www.geologia-croatica.hr Journal of the Croatian Geological Survey and the Croatian Geological Society 2025 | 78/3 | 205–220 | 9 Figs. | 4 Tabs. | 1 Suppl. | Article history: Manuscript received: January 15, 2025 Revised manuscript accepted: July 15, 2025 Available online: October 13, 2025 Keywords: Mollusca, Foraminifera, Ostracoda, fossil assemblages, palaeoecology, community responses, human impact 1. INTRODUCTION 1.1. Combining Conservation and Stratigraphic Palaeobiology Conservation palaeobiology aims to expand the temporal per- spective of conservation science by integrating ecology and palaeobiology to protect ecosystems and enhance biodiversity and to evaluate ecological responses to both human activities and natural environmental changes (JACKSON, 2001; WIL- LIS & BIRKS, 2006; JACKSON & HOBBS, 2009; DIETL & FLESSA, 2011). The differences between recent benthic com- munities and palaeoecological baselines observed in the Pleis- tocene and Holocene fossil records can be used to assess the magnitude and timing of human impacts on populations, com- munities and ecosystems (DIETL & FLESSA, 2011). The in- terdisciplinary nature of conservation palaeobiology provides a comprehensive approach to understand ecosystem changes by incorporating palaeobiological, archaeological, and histor- Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments (northern Adriatic Sea) Michaela Berensmeier1,*, Adam Tomašových2, Rafał Nawrot1, Vlasta Ćosović3, Eva Hasenzagel1, Petra Heinz1, Martin Zuschin1 1 University of Vienna, Institute for Palaeontology, Josef-Holaubek-Platz 2 (UZA II), 1090 Vienna, Austria; (*corresponding author: michaela.berensmeier@univie.ac.at; rafal.nawrot@univie.ac.at; eva.hasenzagel@gmx.at; petra.heinz@univie.ac.at; martin.zuschin@univie.ac.at) 2 Slovak Academy of Sciences, Earth Science Institute, Dubravska cesta 9, Bratislava, 84005, Slovakia; (adam.tomasovych@savba.sk) 3 University of Zagreb, Faculty of Science, Department of Geology, Horvatovac 102b, 10000 Zagreb, Croatia; (vcosovic@geol.pmf.hr) doi: 10.4154/gc.2025.16 Abstract Human-induced changes in sedimentation rates, pollution, and eutrophication significantly transformed the benthic communities in the northern Adriatic Sea during the late Holocene, particularly in recent centuries. Fossil assemblages from sediment cores record these changes but are often affected by stratigraphic condensation and mixing. Here, we show that molluscan, foraminiferal, and ostracod assemblages preserved in a condensed sedi- ment core collected off the Po delta at 31 m water depth, still archive information about the composition of benthic communities prior to anthropogenic changes. All three groups exhibit a similar trend in total abundance (density): a gradual increase peaking in a 10 cm-thick shelly lag (≈ 2 – 6 kyr BP) with millennial time-averaging, followed by a significant decline in the uppermost 5 cm of highstand silts (representing the past 2 kyr BP) with centennial time-averaging. The molluscan assemblage in the shelly lag integrates across several base- line community states. The assemblage mainly comprises shallow-subtidal filter feeders and soft-bottom infauna, with the bivalve Varicorbula gibba and the gastropod Turritellinella tricarinata dominating and increasing in proportional abundance in the highstand silts. The ostracod and benthic foraminiferal assemblages in the shelly lag are dominated by oppor- tunistic species such as Cytheridea neapolitana, Ammonia beccarii, and Haynesina germanica. In contrast, the foraminifera Nonionella sp. and the ostracod Loxoconcha sp. increase in proportional abundance in the highstand silts, characterised by an increase in filter-feeders among the molluscs, infaunalisation and a decrease in epiphytic species. Although an increase in net sediment accumulation primarily causes the decline in fossil density in the uppermost part of the core, upward changes in the relative abundance of species and functional groups reveal a difference between the baseline and impacted community states. Therefore, the time-averaged fossil assemblage in the shelly lag provides a valuable long-term record of an ecosystem in the region before human impact. ical data. Even the youngest fossil records can be affected by biases due to the variable completeness and resolution of fos- sil assemblages (e.g., SCARPONI et al., 2017; TOMAŠOVÝCH et al., 2023; NAWROT et al., 2024; ZECCHIN et al., 2024). These biases can impact our understanding of temporal popu- lation and community dynamics (AGER, 1973; PATZKOWSKY & HOLLAND, 2012). Stratigraphic palaeobiology is a sub- discipline of palaeobiology that examines spatial and temporal patterns of fossil distribution in a stratigraphic context. This approach explicitly recognises that both biological processes, such as speciation, extinction, and community assembly, as well as stratigraphic processes, including sedimentation, ero- sion, and condensation, impact the fossil record (DRESOW, 2023). Consequently, the integration of conservation palaeo- biology with the toolkit used by stratigraphic palaeobiology can provide more rigorous reconstructions of the recent geo- logical past and increase the reliability of the data on the long- term history of human impact on ecosystems. G eo lo gi a C ro at ic a 206 Geologia Croatica 78/3 1.2. Stressors in the northern Adriatic The northern Adriatic Sea is a dynamic and diverse ecosystem that has experienced significant changes due to the combined effects of natural processes and human activities over the past millennia. This has led to considerable alterations in the com- position of marine communities, making it one of the most dis- turbed continental shelves on Earth (LOTZE et al., 2011; GALLMETZER et al., 2019; HASELMAIR et al., 2021; SCARPONI et al., 2022) (see Table 1). It is essential to analyse fossil assemblages to understand the long-term effects on local ecosystems, as systematic ecological monitoring only began in the 20th century. Data from the Holocene fossil record show that the faunal composition in the northern Adriatic has changed significantly over the past millennia and centuries (e.g., TOMAŠOVÝCH et al., 2017, 2018; GALLMETZER et al., 2019; SCARPONI et al., 