credit market and the customs regime. This article argues that cup proposals for state-led development aimed to satisfy the divergent moral demands of agriculturalists (zürrāʿ) and merchants (tüccar) in the tobacco-producing zones of the empire.

Looking at the modern landscape of submediterranean Greece through a palaeoecological lens

The importance of understanding the long-lasting legacy of past land use on modern ecosystems has long been acknowledged. However, the magnitude and persistence of such legacies have been assessed only occasionally. Northern Greece has been a gateway of farming into mainland Europe during the Neolithic, thus providing a perfect setting to assess the potential impact of land-use history on present-day ecosystems. Additionally, the marked Holocene climatic variability of the southern Balkans makes it possible to investigate climate-vegetation-land use interactions over long timescales. Here, we have studied a sediment record from Limni Vegoritis (Northern Greece) spanning the past ∼9000 years using palaeoecological proxies (pollen, spores, stomata, microscopic charcoal). We aimed to reconstruct long-term vegetation dynamics in submediterranean Greece, to assess the environmental factors controlling them and to establish the legacies of the long history of land use in the modern landscape. We found that the Early Holocene afforestation, mainly oak woodlands, was delayed because of suboptimal moisture conditions. Later, colder and drier conditions during the rapid climate change centred around the ‘8.2 ka event’ triggered woodland opening and the spread of wooded (Juniperus) steppe vegetation. First indicators of farming activities are recorded during this period, but their abundances are too low to explain the concurrent large deforestation episode. Later, pinewoods (probably dominated by Pinus nigra) with deciduous Quercus spread and dominated the landscape for several millennia. These forests experienced repeated multi-centennial setback-recovery episodes associated with land-use intensification, but pines eventually declined ∼2500–2000 years ago during Classical times under heavy land use comprising intense pastoralism. This was the starting point for the present-day landscape, where the main ‘foundation’ taxon of the ancient forests (Pinus cf. nigra) is missing, therefore attesting to the strong imprint that historical land use has left on the modern landscape.

Carpinus austrobalcanica – A new highly polyploid species from the Balkan Peninsula closely related to European hornbeam

The Balkan Peninsula was the primary glacial refugium for many temperate tree species and contributed overproportionally to the postglacial recolonisation of central and eastern Europe. This is also the case for Carpinus betulus, the European hornbeam, whose main glacial refugium was in the Balkans. During our fieldwork in the southern Balkans, we discovered trees with a morphology similar to that of C. betulus, but differing in their rough, highly rigged bark and the shape of their fruit involucre. The aim of this study was to investigate the evolutionary origin and differentiation of these morphologically distinct populations of C. betulus from the North Pindus Mountain range in northern Greece and southern Albania using an integrative approach. Our study combined phylogenetic analyses of plastid and nuclear internal transcribed spacer sequences, amplified fragment length polymorphism fingerprinting, relative genome size estimation, and multivariate morphometric analyses. After establishing the genetic and morphological divergence of the aforementioned populations, we described them as a new species, Carpinus austrobalcanica D.Lakušić, Kuzmanović, Stevanoski, Schönsw. & Frajman, sp. nov. We provide diagnosis, description, geographical distribution, and conservation status of this enigmatic newly described tree species locally endemic to the southern Balkans.

A new perspective on the molecular dating of the stone crayfish with an extended phylogeographic information on the species

Reconstructing the origin and historical biogeography of the Austropotamobius torrentium is hampered by insufficient phylogeographic coverage of the Balkans and deep contradictions in previous molecular dating. The present work extends the phylogeographic coverage to Serbia, a country crucial for understanding the species southward dispersal. Our analysis revealed that the Southern Balkans lineage occurs in most of the country, the Central and southeastern Europe lineage is restricted to the southwest and northeast of the country, while a single population in the north of the country harbors the Lika and Dalmatia lineage, which was previously thought to be restricted to the northern-central Dinarides. Dataset expansion led to revised phylogenetic relationships, which indicated that the Apuseni lineage is not nested within Northern-central Dinarides lineages but arose after the most basal split within Austropotamobius torrentium. This ‘Apuseni first’ phylogeny provides a new perspective for molecular dating, according to which the split between Austropotamobius pallipes and A. torrentium took place in the Late Oligocene, while the formation of the phyletic lineages and the dispersal from the Dinarides to Serbia occurred in the late Miocene and is probably associated with the complex and protracted process of disintegration of the Neogene freshwater lakes in southeastern Europe.

