2024 | 77/1 | 29–39 | 11 Figs. | 2 Suppl. | www.geologia-croatica.hr Journal of the Croatian Geological Survey and the Croatian Geological Society 1. INTRODUCTION 1.1. General background Caves often have well preserved sediments that contain valuable palaeontological and archaeological materials, and consequently, cave sediments are interesting because they may contain impor- tant records of past environments. In Croatia, the palynological analysis of cave sediments has rarely been performed (KUREČIĆ et al., 2021). Cave palynological investigations in the surround- ing region are also scarce (DOLÁKOVÁ, 2014; D’AGOSTINO et al., 2022). Even though palynology is very promising, it is also very challenging. Pollen taphonomy in caves depends on a di- verse number of factors: primarily production, dispersal, trans- port, deposition and post-depositional processes (COLES et al., 1989; HUNT & FIACCONI, 2018). The impact of animals and humans upon the composition of palynomorph assemblages is also very important. Therefore, the aim of this study was to ap- ply a combination of palynofacies and palynomorph analyses in cave sediment research. More specific objectives were: (i) to ob- tain additional information about the local vegetation plant cover in the past of the Adriatic area, (ii) to obtain information about temporal changes in the vegetation of that area, and (iii) to enable a better understanding of the post-depositional processes affect- ing the palynomorph assemblage. 1.2. Location, physical environment and vegetation Spila nad Procjepom cave (also known as the Briježica or Špilja kod Nerezinoga dola) is situated on the southern slope of Briježina Hill at 210 m a.s.l. overlooking Procjep Bay in the Mljet National Park (Fig. 1). The cave is a 35 m long chamber partly filled with sediments, formed in the Jurassic–Cretaceous dolostones with First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) Koraljka Bakrač 1, Olena Sirenko 1,5, Dario Hruševar 2,*, Ivona Baniček1, Vibor Novak3, Nataša Kletečki4 and Ankica Oros Sršen3 1 Croatian Geological Survey, Department of Geology, Sachsova 2, HR-10000 Zagreb, Croatia; (kbakrac@hgi-cgs.hr, osirenko@hgi-cgs.hr, ibanicek@hgi-cgs.hr) 2 University of Zagreb, Faculty of Science, Department of Biology, Horvatovac 102a, HR-10000 Zagreb, Croatia; (*corresponding author: dario.hrusevar@biol.pmf.unizg.hr) 3 Croatian Academy of Sciences and Arts, Institute for Quaternary Palaeontology and Geology, Ante Kovačića 5, HR-10000 Zagreb, Croatia; (aos@hazu.hr, vnovak@hazu.hr) 4 Elementary School Bogumil Toni, Perkovčeva 90, HR-10430 Samobor, Croatia; (natasa.kletecki@gmail.com) 5 The National Academy of Sciences of Ukraine, Institute of Geological Science, K55b O. Gonchara str., 01054 Kyiv, Ukraine; (o_sirenko@ukr.net) doi: 10.4154/gc.2024.04 Abstract This paper presents the first results of palynological research from the Spila nad Procjepom cave, situated in the Mljet National Park, Croatia. The palynological data obtained, enables a partial insight into the local vegetation cover, temporal changes in the vegetation during the ac- cumulation of studied deposits (at ca. 3500 cal years BP), and post-depositional processes that influenced the palynomorph assemblage. Results of palynofacies analysis indicate changes from fluvial (channel deposits), through palustrine to terrestrial environments. Although the interpre- tation of changes in plant cover, due to the lack of statistical significance, should be taken with caution, preserved pollen types confirm the dominance of the Mediterranean evergreen forest vegetation on Mljet island. Moreover, a high proportion of non-arboreal pollen (NAP) indicates some level of forest degradation, ranging from Mediterranean open forest to degraded maquis. Abundant charcoal additionaly confirms that the cave was inhabited by humans. limestone intercalations (HUSINEC, 2002). The entrance (di- mension 6 x 2 m) is oriented towards the southwest and the open sea. The entrance part of the cave is dry with a few remains of the flowstones that are no longer active, while the back of the cave is hydrologically active where drip water is still circulating, and flowstones are growing. The sediments studied in this research were sampled from the SE profile of test pit Sonda II in the back part of the cave, in the vicinity of a large flowstone (Fig. 2). The area of the Mljet National Park, as well as the entire island, be- longs to the macro-geomorphological region of South Dalmatia, with the archipelago and the meso-geomorphological region of the South Dalmatian Archipelago (BOGNAR, 1999). According to Köppen’s classification, the island of Mljet belongs to the Med- iterranean climate with hot summers (Csa) (ŠEGOTA & FILIPČIĆ, 2003). It means that winters are mild with abundant rainfall, in opposition to the long, dry and hot summers with fre- quent sunny days. The average annual air temperature on the is- land is 16.7 °C and it is one of the warmest areas in Croatia. The amplitude of average monthly air temperature values indicates the large thermal influence of the sea. Spring is cooler (due to the marine cooling effect) than the relatively warm autumn. The