Dinarides Research Articles

Microclimate of Brown Bear (Ursus arctos L.) Dens and Denning Area

The aim of this study was to determine the microclimate of brown bear dens depending on their size, their status (active or inactive), and the location of the den. The study included five dens of different sizes and locations in the Velebit Mountains, in the Dinaric Alps. The measurements of air temperature (°C) and relative humidity (%) in dens and the forest stands were carried out over 182 days. The absolute minimum air temperature in inactive dens was between −2.88 °C and −0.38 °C and belonged to the dens with shorter tunnels and without chambers. The exception was inactive den 1, which was situated in a thermophilic forest. No negative absolute minimum air temperatures were recorded in active dens. The absolute minimum air temperatures in the forest stands were lower than in the dens. The relative humidity was significantly higher in all dens than in the forest stands. Dens with a large entrance opening and a short tunnel were strongly influenced by the local microclimate of the forest stand. The greatest temperature difference (den–forest) was between elongated dens with chambers and the associated forest stand, while the smallest temperature differences occurred in dens with a short tunnel, without a chamber, and at the northern exposure. The greatest differences in relative humidity (den–forest) were found in dens with chambers. The den temperatures correlated with the air temperatures in the forest stands.

Overview of mean and extreme precipitation climate changes across the Dinaric Alps in the latest EURO-CORDEX ensemble

Overview of mean and extreme precipitation climate changes across the Dinaric Alps in the latest EURO-CORDEX ensemble

Integrating geodiversity in animal spatial ecology: microhabitat selection of Eurasian lynx (Lynx lynx) and European wildcat (Felis silvestris) in a karst landscape

Geodiversity, encompassing various geophysical elements, can have an important impact on species distribution and affect animal behaviour patterns. Although many wild felids are attracted to rugged terrain and conspicuous relief features, most previous research was limited to general topographical characteristics (e.g., slope or terrain ruggedness) and rarely considered the effects of specific microhabitat characteristics. This gap is primarily due to the limited availability of high-resolution digital terrain models (DTMs) and relief features data at larger scales. However, LiDAR DTMs can be used in combination with various automatic methods to detect relief features, enabling non-contact and accurate mapping of large, remote and densely-forested areas. Here, we investigated the selection patterns of various karstic relief features, as well as topographic, anthropogenic and vegetation characteristics, by two sympatric felids, the Eurasian lynx (Lynx lynx) and the European wildcat (Felis silvestris), in the Dinaric Mountains, Slovenia. We used LiDAR DTM to calculate topographic characteristics and detect karst relief features based on automatic methods. We compared the selection of these features between the GPS-collared lynx and wildcats under a use-availability approach. We also investigated the differences in the selection of these features by lynx based on their origin and experience (remnant vs. translocated and naive vs. experienced, respectively). We observed significant impact of relief features on space use by both felids and detected distinct selection patterns between the two species. Lynx selected rugged terrain and proximity of caves, cliffs, karst depressions, ridges, small rocky outcrops, and roads, but avoided human settlements and forest edges. Wildcats selected areas with lower surface slope, closer to main roads, forest edges, caves and ridges, but avoided cliffs, forest roads and human settlements. We observed stronger selection/avoidance patterns among the translocated compared to the remnant lynx, while the differences in experience levels were less important. Our study demonstrates the potential of integrating remote sensing techniques and information on geodiversity into the study of animal spatial ecology. Furthermore, our results indicate that specific relief features provide important abiotic microhabitats for felids and may influence habitat segregation between sympatric species. Our findings provide further evidence for the importance of geodiversity conservation and the need to incorporate abiotic microhabitat features in wildlife habitat selection studies.

Holocene ecosystem and temperature development inferred from invertebrate remains in Zminje Jezero (Dinaric Alps, Montenegro)