2022). While some species have drastically decreased in abundance, a few opportunistic spe- cies survived and proliferated (GALLMETZER et al., 2017; HASELMAIR et al., 2021). However, most studies that have examined Holocene changes in the composition of benthic as- semblages within sediment cores have focused on single taxo- nomic groups, such as macrofauna (e.g., molluscs) (SCAR- PONI et al., 2017; GALLMETZER et al., 2017; MAUTNER et al., 2018) or microfauna (e.g., foraminifera or ostracods) (BAR- MAWIDJAJA et al., 1995; VIDOVIĆ et al., 2016; PICONE et al., 2008). For instance, benthic foraminifera in the sediments of the Po delta have recorded increased eutrophication and a decline in oxygen availability, which are linked to anthropo- genic changes in the Po outflow (BARMAWIDJAJA et al., 1995). Comprehensive quantitative studies that assess the abun- dance and diversity of multiple phyla simultaneously remain rare (see, e.g., D’AMICO et al., 2013; ROSSI et al., 2021; AZZARONE et al., 2020). Furthermore, few multi-taxon stud- ies connect biotic signals to sediment parameters such as grain- sizes or organic matter content (BARBIERI et al., 2019, 2021). 1.3. Objectives of the present study Quaternary sequences provide a significant advantage for studying palaeoecological patterns within a sequence strati- graphic framework. The assemblages primarily consist of ex- tant taxa with well-documented ecological, biogeographic, and phylogenetic relationships. This palaeoecological data can be compared with assemblage states observed in modern envi- ronments, offering insights into how human impacts affect the functioning of contemporary ecosystems. In this study, we take a comprehensive approach to exam- ine Holocene changes (over the past ≈ 10,000 years) in the composition of assemblages formed by three different phyla, all within the context of the sequence stratigraphic framework, at a site located in the central part of the northern Adriatic Sea. This method, which combines conservation and stratigraphic palaeobiology, has been successfully applied at other Holocene locations in the northern Adriatic Sea (e.g., WITTMER et al., 2014; GALLMETZER et al., 2019; BARBIERI et al., 2019). However, none of the previous studies directly compared re- sponses of macro- and microfaunal assemblages to natural and human-induced environment changes. BERENSMEIER et al. (2023) described the geochronol- ogy, geochemistry, and molluscan assemblages at this coring location. Despite intensive stratigraphic condensation and mixing of the fossil record at this site, the assemblages and sediments enriched in metals and organic carbon in the up- permost core increments reveal evidence of human impacts. Furthermore, the core-chronological data from this study, based on molluscs, was used in a palaeoecological study of core data from the northern Adriatic by TOMAŠOVÝCH et al. (2024). They found that species diversity in fossil assem- blages increases with the time averaging of samples, which is influenced by varying sediment accumulation. The main objectives of the present study are to: (i) de- scribe the faunal composition of molluscan, benthic foramini- feral, and ostracod fossil assemblages; (ii) analyse their diver- sity and species composition within the stratigraphic context; and (iii) assess whether the three types of assemblages respond differently to human impacts. 2. MATERIALS AND METHODS 2.1. Sampling locality and stratigraphic framework In June 2017, the team of F. S. Poseidon-cruise no. 514 took the gravity core POS514-GC25-5 in the western part of the northern Adriatic Sea, ≈ 32 km to the East of the Po delta Pila Table 1. Selected stressors impacting the benthic ecosystem in the northern Adriatic. Stressor Causes (simplified) Impact on the benthic ecosystem References (selection) Hypoxia and anoxia Human-induced eutrophication in combination with ocean stratification Mass mortalities; alteration of sediment biogeochemistry; alteration of faunal composition STACHOWITSCH, 1984; JUSTIĆ, 1991; STACHOWITSCH, 2014 Mucilage aggregates Diatom-blooms after human-induced eutrophication Anoxia; mass mortalities STACHOWITSCH et al., 1990; DEVESCOVI & IVEŠA, 2007; KRAUS & IVOŠEVIĆ DENARDIS, 2023 Chemical pollution Human-induced influx of heavy metals, organic pollutants or other Alteration of faunal composition; mass mortalities Molluscs: GALLMETZER et al., 2017; SCHNEDL et al., 2018 Benthic foraminifera: FRONTALINI & COCCIONI, 2011; MELIS et al., 2019 Ostracods: BERGIN et al., 2006 Alien species Shipping, aquaculture activities, or a natural range expansion Competitive interactions with native species; alteration of the faunal composition; parasitism ORLANDO-BONACA, 2010; OCCHIPINTI-AMBROGI et al., 2010; CROCETTA, 2011 Bottom trawling Still used fishing technique Alteration of faunal composition; decrease in faunal densities; habitat loss KOLLMANN & STACHOWITSCH, 2008; PITCHER et al., 2022 Heat waves Human-induced climate change Alteration on faunal composition; mortality of temperature sensitive species; alien invasions, habitat loss PAIRAUD et al., 2014; GÓMEZ-GRAS et al., 2022 G eologia C roatica 207Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... mouth, at 31 m water depth (Fig. 1). This 3-m-long core, 16 cm in diameter, comprises Holocene sediments, of which the upper 75 cm represent the Holocene marine transgression. The sedimentology, stratigraphy, and geochemistry of the core and its sequence-stratigraphic framework and facies are described in BERENSMEIER et al. (2023) and follow the regional-scale Holocene facies architecture of the Po coastal plain and delta (e.g., AMOROSI et al., 2017; BRUNO et al., 2017; CAMPO et al., 2017). Based on faunal content and core chronology, the studied core section is divided into four facies from bottom to top: (i) shell-poor, bioturbated sandy silts (40 – 75 cm), (ii) silty sands (22.5 – 40 cm), and (iii) a shelly lag (10 – 22.5 cm), which together represent the transgressive systems tract, and (iv) shell-poor, bioturbated muddy silts belonging to the highstand systems tract (0 – 10 cm) (Fig. 2). The assignment of the facies to the depositional sequences was discussed in BERENSMEIER et al. (2023), who documented two