low- est mean monthly temperature is measured in January and Feb- ruary (8.8 °C each) and the highest values in July and August (26.0 °C and 25.8 °C, respectively). Average annual precipitation varies between 815 mm and 1021 mm, depending on the location and altitude (ŠPANJOL et al., 2016). Due to its geographical po- sition, the island of Mljet is located in the Mediterranean Biogeo- graphical Region. Plant cover of the island belongs to the Steno- mediterranean (near coastline), Eumediterranean (inland area) and Hemimediterranean (inland higher altitudes) vegetation zones (TRINAJSTIĆ, 1998; ŠPANJOL et al., 2016). Only in the Article history: Manuscript recieved: December 08, 2023 Revised manuscript accepted: February 13, 2024 Available online: February 27, 2024 Keywords: Adriatic Sea, Holocene, hydrological changes, Mediterranean Biogeographical Region, palynofacies, palynomorphs, palaeoenvironment, pollen mailto:ibanicek@hgi-cgs.hr mailto:dario.hrusevar@biol.pmf.unizg.hr mailto:vnovak@hazu.hr mailto:natasa.kletecki@gmail.com mailto:o_sirenko@ukr.net G eo lo gi a C ro at ic a Geologia Croatica 77/130 latter zone, do deciduous plants with a focus on Ostrya carpini- folia SCOP. have an important role (TRINAJSTIĆ, 1998). The other two vegetation zones are dominantly evergreen. So even though the island is nowadays almost completely covered with evergreen forest vegetation, it was not the case in the past. For- est habitats are the dominant type of vegetation, occupying more than three quarters of the island area (MESIĆ et al., 2009). Today the dominant forest trees are holm oak (Quercus ilex L.) and Aleppo pine (Pinus halepensis L.). Most of the National Park is overgrown by maquis (ŠPANJOL et al., 2016), and various deg- radation stages of holm oak forests (UGARKOVIĆ et al., 2019), while garrigues vegetation occupies only a small area (MESIĆ et al., 2009). The most abundant species of the maquis layer are ev- ergreen trees, such as the strawberry tree (Arbutus unedo L.), mock privet (Phillyrea media L.), tree heath (Erica arborea L.), Chios mastic (Pistacia lentiscus L.), common myrtle (Myrtus communis L.) and laurustinus (Viburnum tinus L.), followed by carob (Ceratonia siliqua L.), olive (Olea europaea L. var. sylves- tris BROT.), laurel (Laurus nobilis L.), and others. Among coni- fers, the most abundant is the genus Juniperus represented by brown-berried juniper (Juniperus oxycedrus L. ssp. oxycedrus), large-berried juniper (Juniperus oxycedrus L. ssp. macrocarpa (SM.) BALL) and Phoenicean juniper (Juniperus phoenicea L.). Pine forests, after maquis, represent the most abundant form of forest vegetation in the Park, while the large preserved Aleppo pine trees testify to the widespread appearance of the former au- tochthonous pine forests of Mljet (ŠPANJOL et al., 2016). 1.3. Previous palynological and archaeological research The island of Mljet has been palynologically investigated several times, e.g. BEUG (1967), JAHNS & VAN DEN BOGAARD (1998), and JAHNS (2002) so the Holocene vegetation changes are mostly well known. According to JAHNS (2002) who brings a synthesis of palaeovegetational changes, in the period preced- ing the Common Era, four main steps in vegetation development can be recognized: the deciduous Quercus woodland was re- Figure 1. Geographical location (red dot) and the entrance of the Spila nad Procjepom cave (island of Mljet). Figure 2. Plan of the Spila nad Procjepom cave with locations of the test pits Sonda I (S I) and Sonda II (SII). Sampled profile of SII is marked with a bold line. G eologia C roatica Bakrač et al.: First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) 31 placed by the evergreen Juniperus-Phillyrea community before ca. 7500 years BP, the latter was succeeded by the evergreen Quercus ilex woodland ca. 6000 years BP, followed by the Pinus dominated forest shortly after ca. 3000 years BP. Archaeologically and palaeontologically, the Spila nad Procjepom cave was investigated during five campaigns over the last 12 years (from 2010 to 2022) in collaboration with the Public Institution of the “Mljet National Park”. Two test pits were dug: Sonda I (“S I”) near the entrance and Sonda II (“S II”) in the in- ner part of the cave. While the silty, topmost sediments were rich in the remains of domesticated animals (mainly sheep and goats), marine fauna (fish and molluscs) and archaeological material (mainly pottery) spanning from the Copper Age to Medieval time (OROS SRŠEN et al., 2013; MAUCH LENARDIĆ et al., 2017), the flowstone discovered beneath provided a much older age. The sediment deposition in the cave lasted at least from the Last In- terglacial (MIS 5e), however, the breccia and the dolostone debris beneath could be of MIS 6 age or older (OROS SRŠEN et al., 2017). 