Pollen and Capsule Morphological Studies on Verbascum Hybrids (Scrophulariaceae) in Mardin, Turkey and Their Taxonomic Importance

Verbascum (Scrophulariaceae, Lamiales), usually known as “Mullein”, is widely distributed in Eurasia and North Africa, but the highest taxa diversity is in Anatolia, the southern Balkans, the Middle East, the Caucasus and northwestern Iran and is represented by approximately 360 species in the World. As hybridization is very dense in the genus Vesbascum, the high frequency of morphological variations has been intricated in species delimitation. In this study, pollen and capsule morphology of three hybrid Verbascum taxa [Verbascum x calcicolum Hub.-Mor., V. x kotschyoides Hub.-Mor., and V. x nusaybinense Hub.- Mor.] were investigated. They are hybrids wild distributed in the province of Mardin in Southeastern Anatolia. Pollen slides were prepared according to the Woodhouse method, pollen and seed morphologies were examined with microscope. In general terms, aperture types were tricolporate, but also tricolpate aperture types were observed. The shapes of the pollen grains were determined as oblate-spheroidal and subprolate. The sculpture was reticulate in all cases. The seeds not developed. The capsules are pubescent, densely covered by branched hairs, or loosely by glandular hairs, depending on species. The results of this study showed that morphological features, such as pollen shape, polar axis length (P), equatorial diameter (E), aperture type, exine and intine thickness, colpus, and por measurement, show differences that can be taken into consideration in the systematic discrimination within the taxa. The capsule showed that substantial taxonomic insight can be gained from examining the capsule characteristics of Verbascum, particularly at the species level.

Natural asphalt on Late Neolithic (5000–4500 BCE) potsherds from southeastern Albania: A geochemical study

Nine archaeological bituminous potsherds from the Neolithic of Maliq and Kamnik (5000–4500 BCE) in Albania were geochemically compared for the first time to four natural asphalts from the famous present-day mine of Selenicë. The bitumen on potsherds originates from Selenicë, a unique source in Albania and the southern Balkans. Carbon isotopes on the asphaltene chromatographic fraction confirm that the bitumen on the potsherds came from Selenicë. Carbon and hydrogen isotopes on saturates, aromatics, NSO, and asphaltenes show changes that may be due to biodegradation-oxidation effects and also the occurrence of organic remains from the content of the vessels mixed with bitumens. Steranes, terpanes, and aromatics confirm the increased biodegradation of bitumen on potsherds when compared to natural asphalts references.

Early Neolithic pastoral land use at Alsónyék-Bátaszék, Hungary (Starčevo culture): New insights from stable isotope ratios

The earliest introduction of livestock (cattle, goats, sheep, pigs) into the Carpathian Basin was an important step towards farming expansion into continental Europe. This spread beyond the environments of the southern Balkans was accompanied by a reduction in the spectrum of cultivated crops, changes in the relative representation of different domestic animals, and, most likely, adaptations of husbandry practices. How the earliest farmers in the Carpathian Basin kept their domestic stock is still understudied. We explored early animal management and land use strategies at the Starčevo settlement at Alsónyék-Bátaszék, Hungary (Early Neolithic, ca. 5800–5600 cal BC). Settled at the intersection of wide alluvial plains, waterlogged meadows and marshes to the east, and forested hills to the west, early farmers at Alsónyék had a wide variety of options for nourishing their livestock. We performed stable isotope ratio analysis of bone collagen (n = 99; δ13C, δ15N) and tooth enamel (nteeth = 28, sequentially sampled for δ13C and δ18O) from wild and domestic animals to locate them in the landscape and investigate herding practices on a seasonal scale. The bone collagen isotope ratios mostly indicate feeding in open environments. However, results from the sequential analysis of cattle and sheep enamel suggest diverse dietary strategies for winters, including consumption of forest resources, consumption of summer hay and grazing in an open environment. Most pigs appear to have had herbivorous diets, but several individuals likely supplemented their diet with animal protein. Stable isotope ratio results from the Lengyel phase at Alsónyék (ca. 4800–4300 cal BC) suggest more access to animal protein for pigs, and feeding in more open areas by wild boar, red deer and cattle compared to the Starčevo phase. This study’s results demonstrate considerable variability in early animal husbandry practices at Alsónyék.