major stratigraphic surfaces in the record: the maximum flooding surface and the wave ravinement surface (Fig. 2). These two surfaces define the general sequence stratigraphic architecture of the studied core, which is consistent with the regional architecture in the north- western Adriatic (e.g., AMOROSI et al., 2017; SCARPONI et al., 2017). While the maximum flooding surface marks the transition from a transgression to regression (AMOROSI & COLALONGO, 2009), the wave ravinement surface forms in shallow-water settings during transgression (ZECCHIN et al., 2019, and references therein). The core was sliced into 2.5-cm thick increments in the upper part (0 – 22.5-cm core depth) and into 5-cm-thick incre- ments in the remaining part. Bulk sediment geochemistry was shown in BERENSMEIER et al. (2023). Increased nitrogen and heavy metal concentrations, such as mercury, are detected in core depths 0 – 10 cm, suggesting eutrophication and pol- lution impacting the study area in the last centuries. The upper 75 cm of the core were deposited under very low net sedimen- tation rates over the past ≈ 10,000 years, as indicated by the ages of bivalve shells (Varicorbula gibba, Lentidium mediter- raneum) dated with radiocarbon-calibrated amino acid race- Figure 1. Location of the sampling station in the northern Adriatic Sea. a Satellite image of Italy and its surrounding sub-basins of the Mediterranean Sea. MODIS satellite image (Terra) of the northern Adriatic coast (EOIMAGES, 2024), collected on day 335, 2002, at 1030 Greenwich Mean Time (GMT); b Bathymetric map of the study area, including the core position, ca. 30 km off the Po della Pila mouth. Approximate core/sampling station from other studies: a – GALLMETZER et al., 2019, core Venice; b – GALLMETZER et al., 2019, NAWROT et al., 2022, cores Piran 1-2; c – TOMAŠOVÝCH et al., 2017, 2018, core Panzano; d – TOMAŠOVÝCH et al., 2017, core Po 3; e – TOMAŠOVÝCH et al., 2017, core Po 4; f – BARMAWIDJAJA et al., 1995, core 108; g – HRS-BREN- KO, 2006, station SJ101; h – PICONE et al., 2008, core AAL87-19; i – PICONE et al., 2008, core AMA88-39. Figure 2. Stratigraphic framework and geochronological data of the core from BERENSMEIER et al. (2023). Median ages are based on age-dated shells of Varicorbula gibba, Lentidium mediterraneum, and plant remains, all ages in years before the year the core was collected (AD2017). G eo lo gi a C ro at ic a 208 Geologia Croatica 78/3 mization (AAR) methods and 210Pb dating of the sediments (BERENSMEIER et al., 2023). The median postmortem ages (Varicorbula gibba, Lentidium mediterraneum, and coralline algae) increase down-core (BERENSMEIER et al., 2023). With the exception of centennial time-averaging in the upper- most increment (interquartile age range (IQR) of 300 years at 0 – 2.5 cm), the majority of the subsurface record at this site is affected by millennial time-averaging (3,400 years at 5 – 7.5 cm, 1,900 years at 10 – 12.5 cm and 3,000 years at 17.5 – 20 cm) and a sedimentation rate of ≈ 0.1 mm/yr (BERENS- MEIER et al., 2023). The assemblages formed by all skeletal remains in the gravity core are termed here as fossil assem- blages. The term subfossil is not used here despite shell ages younger than 10,000 years. 2.2. Palaeoecological analyses The benthic foraminiferal and ostracod assemblage analysis was performed on subsamples taken from each increment. These subsamples had a volume of circa 30 cm3 in the uppermost 22.5 cm and 60 cm3 in the remaining part of the core. Each subsample was sieved through a 63 µm mesh size and standardised using a microsplitter. A minimum number of 300 benthic foraminiferal tests was counted in each sample of the upper 50 cm of the core. Below that depth, at least 250 tests per increment were counted due to low faunal density. Another subsample from each increment, representing the same volume of sediment used for foraminiferal analysis, was sieved for sieve size fractions 1 mm – 250 µm and 250 – 125 µm, in which all adult ostracod valves were identified and counted. All material >1 mm, including subsamples for benthic foraminifera and ostracods, was used to count molluscan remains. For each bivalve species, the higher number of single valves (either right or left) was added to the number of articulated specimens to get the minimum number of unique individuals in each increment. Gastropods were counted if at least 50% of the shell was preserved and the aperture was present. Only scaphopods with unbroken shell openings were counted. In polyplacophorans, abundance was assessed by adding the number of head valves (each representing one individual) to the number of other plates divided by eight, with decimal places rounded to integers (see Supplement 1 for a complete list of species and their abundances in each core increment). The focus was on the marine species; freshwater or ter- restrial taxa (mainly occurring at the base of the core) are rare and reworked and thus were omitted from analyses. The clas- sification of benthic foraminifera at the species level is based on the criteria of LOEBLICH & TAPPAN (1987). The result- ing species names were standardised according to the World Register of Marine Species (WoRMS, AHYONG et al., 2025). Due to poor preservation of the tests, the most common ben- thic foraminifera identified as Haynesina germanica and Haynesina sp. were summarised as Haynesina spp. Small tests of Ammonia sp. that were not identified to the species level were pooled with other Ammonia species (A. bec- carii, A. inflata, A. tepida) into Ammonia spp. Similarly, Tex- tularia agglutinans, T. bocki, and T. conica were pooled into Textularia spp. Faunal densities of each taxon were calculated as the num- ber of unique individuals divided by the volume of the entire or subsampled increment. Species diversity is reported as (i) the total number of species in each increment or subsample, (ii) the Shannon diversity index (effective number of species, exponential of the Shannon entropy H), and (iii) the Fisher’s α index. The two calculated diversity indices, Shannon and Fish- er’s α, capture different aspects of diversity. The Shannon di- versity index is a measure of entropy (uncertainty in species identity of a randomly