2. MATERIALS AND METHODS 2.1. Fieldwork Three palynological samples were taken from the Sonda II (“S II”) test pit in order to investigate the preservation of palaeobot- anical remains in the cave deposits of Spila nad Procjepom. Test pit “S II” was chosen due to its well-preserved layers. The front- layer of the soil was cleared to a depth of 5 cm, and discarded to eliminate potential contaminants. Three samples were taken from the southeastern profile of the pit (Fig. 3), from different sublay- ers of Layer 4 that were chosen for its well-preserved fine-grained sediments. Layer 4 is mostly grayish brown silty clay (10YR-4/2) with various amounts of charcoal, hearth ash, as well as some burnt clay. All sublayers contain excellently preserved Holocene palaeontological/zooarchaeological remains, including sheep and goats and marine molluscs, as well as prehistoric archaeological material: pottery and a few stone tools (OROS SRŠEN et al., 2013). Samples for palynology were taken from the grayish-brown sed- iment that represents deposits less affected by human-produced fire (hearths – white/orange inclusions of profile on Fig. 3): • Sample P1 – Layer 4, sublayer 3: Overall sediment is yel- lowish-brown silty clay with charcoal particles and intercalations of grayish clay. Visible burrowing holes point to recent bioturba- tion. Sample P1 was taken at a depth of 0,60 m from the pit sur- face (sediment top). • Sample P2 – Layer 4, sublayer 4: Mostly reddish-brown silty clay with numerous charcoal hearth remains, with interca- lations of dark brown to gray silty clay. Sample P2 was taken at a depth of 0,80 m from the pit surface (sediment top). • Sample P3 – Layer 4, sublayer 4: Mostly grayish-brown silty clay with reddish and yellowish intercalations and traces of ash. Sample P3 was taken at a depth of 1,10 m from the pit sur- face (sediment top). One charcoal sample (Z – 5203) was collected from the hearth in Layer 4, sublayer 4 at 0,90 m from the pit surface dur- ing the excavation in 2012. The position of the hearth is 10 cm below sample P2. The charcoal sample was sent for radiocarbon dating (AMS 14C) at the Institute of Ruđer Bošković in collabo- ration with the University of Georgia, Center for Applied Isotope Studies. It is calibrated by Intcal20.14c (REIMER et al., 2020), with the error at the 1σ-level. 2.2. Laboratory work 2.2.1. Sample preparation The samples were processed for palynological analysis in the lab- oratory of the Croatian Geological Survey. For each sample ap- Figure 3. Sampling spots (P1, P2, P3) of Layer 4 (Sonda II test pit). G eo lo gi a C ro at ic a Geologia Croatica 77/132 prox. 18-30 g sediment was cleaned and crushed. Two Lycopodium tablets (Batch no. 280821291; 13761 spores/tablet) were added to each sample at the start of the processing to calculate palynomorph concentration according to MAHER (1981). The preparation pro- cedure included treatment with cold HCl (20%) to remove the car- bonates, and sodium pyrophosphate (Na4P2O7) to prevent coagu- lation, according to the standard techniques described in MOORE et al. (1991). Heavy liquid (ZnCl2, specific gravity 2.1 kg/l) was applied to separate the organic matter from the undissolved inor- ganic fraction. The organic residue was sieved through a 10 µm mesh. For palynofacies analysis slides were mounted in glycerin jelly, and the rest of the material was mixed in silicon oil for pa- lynomorph analysis. Microscopic analyses were performed using a Leica DM2500 microscope at x50, x100, x200, x400 and x630 magnifications combined with the differential interference con- trast (DIC). Photomicrographs were taken using a Leica MC190 HD camera connected to the Leica LAS EZ software. Sediment samples, organic residues and palynological slides are curated at the Department of Geology, Croatian Geological Survey. 2.2.2. Palynofacies analysis For palynofacies analysis, a minimum of 300 sedimentary or- ganic particles were counted. There are three main categories: amorphous organic matter (AOM), phytoclasts and palyno- morphs (TYSON, 1995). In this paper, we distinguish subcatego- ries: preserved phytoclasts (translucent, yellow-green plant tis- sue, and brown-black biostructured wood) and transformed phytoclasts (amorphous particles, gelified particles, and char- coal). Amorphous particles (AP) appear with diffuse but recog- nizable outlines that occasionally have residual internal struc- tures. Gelified particles (GP) present a homogeneous texture, variable colour (brown to amber), true outlines, and commonly angular shape, only occasionally with dulled angles. Charcoal particles are completely opaque, angular, and usually planar, black fragments. Samples are plotted in ternary diagrams using the software Past 4.13 (HAMMER et al., 2001). We used differ- ent types of diagrams to present the results. One of the more com- prehensive diagrams used in palynofacies studies is the APP dia- gram (AOM-Phytoclast-Palynomorph ternary diagram) presenting differences in relative proximity to terrestrial organic matter sources, transport paths, and the redox status of the depo- sitional environments (TYSON, 1995). We also used two more ternary diagrams: “Amorphous OM/Preserved Phytoclasts/ Transformed Phytoclasts” and “Opaque/Amorphous/Gelified Particles” (OP/AP/GP) to estimate the origin of the sedimentary organic matter (SOM) (aquatic/terrestrial), its source areas (veg- etation, soils, river), and controls of early diagenetic changes (aer- obic/anaerobic conditions, advanced oxidation) introduced by SEBAG et al. (2006b). To illustrate the origin (i.e. terrestrial or aquatic) and state of degradation of the organic particles samples, SEBAG et al. (2006b) proposed four optical indices: (i) “AOM contents” to quantify the aquatic contribution; (ii) “preserved phytoclasts / transformed phytoclasts ratio” to calculate the de- gree of degradation of the terrestrial fraction; (iii) “OP contents” to distinguish the allochthonous fraction from the fluvial origin; (iv) “GP/AP ratio” to differentiate the pedogenic or subaquatic degradation of terrestrial plant debris. To obtain information about the origin of the terrestrial fraction, we plotted a diagram with “OP contents” and “GP/AP ratio”. In order to have an over- view of the organic particle amounts in the sediments, we intro- duced “OM concentration” (particles/g) that is calculated by the following formula: OC = ((number of Lycopodium spores added) x (number of particles counted)) / ((number of Lycopodium spores counted) x (weight of dry sediment processed in grams)). 2.2.3. Charcoal analysis Samples for microscopic charcoal analysis were prepared as part of routine pollen analysis. They were counted as palynofacies par- ticles. Particles in the < 50 µm size bracket were not measured be- cause of their great abundance and minimal influence on the total charcoal sum, according to WHITLOCK & LARSEN (2001). Macroscopic charcoal (> 100 µm) was analysed from sediment sieved at 125 µm during the preparation procedure for pollen anal- ysis. All particles that were black with shiny surfaces were con- sidered to represent charcoal fragments. A qualitative approach was used to gain a general view of the fire history. Identification of pollen and plant spores followed standard keys, eg. MOORE et al. (1991) and BEUG (2015). A minimum of 100 palynomorphs were counted (five to six slides per sample) because the abundance was very low. All terrestrial pollen and spores were included in the pollen calculation sum. Indeterminable pollen counts were in- cluded in the total pollen sum because exclusion could result in bias in favour of well-preserved types (KAPP et al., 2000). When calculating the palynomorph percentages, the sum of all estab- lished pollen grains and spores, excluding algae and modern spores, was taken as 100%. The determination of pollen and spores was carried out according to the Engler classification system. Pa- lynological results, including changes in the arboreal/non-arboreal pollen (AP/NAP, i.e. woody vs. herbaceous plants), were presented as pollen percentage histograms, tables, and plates. 3. RESULTS 3.1. Radiocarbon dating The radiocarbon date of one charcoal sample (Z – 5203/UGAMS 14722) has yielded a date of 3160±25 years BP (3276 – 3448 cal years BP/ 1439 cal years BC). 3.2. Palynofacies Organic matter concentrations are relatively high ranging from 53.254 (SP22 P2) to 155.923 (SP22 P3) particles per gram. Pa- Figure 4. Phytoclasts ratio. G eologia C roatica Bakrač et al.: First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) 33 lynofacies of all three studied samples are dominated by phyto- clasts (Suppl. 1), mostly degraded woody tissue. Translucent, yel- low-green plant tissue (cuticle and membranous fragments) and amorphous organic matter are very rare (Fig. 4). In all three sam- ples macroscopic charcoal (>100 µm) is abundant (Fig. 5 a,c,e). In sample SP22 P3 charcoal is the most abundant particle type. Its ratio decreases in sample SP22 P2 and it has the lowest abun- dance in palynofacies of sample SP22 P1. At the same time, in sample SP22 P1 the degraded wood ratio increases, as well as the brown-black biostructured wood clasts (Fig. 5). 3.3. Palynomorphs Palynomorph concentrations are relatively low ranging from 1366 to 3225 palynomorphs per gram (Suppl. 2). The obtained materials made it possible to establish three spore-pollen com- plexes that characterize the deposits of the Spila nad Procjepom cave. Selected samples of palynomorphs are given in Fig. 6. In the established complexes, the pollen of woody and herbaceous plants is approximately equal in proportions. However, they have several differences (Suppl. 2). It is important to note that in the macerates of all samples, redeposited pollen from more ancient deposits was noted in different amounts, as well as modern spores of Polypodiaceae 3.3.1. SP22 P3 The palynomorph assemblage of sample SP22 P3 comprises 33.8% AP and 35.3% NAP. Distinctive features of the assem- blage are the highest percentage of spores (20.7%) of all three samples, belonging mainly to Polypodiaceae, the lowest amount of Ericaceae pollen (1.7%), as well as the highest content and taxonomic diversity of pollen from aquatic plants (Suppl. 2). The group of woody plants is dominated by Oleaceae pollen (23.2%) and closely followed by Pinus pollen (13.9%). The group of her- baceous plants is dominated by Cichoriaceae pollen (12.2%). The pollen of Poaceae and Asteraceae was recorded in approximately equal proportions, at 6.9% and 5.2%, respectively, with Valeri- ana pollen at 2.6%. Single pollen grains of Caryophyllaceae and Lamiaceae were also noted (Suppl. 