AbstractLake Zminje Jezero (1535 m a.s.l.) in Montenegro was studied for chironomid and other aquatic invertebrate remains in a sediment sequence dating back to 12,000 calibrated 14C years before present (cal yr BP), providing, to our knowledge, the first lake-sediment record studied for chironomids and other associated chitinous aquatic invertebrate remains in the Dinaric Alps. Changes in chironomid and other invertebrate remains along the record make it possible to constrain changes of relevant environmental variables for aquatic invertebrates in the lake including temperature, oxygen availability, trophic status and water depth. The results suggest moderate changes in trophic conditions with chironomid assemblages indicating meso- to eutrophic conditions during the analysed interval. Invertebrate assemblages were typical for lakes with relatively high oxygen availability with a minor trend during the middle to late Holocene to conditions observed in lakes that are more hypoxic. A change in temperature is suggested in the earlier part of the record when the share of chironomid taxa adapted to warmer conditions increased, at the Younger Dryas to Holocene transition, whereas later no particularly pronounced shifts in temperature-sensitive taxa were observed. July air temperatures were estimated based on chironomid assemblages using a transfer function based on calibration data consisting of 117 lakes sampled in the Swiss Alps and northern Switzerland. The results suggest that temperatures rapidly increased by 5 °C at the onset of the Holocene (11,500 cal yr BP) leading to a relatively warm early to mid-Holocene and thereafter slightly decreased during the late Holocene. Reconstructed temperatures are discussed together with a previously published pollen record for our study site and are broadly consistent with other chironomid-based temperature reconstructions from Central, Eastern and Southern Europe. From ca. 3000 cal yr BP sedimentation rates increased and from ca. 500 cal yr BP onwards pollen data suggest that the vegetation and lake catchment were affected by human activities, possibly influencing chironomid and other invertebrate assemblages in the lake and thereby also reconstructed temperatures. Overall, our results show that combining analyses of chironomid and other invertebrate assemblages can provide valuable insights into long-term environmental changes and can provide temperature reconstructions for small mountain lakes in the Balkans, such as Zminje Jezero.

Flooding and flood water storage in karst systems of the Mediterranean region

Flooding is a recurring natural phenomenon that can have both life-giving and destructive aspects. In natural environments, floods are often an important element of the seasonal hydrologic cycle that provides water and nutrients to soil, supporting unique, rich and diverse ecosystems. However, flood events can also represent a destructive force that can endanger lives and cause significant damage in urban areas. Karst areas, in particular, are unique because of their special hydraulic characteristics in terms of flood occurrence, the dependence of ecosystems on such events, and attempts to actively store and manage floods. In this article, the hydraulic response of karst aquifers to heavy precipitation events, flood generation, and engineering interventions for flood control are discussed using several examples from karst areas in the Mediterranean region. Flooding mechanisms and regulatory structures in karst poljes are considered using several typical examples from the Dinaric mountain range. In addition, different variants of groundwater abstraction for increasing storage capacity and flood control are presented using examples from France and Montenegro. Managed aquifer recharge in karst areas and adjacent aquifers is demonstrated with examples from Jordan and Algeria. Finally, failed attempts at flood storage in karst reservoirs are presented with examples from Spain and Montenegro. These examples of flood retention in karst areas show the wide range of planning and technical measures and remind us of possible risks and failures in implementation as well as some positive and negative impacts on the environment and especially on ecosystems.

Leaf phenotypic plasticity of European ash (Fraxinus excelsior) at its northern range in Dinaric Alps

The Dinaric Alps have been recognised on numerous occasions as a biodiversity hotspot. They host a variety of species with great importance in sustainable forestry operations and nature conservation. One such species is the European ash (Fraxinus excelsior L., Oleaceae), a broad-leaved, wind-pollinated and wind-dispersed forest tree. In this paper, we aimed to determine the morphological variability of the European ash populations of the northern Dinaric Alps. For this purpose, leaf samples from 10 individuals in seven populations were collected. Morphometric analysis of intra- and interpopulation variability was conducted using 19 morphological leaf traits. We determined great variability of trees within, and small variability among, populations. The variables that best discriminated studied populations were those relating to leaflet shape. Based on these variables, populations were grouped into two clusters. The first cluster encompassed individuals with acute leaflets, found in drier and rockier habitats, whereas the second cluster, defined by more rounded leaflets, was found in mesophilous and nutrient-rich habitats. However, this research revealed no influence of geographical or bioclimatic distances on morphological variability, which indicates that the rockiness and soil are most likely two predominant factors in shaping the phenotypic plasticity of European ash populations. These results are of great significance in the planning of future forest breeding programs, as populations from drier habitats are likely to persist and spread due to their adaptation to water scarcity, which will become more pronounced in the future.