selected individual) and considers the equal distribution of individuals across species, taking into account the abundance of each species (JOST, 2007). Fisher’s α index is more focused on the absolute number of species and the abundance of rare species (SCHULTE et al., 2005). We calculate both indices to show how the diversity and heteroge- neity of the three fossil assemblages (mollusc, benthic fo- raminifera and ostracod) changed in the stratigraphic context. To account for variability in fossil abundance, we used rarefac- tion to compare species richness and Shannon diversity be- tween increments. Mean values of rarefied species richness and Shannon diversity expected at 50 specimens with corre- sponding 95% confidence intervals (based on the percentile method) were calculated based on 1,000 resampling iterations. The multivariate analyses are based on proportional spe- cies and genera abundances. Non-metric multidimensional scaling (nMDS) based on Bray-Curtis distances and square- root transformed relative abundances was used to visualise changes in the assemblage structure along the core for each of the three fossil assemblages (molluscs, benthic foraminifera and ostracods) separately, (using only samples containing a minimum number of 50 individuals per increment) following KOWALEWSKI (2015). These ordinations visualise the changes in the assemblage structure and the differences be- tween the subsurface increments, (undisturbed by human im- pacts) and the uppermost core increments affected by pollution and human impacts. In addition to the nMDS, constrained hierarchical cluster analyses were carried out to evaluate whether groupings of in- Table 2. Mean relative abundances of feeding guilds and habitat prefer- ences of the molluscan assemblages in the depositional sequences: high- stand silts, 0 – 10 cm and the shelly lag 10 – 22.5 cm. All values are given in %. Feeding guilds of the molluscs Shelly lag Highstand silts Filter feeder 69 73 Detritus feeder 13 14 Carnivore 11 6 Scavenger 4 3 Grazer 2 2 Chemosymbiont 1 2 Habitat preferences Shelly lag Highstand silts Infauna 77 80 Epifauna, hard bottom 9 8 Epifauna, soft or hard bottom 8 6 Semi-infauna 2 1 Epifauna on vegetation 2 1 Epibiont/ectoparasite 1 3 G eologia C roatica 209Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... crements with similar faunal composition correspond to dis- tinct stratigraphic units. The cluster analysis considers the temporal sequence of samples (PATZKOWSKY & HOL- LAND, 2012). For this purpose, the CONISS algorithm from the “chclust” function of the “rioja” package was applied to the Bray-Curtis distance matrix (JUGGINS, 2024). Since the core comprises undisturbed and disturbed peri- ods in the depositional history of the northern Adriatic, we have calculated biotic indices measuring the ecological quality of benthic habitats based on the fossil assemblages. The Marine Biotic Index (AMBI) of BORJA et al. (2000), based on the tol- erance of macrobenthic taxa towards pollution, was applied to the molluscan assemblages to determine temporal changes in the Ecological Quality Status along the stratigraphic succes- sion. Version 6.0 of AMBI software was used (AMBI, 2024). The Foram Stress Index (FSI, after DIMIZA et al., 2016) was calculated to evaluate whether foraminiferal assemblages re- cord environmental stressors. The FSI is based on the relative abundances of sensitive and stress-tolerant (opportunistic) spe- cies according to their organic matter sensitivity (JORISSEN et al., 2018; DIMIZA et al., 2016). To classify the foraminiferal species according to their stress sensitivity or tolerance, the methods of JORISSEN et al. (2018) and ŽVAB ROŽIČ et al. (2022) and the reference therein were followed. The enhanced Benthic Foraminifera Oxygen Index (eBFOI) was calculated based on relative abundances of species typical of oxic, suboxic or dysoxic environments (KRANNER et al., 2022). All statis- tical analyses were performed in R 4.0.3 and RStudio version 2023.12.0.369 (R CORE TEAM, 2021; POSIT TEAM, 2023) using the “vegan” package (OKSANEN et al., 2025). 3. RESULTS 3.1. Species composition Three benthic phyla dominate the core’s fossil content in dif- ferent grain size fractions: Mollusca dominate the >1 mm size fraction, Ostracoda the125 µm – 1 mm, and Foraminifera the 63 µm – 125 µm. These three groups share the same up-core pattern in densities: (i) low densities below the shelly lag (75 – 22.5 cm), (ii) a maximum density in the shelly lag (22.5 – 10 cm), and (iii) an abrupt decrease in the top 10 cm (Fig. 3). The molluscan total assemblage comprises 3180 bivalve individuals (56 species), 1009 gastropods (43 species), 102 scaphopods (1 species), and 12 polyplacophorans (2 species). The most abundant species is the shallow-burrowing, suspen- sion-feeding bivalve Varicorbula gibba that can also feed on resuspended detritus and benthic diatoms (YONGE, 1946; KIØRBOE & MØHLENBERG, 1981; HRS-BRENKO, 2006), and the semi-infaunal, filter-feeding gastropod Turri- tellinella tricarinata (Fig. 4). All scaphopods belong to the in- faunal micro-carnivore Antalis inaequicostata. Polyplacopho- rans are represented by Chiton olivaceus and Acanthochitona fascicularis. The 15 most abundant molluscan species com- prise about 70% of the molluscan assemblages. The molluscan density increases gradually from 40 cm up-core, from less than 3, to 134 molluscs per 100 cm3 (Fig. 3). The shelly lag bears the highest densities of all mollusc species (Fig. 4). In the overlying highstand silts, the faunal densities decrease again. Formerly abundant species such as Parvicardium sca- brum, Ostrea spp., or Nucula cf. nucleus decrease in relative abundance, whereas T. tricarinata increases to 16% (Fig. 4). The foraminiferal total assemblage comprises 8937 indi- vidual tests (65 species and 23 genera in open nomenclature). The foraminiferal assemblages are dominated by Rotaliida (Table 3). The most abundant taxa are the rotaliids Haynesina spp. (forming 10 – 36% of the assemblages) and Ammonia spp. (11 – 26%), followed by the textulariid Textularia spp. (4 – 16%). The 15 most common benthic foraminifera form about 72% of the total assemblage (Fig. 5). Foraminiferal densities decrease