2). A characteristic feature of the complex is the highest content and taxonomic diversity of hygrophilous pollen: Сyperaceae (2.6%), Typha latifolia (1.7%), Figure 5. Palynofacies SP 22 P-1 a-b; SP 22 P-2 c-d; SP 22 P-3 e-f. The scale bar is 200 µm for a, c, e and 50 µm for b, d, f. G eo lo gi a C ro at ic a Geologia Croatica 77/134 Liliaceae (1.7%), Alismataceae (0.8%). The highest number of spores of Polypodiaceae was also recorded in sample SP22 P3 (Suppl. 2). 3.3.2. SP22 P2 In the palynomorph assemblage of sample SP22 P2, the AP reaches 43.8%. Compared to the assemblage described above, the amount of Pinus pollen in its composition has decreased to 9.3%. However, the composition of prevailing pollen types, as well as the amount of Oleaceae pollen, remained at the level of the pre- vious sample complex (22.2%). A distinctive feature of the com- plex is the relatively high content of Ericaceae pollen (11.2%) and the appearance of Celastraceae pollen. Compared to the previous complex, the NAP amount has slightly increased (39.8%). This group doubled the amount of Asteraceae pollen (11.9%), with an accompanying high percentage of Cichoriaceae pollen (15.7%). Poaceae pollen content decreased to 5.9%, and only a single Ephedra pollen grain was noted. Compared to the previous pa- lynomorph assemblage, the amount of Caryophyllaceae pollen has doubled (1.9%), while Valeriana pollen reached 1.9%. Aquatic plants are represented by Typha latifolia pollen (2.6%). The spores belong mainly to Polypodiaceae and sporadically to Se- laginella. A Pseudoschizaea algae was also recorded (Suppl. 2). 3.3.3. SP22 P1 The palynomorph assemblage of the SP22 P1 sample reaches 44.2% of AP, while this group contains the highest quantities of Pinus pollen (18.5%). The composition of angiosperm pollen is dominated by pollen grains of Oleaceae (18.5%). Ericaceae (4.8%) is of subordinate importance. A distinctive feature of the Figure 6. Palynomorphs from the deposits of the Spila and Procjepom cave: (a)-(b) Pinus, (c) Oleaceae, (d)-(e) Ericaceae, (f )-(g) Asteraceae, (h) Chenopodiaceae, (i)-(j) Poaceae, (k) Caryophyllaceae, (l)-(m) Cichoriaceae, (n) Celastraceae, (o) Artemisia, (p) Liliaceae, (q) Cyperaceae, (r)-(s) Polypodiaceae, (t) Selaginella, (u) Ano- gramma, (v) Redeposited pollen, (w) Pseudoschizaea. The scale bar is 20 μm. G eologia C roatica Bakrač et al.: First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) 35 complex in sample SP22 P1 is the presence of pollen from broad- leaved species: Quercus, Ostrya and Acer (in total 2.4%). The AP and NAP are equal in proportion (44.2%). This group is domi- nated by the pollen of Cichoriaceae (13.2%) and Asteraceae (11.7%). The pollen of Poaceae (8.5%) acts as a subdominant. Pol- len grains of Caryophyllaceae (3.2%) and Artemisia (1.5%), Va- leriana (1.5%), as well as single pollen from Solanaceae and La- miaceae were observed. The group of aquatic plants is represented by the pollen of Typha latifolia (1.5%). Spores belong mainly to Polypodiaceae. Also, single spores of Anogramma, as well as the alga Pseudoschizaea were noted. 4. DISCUSSION 4.1. Depositional environment A relatively small amount of organic residue in all three samples with a high proportion of the opaque or semi-opaque phytoclasts indicates an oxic environment (TYSON, 1995), where other or- ganic compounds are selectively destroyed. The APP diagram (Fig. 7) shows the absolute domination of phytoclasts in all the analysed samples, indicating a pronounced input of terrestrial or- ganic material (TYSON, 1995). The dominance of structured phytoclasts in the total organic residue in all of the analysed sam- ples indicates the pronounced input of terrestrial organic compo- nents. Short fluvial transport is evident in phytoclasts’ structure, angularity and the fact that the clasts are unsorted (TYSON, 1995; MENDONÇA FILHO et al., 2017). Black oxidized phyto- clasts (charcoal) were derived from fires (wildfires or anthropo- genic fires), providing direct evidence of burning (TYSON, 1993; 1995). In the microscopic material, opaque particles are not al- ways charcoal, but rather gelified phytoclasts because higher tem- peratures convert the material to ash through glowing combus- tion, while lower temperatures may only lightly scorch the material, but not char it (WHITLOCK & LARSEN, 2001). Since charcoal is produced between temperatures of 280°C and 500°C, microscopic charcoal abundance increases during local fires. However, other proxy records, such as macroscopic charcoal or lithological changes, are needed to confirm if the fire was local. Abundant macroscopic charcoal (>100 µm) in all of the samples, coupled with charcoal, hearth ash, and some burnt clay in sedi- ment, confirms the local fire event. The “preserved/transformed phytoclasts ratio” allows quantification of the degree of degrada- tion of the terrestrial fraction. In the “AOM/Preserved phyto- clasts/Transformed particles” ternary diagram (Fig. 8), the dom- ination of transformed particles indicates a high plant-derived content (SEBAG et al., 2006a). SEBAG et al. (2006a) concluded that the “Preserved/transformed phytoclast ratio” decreases from production (i.e. litters) to depositional area (ditch, pond, alluvial deposits, etc.). In samples from the Spila nad Procjepom