Dinarides Research Articles

Geomorphology of Blidinje, Dinaric Alps (Bosnia and Herzegovina)

ABSTRACTBlidinje is tectonically derived lowland in the Dinaric Alps within Bosnia and Herzegovina. It is surrounded by Vran, Čvrsnica and Čabulja Mountains. The Blidinje study area is predominantly built of carbonates, where karst, fluvial and glacial geomorphological systems interact. We present a 1:25,000 geomorphological map covering a 100 km2 with an elevation range of 1200–1900 m. The map results from a combination of a fieldwork mapping, interpretation of orthophotos and an analysis of a digital elevation model. This map presents a distribution of landforms and sediments associated with the past and present karst, glacial and fluvial processes, such as different varieties of karst depressions, conical hills, erosional gullies, alluvial fans and large amphitheatre-like moraines. The focus of the presented map is to assist ongoing studies in this part of the Dinaric Alps that seek to understand the dynamics of former glaciers and associated palaeoclimate.

The genetic impact of chamois management in the dinarides

The Dinaric Mountains in Slovenia, Croatia, and Bosnia and Herzegovina provide a unique system to address the effects of past hunting on the genetic structure of northern chamois (Rupicapra rupicapra) and possible hybridization in the contact zone in the Velebit Mountains. The northern Dinaric Mountains should be occupied by alpine chamois (Rupicapra rupicapra rupicapra), whereas the central and southern areas are inhabited by the Balkan chamois (R. rupicapra balcanica). This is the first study to characterize the genetic variation in chamois populations in the area. We used microsatellite and mitochondrial markers to analyze the genetic variation and structure of chamois populations from different geographical areas with different histories. Specifically, we explored the influence of recent human translocations and geographical isolation on the genetic architecture of chamois populations in the assumed contact zone. We successfully genotyped 74 individual samples and the number of alleles/locus ranged from 6 to 20 with a mean of 9.20. Allelic richness across populations ranged from 2.94 in the Prenj Mountains, Bosnia and Herzegovina to 3.56 in the Biokovo Mountains, Croatia. A similar pattern was also observed for heterozygosity, ranging between 0.729 and 0.572, and expected heterozygosity, ranging between 0.762 and 0.644 in the Prenj and Biokovo mountains, respectively. The global genetic distance (FST) for 7 population samples was 0.103 ± 0.047 (range = 0.0156–0.185). The STRUCTURE tree clusters separated samples from the northern Dinaric Mountains from those of the southern Dinaric Mountains into 2 clusters according to geographic location. The results obtained using a Bayesian clustering methodology was similar. By using mtDNA variation in chamois from Slovenia, Croatia, and Bosnia and Herzegovina, the existence of alpine chamois haplotypes in northern areas and Balkan chamois haplotypes in southern areas was confirmed. These results confirm the impact of recent human management (i.e., translocation) into the Velebit Mountains, which established a new contact (hybridization) zone between the subspecies. Therefore, future translocations must be planned carefully to avoid compromising genetic integrity and posing a serious risk to native species, as in this case. © 2016 The Wildlife Society.

The extent of middle Pleistocene ice cap in the coastal Dinaric Mountains of Croatia

Solitary limestone blocks and groups of blocks occur on Risnjak and Velebit Mountains and on the northern Adriatic islands of Krk and Rab. Previous researchers have interpreted some of these as a) erratic blocks, b) corrosional remnants, or c) rockfalls. We have studied their mode of occurrence and composition, and revised previous interpretations of their origin in the light of transport mechanism and depositional processes. After analyzing the context of the block positions and the physical processes responsible for their emplacement, and taking into account their sedimentological context (their association with glaciogenic sediments), we herein propose a glacial origin for most of these blocks. However, some blocks are indeed shaped by sub-soil corrosion, as evidenced by their structure. The interpreted erratic blocks on the inner northern Adriatic Sea islands document the presence of middle Pleistocene glaciation of Dinaric Mountains though not its maximal extent, which is still unclear as the ice terminus was in the area that is inundated by postglacial rise of Adriatic Sea. The reconstructed ice cap area, which extended along the coastal mountains from Risnjak Mt. to south Velebit Mt. and across the range from Lika Polje to Rab Island, is conservatively estimated to be 5400 km2.