significantly at 70 cm, and no benthic foraminifera were found below 77.5 cm. Within the shelly lag, the fo- raminiferal density peaks at ≈ 940 individuals per 1 cm3 (Fig. 3). In the highstand silts, the foraminiferal densities decreased drastically, apart from representatives of Nonionella sp., which were more numerous in this interval (ca. 11% of the foramini- feral assemblages). The ostracod total assemblage comprises 2528 individu- als (12 species and 7 genus-level taxa). Most ostracods belong to marine species, except two non-marine species that attain less than 1% of the total assemblage: Cyprideis torosa (brack- ish) and Candona sp. (freshwater). An additional six taxa con- Figure 3. Faunal densities, species richness, Shannon and Fisher diversity of the ostracod, foraminiferal and molluscan assemblages. Maximum values of fossil density are indicated next to the graphs; faunal densities of all three taxa peak within the shell lag. A decrease in species richness and Shannon diversity in the highstand silts is only recorded in the molluscan assemblages. G eo lo gi a C ro at ic a 210 Geologia Croatica 78/3 stitute less than 1% of the total ostracod assemblage and are not displayed in Figure 6: Aglayocypris sp., Aurila spp. (A. convexa and A. speyeri), Loxoconcha rhomboidea, Semi- cytherura cribriformis, Semicytherura sp., Xestoleberis com- munis. The ostracod assemblages are strongly dominated by the marine species Cytheridea neapolitana, which forms up to 53% of the assemblages. Ostracod densities remain below 10 valves per increment below 40 cm core depth. They increase above 50 cm, reaching the highest density in the shelly lag, followed by a substantial decrease in density in the highstand silts (Fig. 3). The relative abundances show that C. neapolitana remains the dominant species throughout the whole core (Fig. 6). However, several taxa increase in relative abundance in the uppermost layers, including Loxoconcha sp., Cystacythereis sp. and Callistocythere adriatica. 3.2. Functional groups Filter feeders dominate the molluscan assemblages, while other feeding guilds, such as detritus feeders and carnivores, are rare (Table 2; Fig. 7a). When comparing the mean relative abundances of feeding guilds between the shelly lag and highstand silts, filter feeders and detritus feeders increase in the latter. In contrast, higher trophic levels, such as carnivores and scavengers, decrease (Table 2; Fig. 7a). However, it is the infaunal and semi-infaunal species that truly dominate the molluscan assemblages with a relative abundance of ≈ 80%. In comparison, epifaunal molluscs constitute ≈ 20% of the molluscan assemblages (Table 2; Fig. 7b). In the Rotaliida- dominated total foraminiferal assemblage (Fig. 7c–d; Table 3), epifaunal/infaunal and epiphytic/infaunal species are the most common. Similarly, the functional composition of ostracod assemblages remains relatively constant throughout the core and consists mainly of stenohaline species. Brackish and freshwater species are singletons and can be regarded as reworked specimens. All observed ostracod species belong to the subclass Podocopa and are benthic mobile detritus-feeders. 3.3. Species diversity The raw species richness, Shannon and Fisher diversity indi- ces of the molluscan assemblages increase up-core (Fig. 3). The molluscan species richness reaches 68 species in the shelly lag. It decreases up-core to 26 species in the highstand silts. The Hill-transformed Shannon diversity of the molluscan assemblages peaks in the shelly lag at 29.7 and decreases to 16.1 in the highstand silts, and Fisher’s index peaks with 22 at Table 3. Mean relative abundances of benthic foraminiferal orders and habitat preferences in the depositional sequences: highstand silts, 0 – 10 cm and the shelly lag 10 – 22.5 cm. All values are given in %. Orders Shelly lag Highstand silts Rotaliina 80 80 Textulariina 13 13 Miliolina 6 7 Habitat preferences Shelly lag Highstand silts Epifauna 8 6 Epiphytic 20 17 Infauna 3 12 Mixed preferences 69 65 Figure 4. Faunal densities and relative abundances of the most abundant species of the molluscan assemblages. G eologia C roatica 211Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... Figure 5. Faunal densities and relative abundances of the most abundant species of the benthic foraminiferal assemblages. Figure 6. Faunal densities and relative abundances of the most abundant species of the ostracod assemblages. G eo lo gi a C ro at ic a 212 Geologia Croatica 78/3 Figure 7. Feeding guilds a and habitat preferences b of the molluscan assemblages, and orders c and habitat preferences d of the benthic foramini feral assemblages. Each increment has been analysed for its species composition and displayed here as faunal densities (absolute number of individuals/ volume of the sample) and relative abundance (decimal). G eologia C roatica 213Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... 7.5 – 10 cm, and also decreases to 14.8 within the highstand silts. The maximum diversity of the total molluscan assem- blage is recorded within the shelly lag or slightly above, fol- lowed by a remarkable drop in the highstand silts (Fig. 3). Raw foraminiferal species richness ranges from 18 to 29 species in the shelly lag and from 21 to 29 in the highstand silts (Fig. 3). Shannon diversities of the foraminiferal assemblages increase from around 9 in the shelly lag to 13.5 in the high- stand silts (Fig. 3). Fisher’s index is slightly higher in the high- stand silts (ranging from 4.8 and 7.4) than in the shelly lag (range from 4.2 to 5.6) (Fig. 3). Although the foraminiferal densities decrease in the uppermost 10 cm, their species rich- ness and diversity do not decline significantly in the assem- blages of the highstand silts. Ostracod species richness ranges from 12 to 16 taxa in the shelly lag and from 11 to 14 taxa in the highstand silts (Fig. 3). Their Hill-transformed Shannon diversities range from 4.7 to 6.5 in the shelly lag, and decrease in the highstand silts to 3.5. The Fisher’s α of the ostracod assemblages fluctuates from 2.5 to 4 in the shelly lag, and ranges from 2.7 to 4.4 in the high- stand silts (Fig. 3). The species richness and diversity of the ostracod assemblages do not change significantly in the high- stand silts despite the drop in density. The rarefied molluscan diversity reveals a slight decrease in the highstand silts, but the diversity of benthic foraminiferal and ostracod assemblages shows no significant shifts within the record (Fig. 8). 