Simulation of a former ice field with Parallel Ice Sheet Model – Snežnik study case

Abstract. In this paper, we present a reconstruction of climate conditions during the Last Glacial Maximum on a karst plateau Snežnik, which lies in Dinaric Mountains (southern Slovenia) and bears evidence of glaciation. The reconstruction merges geomorphological ice limits, classified as either clear or unclear, and a computer modelling approach based on the Parallel Ice Sheet Model (PISM). Based on extensive numerical experiments where we studied the agreements between simulated and geomorphological ice extent, we propose using a combination of a high-resolution precipitation model that accounts for orographic precipitation combined with a simple elevation-based temperature model. The geomorphological ice extent can be simulated with climate to be around 6 °C colder than the modern day and with a lower-than-modern-day amount of precipitation, which matches other state-of-the art climate reconstructions for the era. The results indicate that an orographic precipitation model is essential for the accurate simulation of the study area, with moist southern winds from the nearby Adriatic Sea having a predominant effect on the precipitation patterns. Finally, this study shows that transforming climate conditions towards wetter and warmer or drier and colder does not significantly change the conditions for glacier formation.

The largest European forest carbon stocks are in the Dinaric Alps old-growth forests: comparison of direct measurements and standardised approaches

BackgroundCarbon (C) sink and stock are among the most important ecosystem services provided by forests in climate change mitigation policies. In this context, old-growth forests constitute an essential reference point for the development of close-to-nature silviculture, including C management techniques. Despite their small extent in Europe, temperate old-growth forests are assumed to be among the most prominent in terms of biomass and C stored. However, monitoring and reporting of C stocks is still poorly understood. To better understand the C stock amount and distribution in temperate old-growth forests, we estimated the C stock of two old-growth stands in the Dinaric Alps applying different assessment methods, including direct and indirect approaches (e.g., field measurements and allometric equations vs. IPCC standard methods). This paper presents the quantification and the distribution of C across the five main forest C pools (i.e., aboveground, belowground, deadwood, litter and soil) in the study areas and the differences between the applied methods.ResultsWe report a very prominent C stock in both study areas (507 Mg C ha− 1), concentrated in a few large trees (36% of C in 5% of trees). Moreover, we found significant differences in C stock estimation between direct and indirect methods. Indeed, the latter tended to underestimate or overestimate depending on the pool considered.ConclusionsComparison of our results with previous studies and data collected in European forests highlights the prominence of temperate forests, among which the Dinaric Alps old-growth forests are the largest. These findings provide an important benchmark for the development of future approaches to the management of the European temperate forests. However, further and deeper research on C stock and fluxes in old-growth stands is of prime importance to understand the potential and limits of the climate mitigation role of forests.

Response of beech and fir to different light intensities along the Carpathian and Dinaric Mountains.

Predicting global change mitigations based on environmental variables, like temperature and water availability, although yielding insightful hypothesis still lacks the integration of environmental responses. Physiological limits should be assessed to obtain a complete representation of a species' fundamental niche. Detailed ecophysiological studies on the response of trees along the latitudinal gradient are rare. They could shed light on the behaviour under different light intensities and other studied traits. The forests of the Dinaric Mountains and the Carpathians represent the largest contiguous forest complexes in south-eastern Europe. In uneven-aged Carpathian (8 plots) and Dinaric Mountain (11 plots) forests, net assimilation (Amax) and maximum quantum yield (Φ) were measured for beech and fir in three predefined light intensity categories according to the indirect site factor (ISF%) obtained by the analysis of hemispherical photographs in managed and old growth forests, all located above 800 m a.s.l. The measurements were carried out under fixed environmental conditions in each light category per plot for three consecutive years. Data from the last 50-year average period from the CRU TS 4.01 dataset were used for the comparison between Amax, Φ, and climate. The highest Φ for beech were observed in the central part of the Dinaric Mountains and in the south westernmost and northwesternmost part of the Carpathians for both beech and fir, while they were highest for fir in the Dinaric Mountains in the northwesternmost part of the study area. The Φ-value of beech decreased in both complexes with increasing mean annual temperature and was highest in the open landscape. For fir in the Carpathians, Φ decreased with increasing mean annual temperature, while in the Dinaric Mountains it increased with higher temperature and showed a more scattered response compared to the Carpathians. Short-term ecophysiological responses of beech and fir were consistent to long-term radial growth observations observed on same locations. The results may provide a basis and an indication of the future response of two tree species in their biogeographical range to climate change in terms of competitiveness, existence and consequently forest management decisions.