Allium goumenissanum (Allium sect. Codonoprasum), a new species for Bulgaria and new localities in Greece: testing an integrated approach to species identification

Allium sect. Codonoprasum represents an evolutionarily young and rapidly radiating group of bulbous geophytes, with a significant proportion of polyploids and minor morphological differences between species. In the last 20 years, dozens of new species of this section have been described from the Mediterranean. Still, very little new information has been obtained on most of them since their description, especially concerning cytogenetic, molecular, and distribution data. Allium goumenissanum is a recently described species of this section, known only from three nearby localities in northern Greece. During the last 10 years, the senior authors' team collected population samples of an unknown species of this section from the southern Balkans, which appeared to be representative of A. goumenissanum. Using an integrated approach employing a range of methods (cytogenic, molecular, morpho- and micromorphological methods), the populations of an unknown species were compared with morphologically similar species A. goumenissanum and A. stamatiadae. AFLP, ITS sequencing and genome size analyses revealed that all the populations of the unknown species represent A. goumenissanum, and that this species is differentiated genetically and in genome size from A. stamatiadae. ITS sequencing indicates an allotriploid origin of A. goumenissanum from unknown ancestors. The characterization of the morphology and reproduction of A. goumenissanum is completed and micromorphological and anatomical characteristics of the leaves of both species are newly published. The distribution of A. goumenissanum, based on new data, now includes a more extensive area of north central Greece and the species has been found as a new one for Bulgaria.

The Spatial Distribution of Crimean-Congo Haemorrhagic Fever and Its Potential Vectors in Europe and Beyond.

Crimean-Congo haemorrhagic fever (CCHF) is considered to be spreading across the globe, with many countries reporting new human CCHF cases in recent decades including Georgia, Türkiye, Albania, and, most recently, Spain. We update a human CCHF distribution map produced in 2015 to include global disease occurrence records to June 2022, and we include the recent records for Europe. The predicted distributions are based on long-established spatial modelling methods and are extended to include all European countries and the surrounding areas. The map produced shows the environmental suitability for the disease, taking into account the distribution of the most important known and potential tick vectors Hyalomma marginatum and Hyalomma lusitanicum, without which the disease cannot occur. This limits the disease's predicted distribution to the Iberian Peninsula, the Mediterranean seaboard, along with Türkiye and the Caucasus, with a more patchy suitability predicted for inland Greece, the southern Balkans, and extending north to north-west France and central Europe. These updated CCHF maps can be used to identify the areas with the highest probability of disease and to therefore target areas where mitigation measures should currently be focused.

Eagle owls’ predation within a highly diversified Late Glacial landscape: remains of pikas and hares (Lagomorpha) from the Loutra Almopias Cave (Central Macedonia, Greece)