3.4. Multivariate patterns in assemblage species composition The NMDS ordination shows an up-core shift in the mollus- can composition along the first NMDS axis, with a larger sep- aration between assemblages at the base of the core on one hand and assemblages from the shelly lag and the highstand silts on the other (Fig. 9a). The constrained clustering indicates that the highstand silts (0 – 10 cm, last 2,000 yrs BP) differ in the molluscan composition relative to the extended shelly lag (10 – 40 cm), while the assemblages in transgressive sediments (2,000 – 7,800 yrs BP) are highly variable in composition (es- pecially at 45 – 60 cm) and are less similar to those from the upper core increments (Fig. 9a). Similar to the molluscs, the NMDS ordination of the foraminiferal assemblage shows an up-core shift along the first NMDS axis and very high disper- sion among assemblages occurring in the lowermost incre- ments (50 – 75 cm, Fig. 9b). The NMDS ordination based on the ostracod assemblages reveals high similarity in the species composition among assemblages within the upper 30 cm of the core, with higher dispersion of assemblages at 30 – 40 cm (Fig. 9c). The Mantel test shows a positive Spearman correla- tion between molluscan and foraminiferal Bray-Curtis dis- similarities (rho = 0.55, p < 0.001), indicating that temporal changes in the composition of assemblages are shared by these two taxa, i.e., core increments with similar molluscan assem- blages are also similar in terms of foraminiferal assemblages. In contrast, there is a low and insignificant correlation between molluscan and ostracod dissimilarities (rho = 0.18, p = 0.24). 3.5. Stress indices The Marine Biotic Index (BORJA et al., 2000) for molluscs is 1.19 for the shelly lag (unpolluted/undisturbed status) and 1.44 for the highstand silts (slightly polluted status). The total fo- raminiferal assemblage comprises 44 sensitive, 15 stress-toler- ant, and 10 species of unknown stress tolerance. The number of stress-tolerant and sensitive species remains relatively constant Figure 8. Trends in rarefied species richness and Shannon diversity along the core. Shading indicates 95% confidence intervals. G eo lo gi a C ro at ic a 214 Geologia Croatica 78/3 throughout the core. However, the relative abundances of Non- ionella and Bulimina increase from the shelly lag to the high- stand silts. The stress-tolerant species include the most abun- dant genera of the total foraminiferal assemblage, Haynesina spp. and Ammonia spp. The Foram Stress Index (FSI, DIMIZA et al., 2016) exceeds a mean value of 3.8 in the shelly lag and Figure 9. Non-metric multidimensional scaling ordination and constrained clustering of the three taxa. a Marine molluscs; b Benthic foraminifera; c Marine ostracoda. All numbers indicate the increment midpoints (core depth) in cm. G eologia C roatica 215Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... 3.9 in the highstand silts. Both values indicate a moderately pol- luted environment. The enhanced Benthic Foraminifera Oxy- gen Index (eBFOI, KRANNER et al., 2022) exceeds a mean value of 68.9 in the shelly lag and 67.7 in the highstand. Both values indicate a well-oxygenated environment. 4. DISCUSSION 4.1. A stratigraphic record affected by condensa- tion, reworking and mixing The gravity core preserves a palaeoecological marine archive of ≈ 10,000 yrs, characterised by centennial and millennial time averaging, affected by slow sedimentation (condensation), re- working, and vertical mixing of shells. Varicorbula gibba shell ages span 300 years in the uppermost core increment. In con- trast, they range in age by several thousand years at the base of the highstand silt and in the shelly lag (mainly 2 – 6 kyr) (BE- RENSMEIER et al., 2023). Median ages of the shallow-subtidal bivalve V. gibba do not overlap between the highstand silt and the shelly lag (median age is 50 years at 0 – 2.5 cm, 1380 years at 5 – 7.5 cm, 2880 years at 10 – 12.5 cm, and 4480 years at 17.5 – 20 cm). Although the overlap in shell ages between the upper and the lower highstand silt increment is limited, age data dem- onstrate that the base of the highstand silts is already mixed with the shelly lag. Postmortem ages of the intertidal bivalve L. mediterraneum do not vary systematically within the upper 60 cm, with all shell ages ranging between 9380 and 10 510 cal years AD2017 (Fig. 2), indicating substantial post-depositional mixing of very old shells across the highstand silts and the shelly lag, induced either by bioturbation and/or by their trans- port from adjacent locations. However, this species is numeri- cally rare and does not significantly alter the overall molluscan composition (BERENSMEIER et al., 2023). To conclude, the shelly lag is characterised by millennial- scale time averaging. It thus integrates across molluscan, fo- raminiferal and ostracod communities that inhabited the sea- floor prior to the anthropogenic impacts in the northern Adriatic Sea (from ≈ 2 kyr BP up to ≈ 6 kyr BP). Although the lowermost increments of the overlying silts are still mixed with the shelly lag, the shell-poor, organic and metal-rich silts at the top of the core are less time-averaged and less affected by mix- ing with the underlying strata. In contrast, as the shelly lag does not contain shells from recent centuries, its fossil assem- blages with molluscs, benthic foraminifera, and ostracods can be representative of pre-impact conditions. However, the high- stand silts probably represent the mixture of pre- and post-im- pact conditions, and even the uppermost increment, averaging across the past ≈ 300 years, can be expected to integrate across distinct phases of the eutrophication history. Although the upward decline in macro- and microfaunal densities between the shelly lag and the highstand silts can re- flect a substantial decline in benthic production, a positive cor- relation between molluscan fossil density and sample size-in- dependent diversity is observed in this core (TOMAŠOVÝCH et al., 2024). This positive correlation indicates that the decline in fossil density reflects an increase in sedimentation rate over the past centuries and thus does not necessarily correspond to any temporal change in standing population densities of mol- luscs, benthic foraminifers or ostracods. Analyses of a total foraminiferal assemblage collected in a sediment core at a muddy site location in the Po Delta not far from our coring lo- cation also indicate that sedimentation rates increased during the 19th and 20th centuries (BARMAWIDJAJA et al., 1995). 