Old-growth forests in the Dinaric Alps of Bosnia-Herzegovina and Montenegro: a continental hot-spot for research and biodiversity

IntroductionAccording to various censuses, Europe has less than 1.5 million ha of old-growth forests (OGF). Most of them are in the boreal zone, while their presence in the temperate zone is residual and fragmented.In the framework of the EU biodiversity strategy, it has been adopted a broad definition of OGF which includes late-seral forests and forests with some management legacies. However, research purposes need to identify strictly defined OGFs characterized by structure, disturbance history, and processes typical and exclusive of the last stage of the forest dynamic.MethodsThe present paper wants to contribute to this debate by presenting a research network of four mixed (Fagus-Abies-Picea) montane OGFs in the Dinaric Alps (Lom, BiH; Janj, BiH; Perućica, BiH; Biogradska Gora, MNE), summarizing 20 years of multidisciplinary research by focusing on the structural characteristics and the disturbance history of the whole network and their coherency with strict OGF indicators. These sites were selected in relatively structurally uniform study areas, where 142 permanent plots have been established since 2002.Results and discussionThe study sites have a high living (747–1,201 m3 ha−1) and coarse woody debris (CWD) biomass (304–410 m3 ha−1), resulting in the highest forest carbon sink at the continental level (398–484 Mg C ha−1). The presence of large and old trees is one of the critical characteristics of the old-growth stage: in Lom and Perućica, there are 19 trees and 14 ha−1 larger than 1 m at breast height, respectively, and 14 trees and 15 trees ha−1 older than 400 years. In the last three centuries, continuous small-scale disturbances have driven forest dynamics, developing stands characterized by gap-phase dynamics and quasi-equilibrium structure. The Dinaric OGF network presents robust indicators of old-growthness, similar structural characteristics, and dynamic processes across all four sites. Identifying this sub-set of OGF using strict criteria is critical for recognizing conservation priorities and for quantifying, along an old-growthness chronosequence, the current structural differences of managed or recently abandoned forests. Besides, only OGF selected with rigorous criteria can act as a reliable reference for ecological restoration and sustainable forest management as a benchmark for carbon sink and for quantifying the impact of climate change on forests.

Molecular analysis of scats revealed diet and prey choice of grey wolves and Eurasian lynx in the contact zone between the Dinaric Mountains and the Alps

A comprehensive understanding of the dietary habits of carnivores is essential to get ecological insights into their role in the ecosystem, potential competition with other carnivorous species, and their effect on prey populations. Genetic analysis of non-invasive samples, such as scats, can supplement behavioural or microscopic diet investigations. The objective of this study was to employ DNA metabarcoding to accurately determine the prey species in grey wolf (Canis lupus) and Eurasian lynx (Lynx lynx) scat samples collected in the Julian Alps and the Dinaric Mountains, Slovenia. The primary prey of wolves were red deer (Cervus elaphus) (detected in 96% scat samples), European roe deer (Capreolus capreolus) (68%), and wild boar (Sus scrofa) (45%). A smaller portion of their diet consisted of mesocarnivores, small mammals, and domestic animals. In contrast, the lynx diet mostly consisted of European roe deer (82%) and red deer (64%). However, small mammals and domestic animals were also present in lynx diet, albeit to a lesser extent. Our findings indicate that the dietary habits of wolves and lynx are influenced by geographical location. Snapshot dietary analyses using metabarcoding are valuable for comprehending the behaviour and ecology of predators, and for devising conservation measures aimed at sustainable management of both their natural habitats and prey populations. However, to gain a more detailed understanding of wolf and lynx dietary habits and ecological impact, it would be essential to conduct long-term genetic monitoring of their diet.

<div>First record of saproxylic beetle Corticeus (= Hypophloeus) unicolor Piller & Mitterpacher, 1783 in Montenegro (Coleoptera: Tenebrionidae) with comments on old-growth forests conservation in the Country</div>

The occurrence of Darkling Beetle Corticeus (=Hypophloeus) unicolor Piller & Mitterpacher, 1783 (Coleoptera: Tenebrionidae), a saproxylic predator beetles, is reported for Montenegro. The species develops in freshly the dead wood of birch, beech, and oak trees. It is probably a predator of the larvae of the beetle Hylecoetus and other wood borers. C. unicolor was found in the Biogradska Gora National Park, in the north-western part of the Bjelasica mountain range in the Dinaric Alps (Montenegro), in October 2022 in an old-growth beech forest at about 1,000 m elevation.

A new subspecies of the subterranean beetle Speoplanes biocovensis Mller, 1934 (Coleoptera: Leiodidae: Cholevinae: Leptodirini) from Bosnia and Herzegovina: the first occurrence of the genus outside Croatia.