Evolution of a fold-thrust belt deforming a unit with pre-existing linear asperities: Insights from analog models

Heterogeneity, whether geometric or rheologic, in crustal material undergoing compression affects the geometry of the structures produced. This study documents the thrust fault geometries produced when discrete linear asperities are introduced into an analog model, scaled to represent bulk upper crustal properties, and compressed. Varying obliquities of the asperities are used, relative to the imposed compression, and the resultant development of thrust fault traces and branch lines in map view is tracked. Once the model runs are completed, cross-sections are created and analyzed. The models show that asperities confined to the base layer promote the clustering of branch lines in the surface thrusts. Strong clustering in branch lines is also noted where several asperities are in close proximity or cross. Slight reverse-sense reactivation of asperities cut through the sedimentary sequence is noted in cross-section, where the asperity and the subsequent thrust belt interact. The model results are comparable to the situation in the Dinaric Alps, where pre-existing faults to the SW of the NE Adriatic Fault Zone contribute to the clustering of branch lines developed in the surface fold-thrust belt. These results can therefore be used to evaluate the evolution of other basement-involved fold-thrust belts worldwide.

Factors affecting the distribution and abundance of orchids in grasslands and herbaceous wetlands

Grasslands, wet meadows, fens, bogs, and marshes represent important ecosystems that host many orchid species. However, research on the joint effect of the multi-scalar abiotic drivers on orchid distribution and abundance in these habitats is limited. In this study, we investigated factors associated with the occurrence and abundance of 42 orchid taxa in grasslands and herbaceous wetlands of 29 mountains and the surrounding low altitude areas in western Serbia (eastern Dinaric Alps, southeastern Europe). Niche analysis of orchids was performed using outlying mean index analysis (OMI). In addition, we used similarity profile analysis (SIMPROF) to classify orchid taxa into ecological groups, multi response permutation procedure (MRPP) to analyse the differences in orchid composition between vegetation types, and indicator species analysis (ISA) to detect orchids with an affinity to a certain type of vegetation and geological bedrock. The most important gradients that separate orchids within the sampled environmental conditions were moisture, temperature, altitude and soil pH. Moreover, this study underlines the importance of certain vegetation types and geological bedrocks in determining the distribution and abundance of orchids. In total, eight ecological groups of orchids were distinguished. The results have shown that 11 orchid taxa are indicators of specific vegetation types, whereas 13 orchid taxa are indicators of specific bedrock types. The study area provides suitable conditions for survival of some rare and endemic orchid species that have southern limits of their distribution in this part of southeastern Europe. The study suggests that the joint effect of ecological factors, especially vegetation communities and bedrock types, is important to consider in the planning and management of orchid species conservation.

Diversification of Luzula sect. Luzula (Juncaceae) on the Balkan Peninsula – a cytogenetic approach

Luzula sect. Luzula is one of the taxonomically most intricate groups of angiosperms, where diversification is mostly driven by true polyploidy and agmatoploidy (fission of chromosomes) leading to three different karyotypes (whole chromosomes – AL, half-sized chromosomes – BL and quarter-sized chromosomes – CL). Morphologically, the taxa are often difficult to distinguish, and genome size data coupled with karyological investigations are currently the most reliable means for their identification. For the Balkan Peninsula, one of the botanically least studied areas of Europe, ten taxa from this section have been reported, but little is known about their distribution. We here estimated the nuclear DNA amount (genome size, GS) using interphase-peak DNA image cytometry and counted the chromosomes for 33 populations of five species of Luzula sect. Luzula mostly from mountainous areas of the Balkan Peninsula and the adjacent southeastern Alps. Five taxa have been confirmed, of which L. divulgatiformis is new for Croatia and L. exspectata for Macedonia and Serbia. The most common species in the Dinaric mountains appears to be diploid (2n = 12 AL) L. taurica, for which the GS (2C = 0.83 pg DNA, fitting well in the range of previously determined values of diploid taxa) has been determined for the first time. The GS and karyotypes of other taxa (L. campestris, L. divulgatiformis, L. exspectata, L. multiflora subsp. multiflora) correspond well to previously published values. An identification key for the Balkan taxa of Luzula sect. Luzula is also included.

