116 ACTA BOT. CROAT. 84 (2), 2025 Acta Bot. Croat. 84 (2), 116–140, 2025 CODEN: ABCRA 25 DOI: 10.37427/botcro-2025-024 ISSN 0365-0588 eISSN 1847-8476 Review article Marine benthic diatoms from the Adriatic Sea (NE Mediterranean): review of investigations and checklist with updated nomenclature Ana Car1*, Aydin Kaleli2 1 University of Dubrovnik, Institute for Marine and Coastal Research, Kneza Damjana Jude 12, HR-20000 Dubrovnik, Croatia 2 Istanbul University, Faculty of Aquatic Sciences, Department of Marine and Freshwater Resources Management, TR-34134 Istanbul, Türkiye Abstract – Despite more than 30 years of investigations, the knowledge of the composition and spatial distribution of marine benthic diatoms along both coasts of the Adriatic Sea is still limited. The distribution and a checklist of marine benthic diatom taxa in the Mediterranean sub-basin of the Adriatic Sea are presented based on literature data. The nomenclature is brought up to date and synonyms are included. Eight hundred and twenty-two species belonging to 70 families and 163 genera have been reported. The number of pennate species represented was 719. Families with the highest number of genera were: Naviculaceae (8), Bacillariaceae (7), Surirellaceae (6), Fragilariaceae (6). The highest number of taxa belonged to the Naviculaceae (95) and Bacillariaceae (88). The most numerous genera were Mastogloia (65 taxa), Navicula (62), Nitzschia (53), Amphora (52), Diploneis (43), Cocconeis (37), Halamphora (24), Achnanthes (22), Licmophora (22), Tryblionella (19), Fallacia (15), Surirella (14), Grammatophora (13), and Lyrella (12). This study focused on the areas of investigation according to the literature reviewed and showed that many areas have to be in- vestigated. Nevertheless, this first comprehensive checklist of marine benthic diatoms in the Adriatic Sea, updated with recent literature, is a valuable tool. However, it is important that this update has been progressive over time. Incor porating data on diatom taxa will improve the overall diversity of microalgae and our knowledge of the ecology of coastal ecosystems. Keywords: Adriatic Sea, Bacillariophyta, biogeography, epilithic, epipelic, epiphytic, Mediterranean Sea, taxonomy Microphytobenthos The microphytobenthos (MPB) colonises the surface sediment layer (2 to 3 mm thick) where strong chemical- physical gradients exist (MacIntyre et al. 1996). In temper- ate regions, the microphytobenthic community is mainly composed of benthic diatoms, which are either colonial or unicellular, either free-living or attached to a substrate through gelatinous protrusions and play an important role in primary production in marine ecosystems (Falkowski et al. 2004). Classification based on sediment typology reveals distinct groups: (i) epipelon, diatoms living on sediment, (ii) epipsammon, diatoms adhering to sand grains, and (iii) epiphyton, diatoms living on other photosynthetic organ- isms (Round et al. 1990). The microphytobenthic commu- nity plays a key ecological role within the aquatic ecosystem, regulating nutrient and oxygen fluxes at the water-sediment interface and contributing significantly to primary produc- tion (MacIntyre et al. 1996). Due to their ecophysiological characteristics, these microorganisms serve as valuable indi cators for water quality assessment and in paleoecologi- cal reconstructions (Stevenson and Pan 1999, Cibic and Blasutto 2011, Barinova et al. 2019, B-Béres et al. 2023). Microphytobenthic populations, particularly in shallow aquatic environments, are frequently enriched by phyto- planktonic species that settle on sediments in conditions devoid of turbulent movement, tidal currents, or water col- umn stratification. Conversely, under conditions of pro- nounced turbulence in the water column, microphytoben- thic organisms may be resuspended, infiltrating the phytoplankton community (Delgado et al. 1991, MacIntyre et al. 1996). Furthermore, identical species may be present both in the water column and in the surface sediments, making it difficult to characterize the typical species in each habitat ecologically (Cibic et al. 2022). In a two-year study * Corresponding author e-mail: ana.car@unidu.hr BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 117 (2003-2004) on MPB biodiversity conducted in Northern Adriatic Sea, on average, 9% of the species in the sediment were planktonic (Cibic et al. 2007a); however, they were pre- dominantly in poor condition. Only species with robust frustules, such as Pseudo-nitzschia seriata, were identifiable and likely still photosynthetically active (Cibic et al. 2007b). In recent years, the application of light microscopy (LM) and scanning electron microscopy (SEM) has enabled the discovery of numerous diatom species in marine coastal waters, greatly improving our understanding of diatom communities in different coastal regions, such as the Black Sea, the Mediterranean Sea or Yellow Sea (Park et al. 2018, Kaleli and Akçaalan 2021, Zidarova et al. 2022). While var- ious authors have previously compiled checklists (Hendey 1974, Viličić et al. 2002, Cibic and Facca 2010, Caraus 2017), the recent and progressing systematics of diatoms has re- sulted in many synonyms and the transfer of taxa to a new species of a genus (Li et al. 2018, Morales et al. 2019). A con- siderable proportion of the studies conducted has predom- inantly focused on planktonic forms of diatoms, thus the publications considered herein generally include both ben- thic and planktonic diatoms. Mediterranean Sea The Mediterranean Sea is a semi-enclosed continental sea that is almost entirely landlocked between Europe, Africa and Asia and consists of several sub-zones, including the Alboran Sea, the Balearic Sea, the Ligurian Sea, the Tyr- rhenian Sea, the Ionian Sea, the Adriatic Sea, the Aegean Sea and the Levantine