cave, the abundance of transformed particles points towards the soil origin. Typical soil assemblages are dominated by transformed phyto- clasts (SEBAG et al., 2006a). The ratio between opaque, amor- phous, and gelified particles is presented in the “Opaque/Amor- phous/Gelified Particles” ternary diagram (Fig. 9). More illustrative is the diagram of the “OP” and “GP/AP ratio” (Fig. 10). The “OP contents” increases with the oxidation of terrestrial Figure 7. Palynofacies (APP) ternary diagram. Figure 8. Amorphous OM/ Preserved Phytoclasts/ Transformed Phytoclasts ter- nary diagram. Figure 9. Opaque/ Amorphous/ Gelifies Particles (OP/AP/GP) ternary diagram. Figure 10. Origin of the terrestrial fraction. G eo lo gi a C ro at ic a Geologia Croatica 77/136 over periods of time (Fig. 11, Suppl. 2). Although our pollen sam- ples are not statistically significant due to the low pollen counts, well represented Oleaceae pollen speak in favour of olive-domi- nated stands at a distance of less than 500 meters from the cave (FLORENZANO et al., 2017). Moreover, the aerobiological re- search conducted in Babino Polje, the largest settlement on the island of Mljet whose inhabitants are mainly involved in the olive and vine cultivation, confirmed the relative dominance of airborn Oleaceae pollen in recent time (VOLARIĆ-MRŠIĆ, 1984). In contrast, pine pollen is not overrepresented as it may be expected due to recent plant cover on the island of Mljet, suggesting that Pinus probably did not participate in the local or extra-local ve- getation. According to CONNOR et al. (2004), pine trees are found near the sampling site only when its pollen comprises 35-50%, and these values were not reached in any of the studied samples. Within the herbaceous plants, pollen from Cichoriaceae, Asteraceae, Poaceae, Chenopodiaceae, and Caryophyllaceae were well represented. These families also include the largest number of drought-resistant taxa which complete their life cycle in one growing season and survive unfavourable conditions in the form of seeds. Additionally, Caryophyllaceae, Poaceae and Chenopodiaceae are indicators of open-ground vegetation in the Mediterranean area (FYFE et al., 2018; IZDEBSKI et al., 2020). Thus, herbaceous vegetation of all of the afore-mentioned taxa is widespread in the recent island flora (REGULA-BEVI- LACQUA & ILIJANI, 1984), as it was during the late Quaternary in the broader Mediterranean area (SUC et al.). Locally, the hydromorphic conditions are marked by the presence of Typha latifolia in all the analysed samples. This aquatic taxon is a com- mon wetland element and refers to water depths within the range of 10–60 (–100) cm (ŠUMBEROVÁ, 2011), and requires a mean July temperature above 15.7 °C (SCHENK et al., 2018). The aquatic-wetland component is highlighted in the SP22 P3 sample by the appearance of Alismataceae and Cyperacea pollen and the highest percentage of spores. Our results correlate very well with the zonation presented for southern Dalmatia by JAHNS (2002) and BASS (2008). JAHNS & VAN DEN BOGAARD (1998) divided the Quercus ilex period into three subzones (C1-C3). Whereas the Q. ilex re- mains at a relatively constant level, the associated vegetation is different. In subzone C1 Juniperus dominate, in subzone C2 Erica and in subzone C3 Pinus. In the samples from Spila nad Procjepom, Ericaceae have greater values than others and we can presume that the sediment in the cave was deposited during the subzone C2 (Quercus ilex – Erica) ranging from approximately 4370 years BP to 2670 years BP (JAHNS, 2002), which is in ac- cordance with our radiocarbon dating. Furthermore, detailed se- dimentological analyses are planned in order to shed more light on the genesis of the sediments and cave. 4.3. Taphonomic processes It is known that periodic wetting and drying of cave deposits as well as human activity (burning) can lead to extremely poor pol- len preservation (HUNT & FIACCONI, 2018). Pollen taphonomy in caves depends solely on transport paths and the depositional environment. Contemporary pollen must have entered the cave via aeolian transport or fluvial input or was brought in by ani- mals/humans. Animals could have easily picked up pollen from foliage in the vicinity of the cave during grazing and brought it to the cave on their fur and feet or in their gut contents (COLES et al., 1989). Daisy family, Asteraceae (entomophilous pollen), which have a great share in the pollen assemblage from Spila nad source materials. The “GP/AP ratio” is used to discriminate sites that have material reworked from the soil as opposed to sites where fresh plant debris falls directly into a basin and decays (SEBAG et al., 2006b). “OP contents” allows the recognition of fluvial (high values) and palustrine (low values) environments. The “GP/AP ratio” allows the distinction of soil (low values) and plant (high values) supplied areas. GP formation is related to the degradation of plant tissues in aquatic environments (pools, ponds, lakes) or anaerobic conditions (catotelm of peat, hydro- morphic soils). It is often observed in hydromorphic soils (NOËL et al., 2001). In the analysed samples, the domination of GP points