Mesolithic harpoons from Odmut, Montenegro: Chronological, contextual, and techno-functional analyses

The (re-)appearance of harpoon technology during the Mesolithic in the southern Dinaric Alps is discussed by presenting the results of contextual, technological and use-wear analyses on the sample of 36 osseous harpoon specimens recovered in Mesolithic and Early Neolithic levels of the rockshelter of Odmut in western Montenegro. The assemblage of harpoons from this site is unique in Balkan prehistory, as there is currently limited evidence for the use of barbed weaponry prior to the regional Late Mesolithic. Technological and functional data related to the introduction of harpoon technology at Odmut and the southern Dinaric Alps are discussed against the background of the appearance of harpoons in the Alpine regions of Switzerland, Italy, and the northern Adriatic. Reasons and mechanisms that might have fostered the spread of harpoons in the circum-Adriatic region and a late use of harpoon technology in Montenegro are discussed.

Mortality in the Eurasian lynx population in Croatia over the course of 40 years

The combination of intensive persecution, habitat loss and prey deficiency led to the extinction of Eurasian lynx (Lynx lynx) in the Dinaric Mountains at the beginning of the 20th century. In 1973, the population was re-established by reintroducing animals from the Slovakian Carpathian Mountains into Slovenia, from where the animals spread into Croatia and Bosnia and Herzegovina. Since the end of the 20th century the reintroduced population has been decreasing, leading us to investigate the principal causes of mortality. Understanding the causes of the declining lynx population in Croatia and elsewhere is important not only in its own right but also because few studies have examined how large carnivore populations have fared under different management regimes. After reintroduction, the first dead lynx was recorded in Croatia in 1978, and from that year until 2013 a total of 232 deaths were recorded. Annual mortality during this period averaged 6.4 deaths per year, ranging from 0 to 17. The number of dead lynxes found every year in the period before the legal protection (1978–1998) was significantly higher than the number of dead animals found in the period after the legal protection (1998–2013). The vast majority of deaths were human-related (92.7%), with only 2.1% not human-related; while the cause of the remaining 5.2% of deaths was undetermined. Shooting was the most dominant cause of death (73.7%), with significantly more males being killed by shooting than females, and significantly more animals shot before the legal protection. Between 1978 and 1998, the year when the lynx became legally protected in Croatia, 10 deaths due to poaching were recorded, accounting for 5% of all deaths in that period and an average of 0.48 poaching cases per year. During 1999–2013, 18 poaching deaths occurred, accounting for 60% of all deaths in that period and an average of 1.2 poaching cases per year. Our findings suggest that the synergy of human-induced mortality, concomitant reduction in genetic variation and possibly prey deficiency may be the principal factors behind the decline in the reintroduced lynx population in Croatia since the end of the 20th century.

Are the dinaric mountains a boundary between continental and mediterranean wild boar populations in Croatia?

Wild boar is the most important large game species in Croatia, with populations subjected to hunting/management programs. There are indications from previous analyses of morphological variation and reproductive performance that population structure of wild boar in Croatia is strongly influenced by the Dinaric mountain range that separates Croatia in two biogeographic regions: Continental and Mediterranean. Of the two main wild boar haplogroups found in Europe, the most widespread is the E1 clade (present from Portugal to Russia), while the E2 clade is found within the Italian peninsula and on Sardinia. Outside Italy, the E2 clade has to date been found only in archaeological samples from the Mediterranean region of Croatia. The aims of this study were to investigate the hypotheses that (i) the Dinaric Mountains act as a natural boundary separating the Continental and Mediterranean wild boar subpopulations in Croatia; and (ii) the mtDNA E2 clade is present in Croatian wild boar. A total of 14 unlinked microsatellite loci were examined in 264 geo-referenced individuals, and the 486-bp fragment of the mtDNA control region sequence was examined in 69 individuals. The results suggest that the region of the Dinaric Mountains. represents a weak boundary between the Continental and Mediterranean subpopulations, and that the predominant factor causing the observed genetic structuring is isolation by distance. Based on these results, it can be proposed that the Croatian wild boar population in the mainland should be managed as a whole. Three variable sites and four haplotypes were obtained from the mtDNA sequences, all of which belonged to the major European clade (E1).