During cave bear excavations in the Loutra Almopias Cave, Pella District, in Central Macedonia, Greece, an ecologically highly diverse Late Glacial faunal assemblage, mainly micromammals and birds, was sampled from an elevated small niche labelled LAC Ia. The lagomorphs are represented by steppe pika (Ochotona pusilla), brown (or European) hare (Lepus europaeus) and mountain hare (Lepus timidus). The site evidences the first record of this last species in Greece and its southernmost evidence so far. The taphonomic patterns of the fossils refer to eagle owls (Bubo bubo) as an accumulating agent. This is congruent with the total LAC Ia species spectrum and the bone element abundance and destruction patterns. Radiocarbon dates of 12,350±40 14C yrs BP (VERA-5631) from a Lepus sp. bone and 16,427±96 14C yrs BP (VERA-7402) from a L. timidus premolar indicate the emplacement of the assemblage in the (very probably early phase of the) Late Glacial. The taphocoenosis reflects a high variety of habitats and biodiversity from periglacial alpine grassland to deciduous forests and Mediterranean open fore-land areas with scattered brush and thick scrub. The position of the eagle owl site is approximately 50 m inside the cave. Along the distance from the Late Pleistocene entrance, the nocturnal raptors had to manage two direction changes within their flight path. That opening was closed later by talus sediments, and thus the fossil owl nest or feeding place has been sealed. The LAC Ia assemblage is so far a unique terrestrial record of Late Glacial biota in the Southern Balkans.

First records of Sirococcus conigenus causing shoot blight on Pinus peuce in Bulgaria

Macedonian pine (Pinus peuce) is a southern Balkan Peninsula endemic, growing in high mountains of Bulgaria, between 1400 and 2100 m a.s.l. Stands of P. peuce form the upper tree line forest areas. During a survey in 2020-2022 on Macedonian pine natural stands and plantations in Bulgaria, symptoms of shoot blight were observed in the Pirin Mts, the Rila Mts and Mt Vitosha. The fungal pathogen Sirococcus conigenus was identified as the causal agent of the disease that appeared for the first time on Pinus peuce in Bulgaria and Balkan Peninsula. Incidence of blighted shoots on individual trees varied, but was as high as 70-80% in the Rila Mts and Mt Vitosha.

Exploring macroinvertebrate biodiversity in the dynamic southern Balkan stream network of the Vjosa using preservative-based DNA metabarcoding

Reliable biodiversity data are crucial for environmental research and management. Unfortunately, data paucity prevails for many regions and organismal groups such as aquatic invertebrates. High-throughput DNA-based identification, in particular DNA metabarcoding, has accelerated biodiversity data generation. However, in the process of metabarcoding, specimens are usually destroyed, precluding later specimen-based analyses. Metabarcoding of DNA released into the preservative ethanol has been proposed as a non-destructive alternative, but proof-of-principle studies have yielded ambiguous results, reporting variance in detection probability for various taxa and methodological biases. In this study, we tested the performance of preservative-based metabarcoding of aquatic invertebrates in comparison to a standard morpho-taxonomic assessment based on samples from one of Europe’s last free-flowing rivers, the Vjosa. Multi-habitat samples were collected at 43 sites in two seasons and stored in ethanol, after fixation in formaldehyde for morpho-taxonomic analyses. Preservative-based DNA metabarcoding detected three times more taxa but failed to detect other taxa found using the standard method. In addition to incomplete reference data and primer bias that likely precluded the detection of specific taxa, preservative-based DNA metabarcoding cannot provide accurate abundance estimates. However, the metabarcoding data revealed distinct small-scale and large-scale community patterns in the Vjosa river network, which were also recovered by quantitative data of the standard approach. Overall, our results indicate that preservative-based metabarcoding provides important biodiversity data, which could be further improved by quantitative validation. The method is robust and reliable, even though samples were taken under harsh field-conditions and stored without cooling. Further, our results emphasise the need for reliable DNA barcoding reference libraries. Building those may be supported by preservative-based metabarcoding that maintains intact vouchers for subsequent specimen-based analyses.

Algorithmic probing: Prompting offensive Google results and their moderation

Google results have been scrutinized over the years for what they privilege, be it the surface web, the powerful, optimized webpages, the personalized and/or their own properties. For some time now, another type of Google returns also has been the source of attention: the offensive result. The following revisits a selection of offensive and other problematic results found by journalists and researchers alike. In a technique termed ‘algorithmic probing’, the prompting queries are re-run to study what has come of these results in Google Web and Image Search but mainly in Google Autocompletion. The question concerns a different kind of privileging – Google's hierarchy of concerns – or the extent to which certain categories as well as languages are moderated and others less so. In all, it was found that Google heavily moderates religion, ethnicities and sexualities (albeit with gaps) but leaves alone stereotypes of gendered professions as well as ageism. It also moderates to a greater degree in English compared to southern European and Balkan languages. The article concludes with a discussion of the stakes of Google's moderation, including its uneven coverage.