A new subterranean leptodirine leiodid beetle, Speoplanes biocovensis bosnicus Vrbica, Vesovi, Raa & uri, ssp. n., is described and diagnosed from a pit in southern Bosnia and Herzegovina. This is the first discovery of the genus Speoplanes Mller, 1911 in this country and the first discovery outside Croatia. Important morphological features of the new taxon are presented, including aspects of the male and female genitalia. The subspecies is endemic to the Dinaric mountain range of Bosnia and Herzegovina. A key to the identification of taxa of the genus Speoplanes is added.

Karyotype Variability in Wild Narcissus poeticus L. Populations from Different Environmental Conditions in the Dinaric Alps.

Narcissus poeticus L. (Amaryllidaceae), a facultative serpentinophyte, is a highly variable species and particularly important ancestor of cultivated daffodils, but is rarely studied in field populations. This study, based on natural populations in the Balkans, focused on karyotype variability, genome size, ploidy and the presence of B chromosomes. Thirteen native populations from different environmental and soil conditions were collected and analyzed using flow cytometry to estimate nuclear genome size, fluorescence in situ hybridization (FISH) for physical mapping of rDNA, fluorochrome labeling (chromomycin and Hoechst) for heterochromatin organization and silver nitrate staining of nucleoli for determining rRNA gene activity. The organization of rDNA and natural triploids is reported here for the first time. The presence of individuals with B chromosomes (in 9/13 populations) and chromosomal rearrangements was also detected. The observed B chromosome showed three different morphotypes. The most frequent submetacentric type showed four different patterns, mainly with active ribosomal genes. The results obtained show that N. poeticus has a dynamic genome with variable genome size due to the presence of polyploidy, B chromosomes and chromosomal rearrangements. It is hypothesized that the observed changes reflect the response of the genome to different environmental conditions, where individuals with B chromosomes appear to have certain adaptive advantages.

Ocra — an Etymological Proposal

The ancient name of the mountain (pass) that lies at the junction of the Julian and Dinaric Alps (nowadays Mt Nanos and the adjoining Razdrto Pass in southwestern Slovenia) is attested as Ocra in Latin (Pliny) and ῎Oκρα in Greek sources (Strabo, Ptolemy). The name is surely Italic and is argued to go back to Proto-Indo-European *H2ok̑reH2- (to *H2ek̑ - ‘to be/become/make sharp’). Given that an o-grade root is not expected in a deverbal adjective in -ro- (i.e. *H2ek̑-ró- ‘sharp’), such a formation is only interpretable as a substantivized feminine form of the possessive adjective *H2ok̑-r-ó- ‘sharp,’ derived from an acrostatic heteroclite neuter abstract noun *H2ók̑-r-/*H2ék̑-n- ‘sharpness.’ As such, Ocra ~ ῎Oκρα is the only unambiguous evidence for the existence of a deverbal abstract with o : e ablaut in Proto-Indo-European. In the addendum a brief etymological account is given of the place-name Acumincum ~ Acimincum from Pannonia Inferior, which likewise goes back ultimately to the PIE root *H2ek̑ -, arguing in favour of the Ptolemaic variant with Acu- (as opposed to Aci-) as the forma difficilior. The proposed etymology starts from PIE *H2ek̑-m̥n-o- > *akumno- ‘rock’ + *-enko- (*akumnenko- ‘rocky (place)’), from which *akumenko- > *akuminko- would then be produced by simple regressive dissimilation.

Specific constructive solution of a log house from southwest Serbia

The Dinaric log cabin was the dominant form of residential house in the Dinaric mountains region in the Balkans. In the valley of the Uvac River, in southwest Serbia, a structural solution with preserved cornice was recorded, which differs significantly from the solution of the Dinaric log cabin. It is characterized by a gable roof (instead of a four-gable one), and the beam on the ridge rests on a specially built structure consisting of a console and a rod. This solution indicates that there was an exchange of construction solutions in the Balkans and Europe, that they were transferred by builders, travellers or settlers. A similar construction solution is applied in Austria, Switzerland and Russia.