Evolution of a karst polje influenced by glaciation: The Gomance piedmont polje (northern Dinaric Alps)

Gomance is a piedmont karst polje in the northern Dinaric Alps presenting geomorphological and sedimentological evidence of past glaciation. During the Pleistocene the polje was situated at the edge of the Snežnik and Gorski Kotar ice fields from where two outlet glaciers reached Gomance. The morphogenesis of the polje was reconstructed by means of geomorphological mapping, sedimentological studies, and ground penetrating radar (GPR) measurements, supported by hand-drillings. With GPR an almost entirely buried moraine system was also imaged and mapped, crucial in reconstructing the polje history. The depression was karstified and well drained without any surface streams before the Last Glaciation. When the glacier front reached the depression, the entire floor became covered by glacial and outwash deposits. Surface runoff dominated over karst drainage in a large part of the polje, particularly where distal outwash deposits with low effective porosity functioned as an aquitard. These deposits diverted surface drainage toward the lowest edge of the polje, which functioned as a ponor front along the entire length. The outwash system of the Gomance polje was active during the Last Glaciation as suggested by radiocarbon-dated outwash deposits.

Variability in growth of trees in uneven-aged stands displays the need for optimizing diversified harvest diameters

This study presents economically optimal management of uneven-aged mixed mountain forests that takes into account tree growth variability. We divided 9846 silver fir (Abies alba), beech (Fagus sylvatica), and spruce (Picea abies) trees measured on 898 forest inventory plots in the Snežnik and Leskova dolina management units (4905 ha, Dinaric mountains, Slovenia) into three growth classes (slow-, medium-, and fast-growing trees) to simulate optimal forest management over a period of 100 years with respect to changing tree growth, stand density, diameter distribution, and tree species composition. We developed a density-dependent and stage and growth-structured matrix transition model which—simultaneous to the long-term stand dynamics projection—scheduled optimal harvesting to maximize the net present value using a nonlinear approach. The ecology of tree species was considered by using tree species-specific and stand-density and diameter-dependent logistic functions for ingrowth, transition, and mortality. The model projected a shift in tree species composition from fir-dominated to beech-dominated forests within 100 years. A change from harvesting slow- and fast-growing trees as if they all had medium growth to growth-sensitive harvesting increased the net revenue and maintained the uneven-aged stand structure. Optimal harvest diameters varied among growth classes, time periods, and tree species according to the economic maturity of individual trees and ranged from 12 (pre-commercial thinning) to 72 cm (target diameter). The simulation highlights the potential of improved bio-economic models for increasing yield from uneven-aged forests and scheduling optimal management regimes with multiple objectives.

Accurate modeling of harvesting is key for projecting future forest dynamics: a case study in the Slovenian mountains

Maintaining the provision of multiple forest ecosystem services requires to take into consideration forest sensitivity and adaptability to a changing environment. In this context, dynamic models are indispensable to assess the combined effects of management and climate change on forest dynamics. We evaluated the importance of implementing different approaches for simulating forest management in the climate-sensitive gap model ForClim and compared its outputs with forest inventory data at multiple sites across the European Alps. The model was then used to study forest dynamics in representative silver fir–European beech stands in the Dinaric Mountains (Slovenia) under current management and different climate scenarios. On average, ForClim accurately predicted the development of basal area and stem numbers, but the type of harvesting algorithm used and the information for stand initialization are key elements that must be defined carefully. Empirical harvesting functions that rigorously impose the number and size of stems to remove fail to reproduce stand dynamics when growth is just slightly under- or overestimated, and thus should be substituted by analytical thinning algorithms that are based on stochastic distribution functions. Long-term simulations revealed that both management and climate change negatively impact conifer growth and regeneration. Under current climate, most of the simulated stands were dominated by European beech at the end of the simulation (i.e., 2150 AD), due to the decline of silver fir and Norway spruce caused mainly by harvesting. This trend was amplified under climate change as growth of European beech was favored by higher temperatures, in contrast to drought-induced growth reductions in both conifers. This forest development scenario is highly undesired by local managers who aim at preserving conifers with high economic value. Overall, our results suggest that maintaining a considerable share of conifers in these forests may not be feasible under climate change, especially at lower elevations where foresters should consider alternative management strategies.

European wildcat populations are subdivided into five main biogeographic groups: consequences of Pleistocene climate changes or recent anthropogenic fragmentation?