Phylogenomics of Anguis and Pseudopus (Squamata, Anguidae) indicates Balkan-Apennine mitochondrial capture associated with the Messinian event

A dated phylogenetic hypothesis on the evolutionary history of the extant taxa of the Western Palearctic lizards Anguis and Pseudopus is revised using genome-wide nuclear DNA and mitogenomes. We found overall concordance between nuclear and mitochondrial DNA phylogenies, with one significant exception – the Apennine A. veronensis. In mitochondrial DNA, this species forms a common clade with the earliest diverging lineage, the southern Balkan endemic A. cephallonica, while it clusters together with A. fragilis in nuclear DNA. The nuclear phylogeny conforms to the morphology, which is relatively similar between A. veronensis and A. fragilis. The most plausible explanation for the mitonuclear discordance is ancient mitochondrial capture from the Balkan ancestor of A. cephallonica to the Apennine population of the A. fragilis-veronensis ancestor. We hypothesize that this capture occurred only in a geographically restricted population. The dating of this presumed mitochondrial introgression and capture coincides with the Messinian event, when the Balkan and Apennine Peninsulas were presumably largely connected. The dated nuclear phylogenomic reconstruction estimated the divergence of A. cephallonica around 12 Mya, while the sister clade representing the A. fragilis species complex consisting of the sister species A. fragilis–A. veronensis and A. colchica–A. graeca further diversified around 7 Mya. The depth of nuclear divergence among the evolutionary lineages of Pseudopus (0.5–1.2 Mya) supports their subspecies status.

Ecological niche modeling of the Macedonian mouse, Mus macedonicus (Mammalia, Rodentia), under climate change conditions

Abstract Mus macedonicus Petrov & Ružic, 1983, also known as the Macedonian mouse or Balkan short-tailed mouse, lives in the southern Balkans and the Middle East. While this species is common in Mediterranean ecosystems and is listed as “least concern” by the IUCN, little is known about how its distribution may shift with climate change. This study explores the ‘species’ potential distribution in three different periods: during the Last Glacial Maximum (LGM) and under current and future climate scenarios, using Maximum Entropy modelling. Modelling was based on 137 georeferenced occurrence records from Macedonia, Bulgaria, Greece, Turkey, Syria, Lebanon, Israel, Iran, Georgia, and Russia and ten bioclimatic variables from the WorldClim database. We show that a combination of precipitation and temperature variables appear to shape the geographical range of the Macedonian mouse and that its predicted distribution during the LGM is consistent with its survival in multiple refugia, as suggested by previous genetic studies. Modelled future distributions are subtly but significantly different from the current, with population losses and gains in different regions. Our results provide a sound framework for future studies on this model species’ range dynamics, suggesting that the overall geographical range of M. macedonicus is relatively stable in the long term.

The fossil record of turtles from the Pleistocene of Crete (Greece)

The Isle of Crete contains an important endemic fossil fauna that has been studied extensively in the past. An endemic fossil tortoise, Testudo marginata cretensis Bachmayer, Brinkerink & Symeonidis, 1975, had previously been named from Pleistocene cave deposits in the island, presumably being larger than the mainland species and having some important morphological differences as well. In this paper I revise these type specimens and describe for the first time numerous additional fossil specimens from the Pleistocene sites of Gerani IV, Zourida, Koumpes, Liko, Mavro Mouri IV, and Simonelli Caves (near Herakleion), Rethymnon fissure, and from the open site of Sitia in the eastern part of the Island. These specimens belong mostly to the marginated tortoise, but the presence of a terrapin is confirmed as well. These new fossils help to refute the validity of the Cretan subspecies, and change the range of the marginated tortoise during Pleistocene/Holocene times in the southern Balkans.