Fine-Scale Spatial Variability of Stand Structural Features under Selection Management and Strict Protection: An Example from the Dinaric Mountains

Small scattered plots of a few hundred square meters well reflect structural variability at stand level, but not at small spatial scales as the data between plots is missing. Information about structural similarities between managed and unmanaged stands, especially based on large sample plots, is still scarce. Our first objective was to quantify and illustrate structural variability of a selection-managed stand and a corresponding old-growth (OG) stand at small spatial scales. The second goal was to find out if there is a positive autocorrelation among neighboring patches in these stands regarding tree density (N) and basal area (BA). Tree positions and their diameters were recorded in 1.5 ha plots. Structural variation was examined at scales from 0.01 ha to 0.36 ha. Spatial correlation of N and BA was examined by applying experimental semivariograms. The variability of N was similar in both stands, whereas it significantly differed regarding BA (α = 0.05). Semivariance did not detect positive spatial autocorrelation of BA, while adjacent plots appeared to be more similar (autocorrelated) regarding N in both stands. Despite statistical difference regarding BA variability, the selection-managed stand exhibited many structural similarities to the OG stand, which makes it potentially suitable for modulating, if needed, to bring it step closer to an old-growth structure.

Monastic silviculture legacies and current old-growthness of silver fir (Abies alba) forests in the northern Apennines (Italy)

Species-rich mixed silver fir (Abies alba Mill.) forests dominated in the northern Apennines, but climate and, mainly, anthropogenic land use provoked a sharp silver fir decline approximately 5000 years bp. The conservation of the silver fir in this region was mainly due to the establishment of monastic orders that preserved and even planted silver fir for its spiritual and economic value. In 1993, the best silver fir stands were included in the Parco Nazionale delle Foreste Casentinesi (FCNP), Monte Falterona e Campigna, and have been submitted to low-intensive management or strict protection regardless of past land use and cultural history. In this study, we have (1) analyzed the current structure of three silver fir forests that have had different ownership histories and (2) compared the structure of the three Italian forests among them and with two old-growth forests from the Dinaric Alps as a reference of naturalness. The results show that the current structures of the three Italian forests are very different among them and are strictly related to past land use and, mainly, to monastic legacies. Even if the Italian forests have experienced decades of low-intensity management or strict protection, they are currently structurally very different from Dinaric old-growth forests. Developing an old-growth structure in these forests can be very slow and, in some ways, unpredictable. The results also highlight the importance of recognizing protected areas as cultural landscapes that host an important biocultural diversity. The current risk is that by applying almost exclusive biodiversity-centered management and setting difficult or impossible-to-achieve biodiversity goals, total diversity will decrease, and biocultural diversity, the greatest richness of most European parks, will be lost.

Further finding of Bolitophagus reticulatus (Linnaeus, 1767) in Montenegro (Coleoptera: Tenebrionidae) with brief comments on its distribution and conservation

The occurrence of Black Tinder Fungus Beetle Bolitophagus reticulatus (Linnaeus, 1767) (Coleoptera: Tenebrionidae), a saproxylic beetle of primordial and old-growth forests in the Palearctic region, is confirmed for Montenegro. The species has a fragmented distribution in the Alpine region and is not included in the IUCN red list for saproxylic beetles. B. sulcatus was found in the Biogradska Gora National Park, in the north-western part of the Bjelasica mountain range in the Dinaric Alps (Montenegro), in October 2022 while surveying the beetle fauna in an old-growth beech forest at about 1,000 m elevation.

An attempt to identify source areas of clastic deposits from selected caves of the Prokletije Mountains (Montenegro): a mineralogical and U-series geochemistry approach

The Prokletije Mountains/Albanian Alps are the highest part of the Dinaric Mountains. Paradoxically, these mountains and their cave areas remain among the most enigmatic in Europe, having only recently become subjects of scientific research and cave exploration. This paper focuses on the Kolata and Greben massifs, where the highest, and thus the oldest, cave systems are located. Through the analysis of heavy mineral composition, the apatite-tourmaline index, and cluster analysis for mineralogical composition and uranium isotopic data, it becomes possible to identify the source areas for the cave clastic deposits. It also allow to distinguish multiple paleoflow directions, each corresponding to different stages of cave development. The older paleoflow was probably from south to north, whereas the younger one is inferred to have flown from east to west. The change in the paleodirection of sediment transportation was probably caused by Neogene tectonic movements. In the later period, because of the subsequent uplift of the Prokletije Mts. and their fragmentation into individual massifs, the transport to the caves was local and related to deglaciations. The deposition of the speleothem from the oldest caves of the Prokletije Mts. took place in the periods c. 381 and 308 ka (MIS 11 and MIS 9), which suggests that these levels had already passed from the phreatic to the vadose stage. Considering the morphology of the caves studied, we believe that these caves may have begun to form at least in the Middle Pleistocene. There are two generations of younger speleothems first at 163 ± 4 ka (MIS 6) and second from 92 ± 2 to 39.4 ± 2 ka (Weichselian age).