4.2. The shelly lag as a compositional baseline As interpreted above, relative abundances of molluscs, benthic foraminifera, and ostracods in the shelly lag represent a time- averaged compositional baseline for assemblages inhabiting sub-tidal, muddy habitats in the northern Adriatic Sea. The molluscan, foraminiferal and ostracod assemblages in the shelly lag are dominated by opportunistic taxa such as V. gibba, Haynesina spp. and C. neapolitana, that typically tolerate or- ganic enrichment, pollution and some degree of oxygen deple- tion (Table 4). However, the relative abundances of Haynesina spp., and C. neapolitana decline in the overlying highstand silts. Moreover, shelly lag assemblages also include species that are relatively sensitive to pollution and organic enrichment, e.g. the molluscs T. tricarinata, Timoclea ovata, Parvicardium sca- brum, and Papillicardium papillosum (e.g., NERLOVIĆ et al., 2011; TOMAŠOVÝCH et al., 2018) and benthic foraminifera Neoconorbina terquemi, Asterigerinata mamilla or Elphidium crispum. The shell lag assemblage reflects a moderately diverse molluscan community (or set of communities) dominated by filter feeders and infaunal species preferring soft bottoms. The diversity values are similar to the other Holocene assemblages exhibiting millennial time averaging at other sub-tidal loca- tions affected by low sedimentation rates in the northern Adri- atic Sea (GALLMETZER et al., 2019; TOMAŠOVÝCH et al., 2020, 2024; PICONE et al., 2008; BARBIERI et al., 2019). The microfaunal assemblage within the shelly lag is characterised by stenohaline ostracods and a highly diverse shallow-water benthic foraminiferal fauna. Table 4. The dominant species of each total assemblage (molluscan, benthic foraminiferal, and ostracod) and their responses to environmental stressors. Total assemblage Dominant species Response to organic pollution Response to heavy metal pollution Response to oxygen depletion Used as index species Ecogroup Molluscs Varicorbula gibba Very tolerant (HRS-BRENKO, 2006) Tolerant (GALLMETZER et al., 2017) Tolerant (HRS-BRENKO, 2006) MORAITIS et al., 2018; TOMAŠOVÝCH et al., 2017, 2018 Species that are tolerant to organic matter enrichment (Group III after BORJA et al., 2000) Benthic foraminifera Haynesina spp. (mostly H. depressula and H. germanica) Tolerant (VIDOVIĆ et al., 2009; BERGIN et al., 2006; BARBIERI et al., 2019) Tolerant (BERGIN et al., 2006; ŽVAB ROŽIČ et al., 2022) Moderately tolerant (GLOCK, 2023) MELIS et al., 2019; ŽVAB ROŽIČ et al., 2022 Species that are indifferent to or relatively favoured by organic enrichment (Group II and III after JORISSEN et al., 2018) Ostracods Cytheridea neapolitana Tolerant (BARBIERI et al., 2019) Ostracods are, in general, sensitive RUIZ et al., 2005; SALVI et al., 2015 Tolerant (BARBIERI et al., 2019) BARBIERI et al., 2019 Opportunist G eo lo gi a C ro at ic a 216 Geologia Croatica 78/3 4.3. The human-impacted shell-poor silts The relative abundances of infaunal molluscs and benthic fo- raminifera increased in the highstand silts. In contrast, all other groups, particularly the epifaunal molluscs and epiphytic foraminifera, decreased from the shelly lag towards the top of the highstand silts. The foraminiferal assemblages also exhibit a significant increase in the relative abundance of infaunal spe- cies (such as Nonionella sp., Bulimina marginata, and Bulim- ina aculeata) within the muddy highstand silts relative to their abundance in the shelly lag. These taxa are considered to be opportunists by JORISSEN et al. (2018) and DIMIZA et al. (2016). In contrast, the relative abundances of epiphytic spe- cies decreased (such as Asterigerinata mamilla and Neoconor- bina terquemi). BARMAWIDJAJA et al. (1995) described a similar decrease in the relative abundance of epiphytic species close to the sampling area (see Fig. 1). Although V. gibba tol- erates hypoxia (HRS-BRENKO, 2006) and organic matter en- richment (Group III of the Marine Biotic Index, BORJA et al., 2000), shells of this species in the shelly lag are typically smaller than in the overlying highstand silt (BERENSMEIER et al., 2023) where they also increase in proportional abun- dance, indicating stronger eutrophication during deposition of the uppermost core increments compared to the shelly lag. Despite their low thickness and mixing at the base, the highstand silts are sedimentologically and geochemically dis- tinct from the underlying shelly lag. The highstand silts cor- respond to an increase in fine-grained sediment accumulation over the last ≈ 2,000 years, driven by the transition from wave- dominated to the river-dominated, rapidly prograding Po del- taic system with prodelta lobes reaching water depths of 30 m (AMOROSI et al., 2019), further enhanced by a shift of the Po River northwards to its present-day position ca. 800 years ago (CORREGGIARI et al., 2005). This change in sedimentary dynamics ≈ 2 kyr BP was at least partly caused by increased soil erosion and sediment discharge due to the intensification of deforestation and agriculture during the Roman Empire (MASELLI & TRINCARDI, 2013). The shift in habitat pref- erences between the shelly lag and highstand matches the ba- sin-wide trend of infaunalisation observed in live-dead studies (HASELMAIR et al., 2021) and other sediment cores (e.g., GALLMETZER et al., 2019), indicating a substantial change in soft-bottom environments in the northern Adriatic Sea. However, T. tricarinata, a species sensitive to pollution or sea- sonal oxygen depletion, even increases in relative abundance in the highstand silts. Monitoring studies focusing on living assemblages and stratigraphic records from non-condensed cores in the Po prodelta and Gulf of Trieste showed that T. tri- carinata declined, whereas V. gibba increased in abundance (and also in shell size) during the 20th century (station SJ101 in HRS-BRENKO, 2006; TOMAŠOVÝCH et al., 2017, 2018; MANARINI et al., 2019). The increase in Turitellinella’s rela- tive abundance can reflect conditions associated with earlier phases of eutrophication and Po prodelta progradation that were not yet associated with the negative consequences of sea- sonal oxygen depletion or pollution. Despite the stratigraphic condensation, the highstand silts preserve, to some extent, a community state that differs from the shelly lag assemblage based on the observed changes in community composition of all three taxonomic groups. 