Geographic differentiation in male calling song of Isophyamodestior (Orthoptera, Tettigoniidae, Phaneropterinae).

We studied the songs and morphology of the stridulatory file of Isophyamodestior across its complete geographic range, in order to test our hypothesis that the male calling song of the species shows strong differentiation between the northern (Pannonian) and southern (Balkan) parts of its distribution range, reflecting its disjunct distribution. Our analyses confirm this hypothesis, separating analyzed specimens of I.modestior into two main groups - one present in the central part of the Balkan Peninsula (representing Isophyamodestior sensu stricto), with the second group occurring in the Pannonian Basin, Dinarides, Slovenia and NE Italy. The most reliable difference between the groups is the duration of the main syllable, the number of stridulatory teeth and number of pulses in the main syllable, where all values are higher in specimens from the Balkan Peninsula. Additional analyses showed that within the second group, there are differences in analyzed characters between specimens from the Pannonian Basin and specimens from the Dinaric area, the latter ones having intermediate song characteristics, closer to the group from the Balkan Peninsula. Our study shows that detailed bioacoustic analyses can help to unravel patterns of intraspecific differentiation and thus provide a useful tool for taxonomic studies.

Reconstruction of historical forest cover on a 1° grid in central and southeast Europe from AD 1800 to 2000

The natural vegetation has been substantially changed over the last millenium, especially from forest to agricultural areas in Europe. To study the effects of deforestation on carbon and climate system, it is essential to construct spatially precise maps in forest cover. The current representative historical forest data sets are either derivatives of agricultural datasets or are produced based on the deforestation-population relationship. Almost no attempts have been made to use the inherently linked relationship between deforestation and cropland expansion that has run through Europe for millennia. This study created an approach based on the relationship between cultivation intensity and forest fraction to reconstruct forest cover changes in central and southeast Europe from AD 1800 to 2000 by the following steps: (1) To develop the spatial relationship curve (spatial resolution is 1°) between the “cultivation intensity (0–100%)” and the “forest fraction (0–100%)” derived from modern land cover datasets; (2) To generate the national-scale cropland data based on historical records from AD 1800 to 2000, and to allocate them to a 1° pixel; (3) To reconstruct the forest cover using the aforementioned relationship curve and the historical gridded cropland database. The results show that: (1) The forest fraction in central and southeast Europe decreased from 38.4% in AD 1800 to 27.0% in AD 1900, and then increased to 32.5% in AD 2000. (2) In AD 1800, large areas of forests can be primarily found in Luxembourg, Netherlands and Belgium, the south of France, the south of Germany, the Alps region, Poland, and the majority of the Balkans. (3) Throughout the 19th century, the entire region was generally subjected to deforestation, with large areas of forest coverage only being preserved in the Alps and the western and southern Balkans after 100 years of exploration. (4) The forest coverage in most of the study regions increased again at the end of the 20th century.

New insights into the genetic diversity of the Balkan bush-crickets of the Poecilimon ornatus group (Orthoptera: Tettigoniidae)

The Balkan Peninsula is treated as a hotspot of biodiversity with over 40% of European bush-crickets occurring there.PoecilimonFischer, 1853 is one of the largest Palaearctic orthopteran genera containing several species groups. One of them is thePoecilimon ornatusgroup (Schmidt, 1850) with 13 species and 5 subspecies. Among the group, thePoecilimon affiniscomplex is designated as consisting ofP. pseudornatusIngrisch & Pavićević, 2010,P. nonveilleriIngrisch & Pavićević, 2010, and five subspecies ofP. affinis(Frivaldszky, 1868). The aim of this study is to reconstruct the phylogenetic relationships among taxa of theP. ornatusgroup and to elucidate the position of taxa related to theP. affiniscomplex. Molecular phylogeny supported the monophyly of theP. ornatusgroup and showed that their ancestor probably originated in the southern Balkans. The underlying processes are thought to be six dispersals and five vicariance events linked to geological events and climate changes in the Pleistocene. The species delimitation analysis showed mostly nine hypothetical species among the group.