to the degradation of plant tissues in aquatic and/or anaerobic conditions, especially in the sample SP22 P1, derived from hy- dromorphic soils, while SP22 P2 and SP22 P3 indicate a fluvial influence. Comparing the data from this study with data pre- sented by SEBAG et al. (2006b), SP22 P3 could be indicative of a fluvial origin (channel deposits), SP22 P2 of palustrine, and SP22 P1 of terrestrial origin (OF horizons sensu BAIZE & GI- RARD (2009) related to Oe horizons (Orthoeutric) from the World Reference Base (IUSS WORKING GROUP WRB, 2007, 2014)), moderately decomposed organic matter. A similar situa- tion to sample SP22 P3 was observed in the Lower Cerovačka Cave (Mt. Velebit, Croatia) where sample DC-SP 1 indicates a fluvial palaeoenvironment confirmed by sedimentological data (KUREČIĆ et al., 2021). 4.2. Vegetation history Due to a limited number of samples, and the fact that there was no sample taken from the surface near the entrance of the cave, which is a prerequisite for palaeofloristic reconstructions, the pre- sented reconstructions of vegetation have a preliminary chara- cter. Considering the above facts, the materials for the palynolo- gical study did not allow biostratigraphic dating of the analysed sediments, which is only a partial obstacle in the interpretation of local temporal changes in vegetation. In all the studied deposits (SP22 P3, SP22 P2 and SP22 P1), Oleaceae were the principal, dominant component within the woody plants, while the composition of subdominants changed Figure 11. Composition of plant groups during the formation of deposits in the Spila Cave. G eologia C roatica Bakrač et al.: First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) 37 Procjepom cave could have been brought by bees, flies or beetles (HUNT & FIACCONI, 2018). This is in accordance with NA- VARRO et al. (2001) who concluded that the amount of the zo- ophilous pollen (Asteraceae/Cichorioideae) increased from the direction of the cave entrance into the inner parts. Ground-living animals such as foxes, badgers, porcupines and rodents may be significant importers of pollen (HUNT & FI- ACCONI, 2018). In the Spila nad Procjepom cave, a small amount of rodents’ remains were recorded (MAUCH LENARDIĆ et al., 2017). According to CONNOR et al. (2004), Pinus pollen was probably brought from the other areas. Although Quercus ilex forests were recorded in the previous palynological research con- ducted on cores taken from the Mljet lakes (JAHNS & VAN DEN BOGAARD, 1998; JAHNS 2002), and it is native to the island, a statistically non-relevant amount of its pollen was found in the cave sediment. This could be due to preservation or accumulation factors. Quercus pollen could be under-represented because of its susceptibility to corrosion and oxidation (HAVINGA, 1964). He noted that the percentage of Quercus in the original pollen flora is much higher than in pollen found in sediment. An alternative cause for the evergreen oak decline in the vegetation record could be due to anthropogenic influence during the Greco-Roman colo- nization of the eastern Adriatic shores (BASS 2008). On the other hand, Ericaceae pollen is abundantly represented. More plausible than the idea of differential preservation is that animals, like sheep and goats, the remains of which were found in these sediments (BASS, 2008; OROS SRŠEN et al., 2013), influenced the pollen distribution more than wind or water. They could have easily picked up pollen from foliage in the vicinity of the cave during grazing. Findings of the palynomorph Pseudoschizaea, probably related to the Zygnemataceae family, indicate the runoff due to periods of enhanced soil erosion (LEROY et al., 2007) confirmed by palynofacies. ESTIARTE et al. (2008) also point out that Pseu- doschizaea is indicative of erosive processes, particularly when occurring with taxa such as Asteraceae, that are known as mark- ers of edaphic processes. This is also the case in Spila and is con- firmed by the palynofacies indicating the pronounced input of terrestrial organic material. VAN DE SCHOOTBRUGGE et al. (2024) concluded that Chomotriletes is the valid senior synonym of a variety of taxa, including Pseudoschizaea and Concentri- cystes. They considered it as a freshwater organism cyst, dominant in floodplain soils and ephemeral freshwater lakes. Since Polypo- dium is highly resistant to oxidation and corrosion (HAVINGA, 1964) it could be over-represented in samples from the soil. On the other hand, some forms that are not resistant to oxidation and corrosion are poorly preserved and indeterminable. 