Extant populations of the European wildcat are fragmented across the continent, the likely consequence of recent extirpations due to habitat loss and over-hunting. However, their underlying phylogeographic history has never been reconstructed. For testing the hypothesis that the European wildcat survived the Ice Age fragmented in Mediterranean refuges, we assayed the genetic variation at 31 microsatellites in 668 presumptive European wildcats sampled in 15 European countries. Moreover, to evaluate the extent of subspecies/population divergence and identify eventual wild×domestic cat hybrids, we genotyped 26 African wildcats from Sardinia and North Africa and 294 random-bred domestic cats. Results of multivariate analyses and Bayesian clustering confirmed that the European wild and the domestic cats (plus the African wildcats) belong to two well-differentiated clusters (average Ф ST=0.159, r st=0.392, P>0.001; Analysis of molecular variance [AMOVA]). We identified from c. 5% to 10% cryptic hybrids in southern and central European populations. In contrast, wild-living cats in Hungary and Scotland showed deep signatures of genetic admixture and introgression with domestic cats. The European wildcats are subdivided into five main genetic clusters (average Ф ST=0.103, r st=0.143, P>0.001; AMOVA) corresponding to five biogeographic groups, respectively, distributed in the Iberian Peninsula, central Europe, central Germany, Italian Peninsula and the island of Sicily, and in north-eastern Italy and northern Balkan regions (Dinaric Alps). Approximate Bayesian Computation simulations supported late Pleistocene-early Holocene population splittings (from c. 60k to 10kyears ago), contemporary to the last Ice Age climatic changes. These results provide evidences for wildcat Mediterranean refuges in southwestern Europe, but the evolution history of eastern wildcat populations remains to be clarified. Historical genetic subdivisions suggest conservation strategies aimed at enhancing gene flow through the restoration of ecological corridors within each biogeographic units. Concomitantly, the risk of hybridization with free-ranging domestic cats along corridor edges should be carefully monitored.

Phytosociological Description of Ostrya Carpinifolia and Fraxinus Ornus Communities in the Julian Alps and in the Northern Part of the Dinaric Alps (NW and W Slovenia, NE Italy)

Abstract Based on comparisons between similar communities in northeastern Italy, southern Austria, central and southeastern Slovenia, and western Croatia, we classified black hornbeam and flowering ash phytocoenoses on steep rocky sites in the beech forest belt in northwestern and western Slovenia into the association Fraxinio orni-Ostryetum Aichinger 1933 and described its new subassociation -phyteumatetosum columnae in the foothills of the Julian Alps and in the northern part of the Dinaric Alps. Black hornbeam and flowering ash stands on steep shady slopes with a higher proportion of diagnostic species of beech and spruce forests are classified into the new association Rhododendro hirsuti-Ostryetum Franz ex Dakskobler, ass. nov. hoc loco, new subassociation -mercurialietosum perennis and the provisional variant var. Hemerocallis lilioasphodelus whose stands are floristically rather similar to the stands of the association Hemerocallido-Ostryetum.

Phytosociological description of hay meadows with dominating Trisetum flavescens in the lower montane belt of north-western and western Slovenia

We conducted a phytosociological study into hay meadows on former fields on original sites of beech forests form the alliance Aremonio-Fagion in the lower montane belt of the northwestern and western Slovenia (southern Julian Alps, northern part of the Dinaric Alps) and compared them to similar, previously described meadows in Slovenia and northwestern Italy. Based on this comparison they are classified into the new association Rhinantho freynii-Trisetetum flavescentis and new habitat type, southeastern-Alpine-northern-Illyrian lower montane hay meadows – 38.239-S1.

Impacts of Climate Change on the Hydrological Regime of the Danube River and Its Tributaries Using an Ensemble of Climate Scenarios

Information about the potential impacts of climate change on river runoff is needed to prepare efficient adaptation strategies. This study presents scenario projections for the future hydrological runoff regime in the Danube River Basin. The eco-hydrological watershed model Soil and Water Integrated Model (SWIM) was applied for the entire Danube River catchment, considering 1224 subbasins. After calibration and validation of the model, a set of high-resolution climate projections (bias-corrected and non-bias-corrected) served as meteorological drivers with which future daily river discharge under different climate warming scenario conditions was simulated. Despite existing uncertainties, robust trends could be identified. In the next 30 years, the seasonal stream-flow regime of the Danube and its tributaries is projected to change considerably. Our results show a general trend towards a decrease in summer runoff for the whole Danube basin and, additionally, in autumn runoff for the Middle and Lower Danube basin, aggravating the existing low flow periods. For the winter and early spring seasons, mainly January–March, an increase in river runoff is projected. Greater uncertainties show up in particular for winter runoff in the Dinaric Alps and the Lower Danube basin. The existing trends become very distinct until the end of the 21st century, especially for snow-influenced river regimes.