4.4. Discrepancy between molluscan and foraminiferal stress indices The Marine Biotic Index based on the molluscan assemblages (BORJA et al., 2000) indicates the transition from an unpol- luted environment recorded in the shelly lag to a slightly pol- luted environment recorded in the highstand stilts. Due to the insufficient percentage of foraminiferal species assigned to ecological categories based on sensitivity to organic carbon content (BORJA et al., 2000; BORJA & MUXIKA, 2005; ALVE et al., 2016), the Foram-AMBI index was not consid- ered for discussion. Therefore, the ecological status was inter- preted only by the FSI (DIMIZA et al., 2016). The values of the FSI (DIMIZA et al., 2016) indicate a slightly polluted en- vironment in both the shelly lag and the highstand silts. Al- though the stress indices based on molluscan and foramini feral assemblages differ to some degree, the associated changes in the relative abundances of species and functional groups in both taxa point to a partial degradation of the ecosystem sta- tus during the deposition of highstand silts relative to the dep- osition of the shelly lag. The discrepancy in molluscan and foraminiferal stress indices of the shelly lag could result from several factors, including (1) differing tolerances of macro- and microfauna to organic enrichment or pollution, with benthic foraminifera or ostracods considered more sensitive to pollu- tion than molluscs (YANKO et al., 2003; RUIZ et al., 2005; SALVI et al., 2015), (2) differences in the degree of time aver- aging because mixing processes are known to be size-selective (however, time averaging of benthic foraminifers and molluscs was documented to be similar at other locations in the north- ern Adriatic Sea, NAWROT et al., 2022), and/or (3) uncertain- ties related to the lack of species-level discrimination. How- ever, the Mantel test demonstrates the positive correlation between the composition of molluscan and foraminiferal as- semblages, suggesting that the living communities of these two taxa responded similarly to environmental changes and that their death assemblages were not subjected to different condensation and mixing despite their differing sizes and or- ganizational levels (single-celled versus multicellular organ- isms). Both molluscan and foraminiferal datasets differ in the number of unassigned species, with 26% foraminiferal taxa and 14% molluscan taxa not determined to species level. The Marine Biotic Index (BORJA et al., 2000) and Foram Stress Index (DIMIZA et al., 2016) are expected to be calculated with less than 1% unassigned species in the molluscan assemblages, and up to 10% unassigned species in the foraminiferal assem- blages. While the molluscan stress index is reliable in this re- gard, the foraminiferal stress index can be biased by the high proportion of individuals undetermined to species level. Even with this uncertainty, the overall shift in the AMBI index is congruent with the increasing relative abundances of pollu- tion-resistant foraminifera, such as Bulimina or Nonionella, even when the FSI does not show any significant change. Moreover, temporal changes in the relative abundances of functional groups align with the results of other palaeoeco- logical studies about benthic foraminifera in the northern Adriatic Sea (e.g., BARMAWIDJAJA et al., 1995; BARBIERI et al., 2019, 2021). G eologia C roatica 217Berensmeier et al.: Late Holocene changes in the composition of foraminiferal, ostracod and molluscan communities in condensed sediments ... The molluscan and foraminiferal fossil assemblages still record these ecological trends, even though they are preserved within the thin, time-averaged stratigraphic succession. 5. CONCLUSION The gravity core collected at ≈ 30 km east of the Po delta in 31 m water depth, in the northern Adriatic Sea, records the composition of the benthic communities that existed in the re- gion prior to human impact (shelly lag, 2 – 6 thousand years ago BP). Although the highstand silts deposited over the past two millennia average across pre- and post-impact community states, they show benthic and geochemical signatures of hu- man impact. To accurately interpret the palaeoecological data, it is essential to combine fossil and stratigraphic information. Even seemingly minor changes in sedimentation rates can sig- nificantly impact the temporal resolution of the fossil record. Although the sediment core examined here is characterised by condensation and mixing, which obscures ecological signals within the fossil assemblages, including molluscs, benthic fo- raminifera, and ostracods, the shelly lag deposited prior to the onset of human impacts remains largely segregated from the highstand silts that archive the most recent changes in the com- munity composition. Changes in species abundances and func- tional groups reflect infaunalisation that coincides with envi- ronmental changes reconstructed from geochemical proxies. Despite the incomplete nature of the fossil record, the assem- blages in the shelly lag thus offer insights into a baseline largely unaffected by human impacts. At the same time, the highstand silts, while still averaging out across the past cen- turies, show fingerprints of eutrophication and pollution, even when the magnitude of the impact is rather minor. 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