5. CONCLUSIONS The obtained data showed the effectiveness and prospects of com- plex palaeontological research in the study of cave sediments. Re- sults of palynological analysis enable a partial insight into the lo- cal plant cover, temporal changes in the vegetation during the accumulation of studied deposits and post-depositional processes that influence the palynomorph assemblage. According to the re- sults of palynofacies analysis, the provenance of the deposited samples ranges from a fluvial environment (channel deposits), through palustrine to hydromorphic soils. The results of the spore- pollen analysis allowed reconstruction of the differences in the composition of the vegetation cover that existed during the accu- mulation of the studied sediments. In all three analysed samples, the proportion of AP is slightly higher than NAP. Moreover, in all samples, Oleaceae pollen forms the basis of forest vegetation. The fact that pine pollen is not overrepresented, and olive pollen is pre- sent in the same quantity, indicates that the former was presum- ably absent from local/extra-local vegetation, and the latter prob- ably grew nearby. Although the interpretation of changes in plant cover, due to the lack of statistical significance, should be taken with caution, preserved pollen types confirm the dominance of the Mediterranean evergreen forest vegetation on the island of Mljet. Additionally, a high proportion of NAP pollen types indi- cate some level of forest degradation, ranging from Mediterranean open forest to degraded maquis. Sediment was most likely depos- ited during the subzone C2 (Quercus ilex – Erica) ranging from approximately 4370 years BP to 2670 years BP which was con- firmed by radiometric dating. The biostratigraphic dating of sed- iments, as well as the detailed reconstructions of palaeoenviron- ments, is possible only with the use of multidisciplinary studies including a broader set of analyses (e. g. palaeomagnetism, cos- mogenic 26Al/10Be burial dating of coarse sediment, environmen- tal magnetic studies of speleothems) and interpretation. Therefore, more detailed research is planned and will continue at the Spilja nad Procjepom cave shortly, to offer a more comprehensive insight into the depositional palaeoenvironment. ACKNOWLEDGMENT This study was supported by the Foundation of the Croatian Academy of Sciences and Arts, Project Preliminarny palaeobo- tanical investigations of the Spila nad Procjepom cave in Na- tional Park Mljet and Public Institution “Mljet National Park” and by the Croatian Geological Survey through programme fund- ing provided by the Croatian Ministry of Science and Education. We would like to thank colleagues from the Institute for Quater- nary Palaeontology and Geology of the Croatian Academy of Sci- ences and Arts and the employees of Public Institution “Mljet National Park” for their help during excavations. Dragica KOVAČIĆ from the Croatian Geological Survey is also thanked for laboratory assistance, and Mateo PETROVIĆ from the Insti- tute for Quaternary Palaeontology and Geology of the Croatian Academy of Sciences and Art for editing the plan and photos. We also thank Maja ANDRIČ and other reviewers for constructive comments that improved this paper. FUNDING Some parts of this research were funded by the Foundation of Croatian Academy of Sciences and Arts (grant number 53- 114/2022), and by the internal research project „ZG-LAB“ at the Croatian Geological Survey, funded by the National Recovery and Resilience Plan 2021–2026 of the European Union – Next- GenerationEU, monitored by the Ministry of Science and Educa- tion of the Republic of Croatia. 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Kluwer Aca- demic Publishers, Dordrecht, 75–97 p. doi: 10.1007/0-306-47668-1_5 G eologia C roatica Bakrač et al.: First palynological results from Spila nad Procjepom cave, Mljet island (Croatia) 39 SAMPLE SP 22 P1 SP 22 P2 SP 22 P3 W O O D Y P H YT O CL A ST S Brown-black biostructured wood 51 21 25 Translucent, yellow-green plant tissue 7 6 1 Degraded wood (AP) 110 44 113 Degraded wood (GP) 196 173 125 Corroded charcoal 9 87 125 Lath-shaped charcoal 2 6 39 Equant-shaped charcoal 5 14 12 Charcoal 16 107 176 Preserved phytoclasts 58 27 26 Transformed phytoclasts 322 324 414 TOTAL PHYTOCLASTS 380 351 440 AOM 4 0 2 PA LY N O M O RP H S Bisaccate pollen 1 3 1 Angiosperm pollen 7 21 3 Spores 6 10 3 Freshwater algae 1 6 0 TOTAL 399 391 449 Lycopodium - batch 280821291 3 11 3 OM “concentration” (particles / g) 120791 53254 155923 SAMPLE SP 22 P1 SP 22 P2 SP 22 P3 spore 9 18 24 7.0% 11.8% 20.7% Pinus subgenus Pinus (Diploxylon) 24 15 16 18.5% 9.8% 13.9% Oleaceae 24 34 27 18.5% 22.2% 23.2% Ericaceae 6 17 2 4.8% 11.2% 1.7% Ostrya 1 0 0 0.8% 0.0% 0.0% Quercus 1 0 0 0.8% 0.0% 0.0% Acer 1 0 0 0.8% 0.0% 0.0% Celastraceae 0 1 0 0.0% 0.7% 0.0% Ephedra 0 1 0 0.0% 0.6% 0.0% Cyperaceae 0 0 3 0.0% 0.0% 2.6% Poaceae 11 9 8 8.5% 5.9% 6.9% Chenopodiaceae 2 0 0 1.6% 0.0% 0.0% Asteraceae 15 17 6 11.7% 11.9% 5.2% Artemisia 2 0 0 1.5% 0.0% 0.0% Cichoriaceae 17 24 14 13.2% 15.7% 12.2% Caryophyllaceae 4 3 1 3.2% 1.9% 0.8% Solanaceae 1 0 0 0.8% 0.0% 0.0% Lamiaceae 1 0 1 0.8% 0.0% 0.8% Valeriana 2 3 3 1.5% 1.9% 2.6% Typha latifolia 2 4 2 1.5% 2.6% 1.7% Liliaceae 0 0 2 0.0% 0.0% 1.7% Alismataceae 0 0 1 0.0% 0.0% 0.8% undiff. 6 7 6 4.5% 4.6% 5.2% Pseudoschizaea 1 1 0 Redeposit pollen 5 20 2 Modern Polypodiaceae 10 22 35 ex. Lycopodium 50 96 120 mass (g) 30.00 18.37 26.30 Ls (18584) 2 2 2 Pollen sum 129 153 116 Concentration 3196 3225 1366 Total sum of pollen and spores 129 153 116 Sum of arboreal pollen 55 67 45 42.6% 43.8% 38.8% Sum of non-arboreal pollen 57 61 41 44.2% 39.8% 35.3% Sum of spores 9 18 24 7.0% 11.8% 20.7% Supplement 1. Palynofacies Supplement 2. The content of palynomorphs