Pleistocene glaciation in Mediterranean Slovenia

Abstract Geomorphological research on glacial features in Slovenia has a long tradition; the first studies were published in the late nineteenth century by Brückner. Evidence of glacial erosion and deposition is preserved in the Slovenian Alps and on the highest plateaus of the Dinaric Mountains, including traces of ice fields and ice caps with outlet glaciers on the edges and some valley glaciers. The prevailing bedrock in all the glaciated areas is limestone, forming complex glacio-karst morphological systems. Modern glacial reconstruction techniques have been applied in the last few years, focusing on the interpretation of palaeoglacial topography and local equilibrium-line altitudes. Both past and present studies, however, lack precise chronological frameworks for the glacial advances, which should be the main goal of future research on glaciation in Slovenia.

Long-distance dispersal connects Dinaric-Balkan and Alpine grey wolf (Canis lupus) populations

In the last two centuries, persecution and deforestation caused grey wolf Canis lupus populations in Europe to decline. Recently, their numbers started to recover although most populations still remain isolated from one another. This study presents the first documented evidence of the successful reconnection of the Dinaric-Balkan and the Alpine wolf populations via long-distance dispersal and subsequent reproduction. A young male wolf radiocollared in the Dinaric Mountains in July 2011 travelled through Slovenia and Austria to the Italian Alps, where he settled in March 2012. During the 98 days of dispersal period, the wolf has travelled a cumulative line distance of 1176 km crossing multiple anthropogenic and natural barriers, and successfully hunting wild prey until he settled 233 km straight line distance from its natal territory. Camera trapping, snow tracking and genetic evidence in the new territory confirmed pairing with a female wolf from the neighbouring Alpine population. In the following year, the pair has produced a first documented “mixed” litter between wolves from the Dinaric-Balkan and the Alpine wolf populations. This case study demonstrates the potential for the future merging of European wolf populations even in human-dominated landscapes and highlights the importance of transboundary cooperation in wolf research and management.

Glaciokarst geomorphology of the Northern Dinaric Alps: Snežnik (Slovenia) and Gorski Kotar (Croatia)

ABSTRACTSnežnik and Gorski Kotar are mountainous areas in the Northern Dinaric Alps. We investigated this remote and densely forested area on the border between Slovenia and Croatia between 2011 and 2014. The result of the fieldwork campaign is a 1:30,000 geomorphological map covering almost 600 km2. The study area mainly consists of Mesozoic age limestone and dolostone. Karstified carbonate lithology results in almost the complete absence of surface run-off and a dominance of typical karst morphology. The extensive karstified area was modified by Quaternary glaciations; therefore, the recent landscape is characterized as a glaciokarst. The focus of the presented map is to provide a basis for further research of Quaternary glaciations in the areas of Snežnik and Gorski Kotar. Along with the map, we propose a legend for glaciokarst landscape mapping.

Flow cytometry, microsatellites and niche models reveal the origins and geographical structure of Alnus glutinosa populations in Europe.

Polyploidy in plants has been studied extensively. In many groups, two or more cytotypes represent separate biological entities with distinct distributions, histories and ecology. This study examines the distribution and origins of cytotypes of Alnus glutinosa in Europe, North Africa and western Asia. A combined approach was used involving flow cytometry and microsatellite analysis of 12 loci in 2200 plants from 209 populations combined with species distribution modelling using MIROC and CCSM climatic models, in order to analyse (1) ploidy and genetic variation, (2) the origin of tetraploid A. glutinosa, considering A. incana as a putative parent, and (3) past distributions of the species. The occurrence of tetraploid populations of A. glutinosa in Europe is determined for the first time. The distribution of tetraploids is far from random, forming two geographically well-delimited clusters located in the Iberian Peninsula and the Dinaric Alps. Based on microsatellite analysis, both tetraploid clusters are probably of autopolyploid origin, with no indication that A. incana was involved in their evolutionary history. A projection of the MIROC distribution model into the Last Glacial Maximum (LGM) showed that (1) populations occurring in the Iberian Peninsula and North Africa were probably interconnected during the LGM and (2) populations occurring in the Dinaric Alps did not exist throughout the last glacial periods, having retreated southwards into lowland areas of the Balkan Peninsula. Newly discovered tetraploid populations are situated in the putative main glacial refugia, and neither of them was likely to have been involved in the colonization of central and northern Europe after glacial withdrawal. This could mean that neither the Iberian Peninsula nor the western part of the Balkan Peninsula served as effective refugial areas for northward post-glacial expansion of A. glutinosa.