Basin. It is a concentration basin that receives relatively low saline Atlantic Water (AW), which flows eastwards in the surface layer and gains salt due to the positive Evaporation-Precipitation balance over the Medi- terranean (Menna et al. 2022). The northeastern Mediter- ranean Sea is influenced by the brackish waters of the Black Sea through the Sea of Marmara. The shallowest part of the Mediterranean Sea is the northernmost third of the Adriatic Sea (i.e., the Northern Adriatic Sea), a shelf with depths up to 100 m (Vilibić et al. 2023). The Mediterranean Sea is con- sidered an oligotrophic basin with a few biological produc- tion and biodiversity hotspots (Myers et al. 2000). The meadows of Posidonia oceanica L. Delile, a species of sea- grass endemic to the Mediterranean Sea, are the most im- portant ecosystem in the Mediterranean in terms of biodi- versity. However, many alien and invasive species have recently been reported along the eastern coasts of the Med- iterranean and Aegean Sea, originating from the Red Sea through the Suez Canal (Zenetos et al. 2005). The Mediter- ranean Sea is an important basin, and its coasts have been heavily influenced by humans through tourism, urbaniza- tion and maritime transport. Additionally, the Mediterra- nean Sea is considered a hot spot of global warming, as it has been changing faster than the global ocean, with recent studies agreeing on increases in sea level, sea surface tem- perature and salinity over the past two decades, which have had a strong impact on the marine environment (Menna et al. 2022). As far as modern diatom studies in the Mediterranean Sea are concerned, one of the pioneering studies was pub- lished by Witkowski et al. (2000). While Witkowski et al. (2000) published a monograph of diatom taxa from various marine coastal sites worldwide, Blanco and Blanco’s mono- graph (2014) showed widespread taxa alongside site-specific taxa in the Mediterranean region. In addition, several checklists with a combination of planktonic species in the Mediterranean Sea have been published. Viličić et al. (2002) published a phytoplankton checklist for the eastern Adri- atic Sea, while Kaleli and Akçaalan (2021) showed the dia- tom flora of the Turkish coasts, Tas (2013) investigated the phytoplankton composition in the Aegean Sea (Tas 2013), and Velâsquez and Cruzado (1995) investigated the diatom flora of the northwestern Mediterranean Sea. Most of the studies used a similar sampling technique, specifically the use of a plankton net designed for phytoplankton sampling, with incidental observations of benthic diatoms. Conse- quently, the use of the plankton net resulted in the observa- tion of relatively large cell sizes for the most commonly cited taxa. In recent decades, the literature on the distribution of MPB species in the Mediterranean Sea has been limited to the coastal regions of Spain and France (Delgado 1989, Barranguet et al., 1996, Barranguet 1997, Riaux-Gobin et al. 1998). While these publications highlight the importance of MPB as a primary producer within coastal or lagoon eco- systems, and report the total MPB expressed as chlorophyll and pigment concentrations, they do not provide informa- tion on the taxonomic composition or present a comprehen- sive taxonomic list. In recent study, Pérez-Burillo et al. (2022) evaluated the advantages and limitations of LM and DNA metabarcoding for the identification of benthic diatom communities in shallow coastal environments of the Mediterranean Sea using biofilm samples from different substrates in the Ebro delta bays (Spain). The compilation of the historic records of marine and brackish diatom species and genera reported for the French coastal areas since 1888 to 2019, with information on the geographical locations and ecology (e.g. sampled substrate, habitat, growth forms) for most taxa by Ribeiro et al. (2022), pro- vides a great dataset for the benthic marine diatoms of French Mediterranean coast. Before the compilation of the MPB checklist for the Ital- ian seas (northern and western Middle Adriatic, and Ligu- rian Sea) by Cibic and Facca (2010), there were no time series data (no continuous observations at specific sites over an extended period with constant abundance) on the MPB in the Italian seas. The knowledge of MPB in Italian seas is low compared to longer-term studies on phytoplankton (Cibic and Facca 2010). The list of Cibic and Facca (2010) encom- passes taxa derived from natural sediment samples collect- ed from various locations, including the Gulf of Trieste ( Sdrigotti et al. 1999, Welker et al. 2002, Cibic et al. 2007a), the Venice Lagoon (Tolomio et al. 1999, 2002, Facca et al. CAR A., KALELI A. 118 ACTA BOT. CROAT. 84 (2), 2025 2002a, b, 2003, 2004, Tolomio 2004, Facca and Sfriso 2007), the Adriatic coast from Ancona to the Po Delta (Totti 2003), the Lesina Lagoon (Gambi et al. 2003) as well as epibiontic microalgae on marine hydroids Eudendrium racemosum in the Ligurian Sea (Romagnoli et al. 2007, 2014), epilithic di- atoms on different artificial hard substrates in the Conero area (Totti et al. 2007), and the first colonisation stages of diatoms on artificial hard substrates at two stations in the Gulf of Trieste (Bartole et al. 1991-94) and the Venice Lagoon (Tolomio and Andreoli 1989, Tolomio et al. 1991). Additionally, several recent papers provided benthic diatom checklists either in the main text or as Supplementary Ma- terial. These studies cover the Gulf of Trieste (Franzo et al. 2014, Rogelja et al. 2018) and the Grado Lagoon (Natali et al. 2023), the Adriatic coast from Ancona to the Po Delta (Accoroni et al. 2016, Cibic et al. 2019), and the Venice Lagoon (Baldassarre et al. 2023). Additionally, microben- thic community structure and trophic status of sediments of the Ionian Sea were studied in the period 2013–2014 ( Rubino et al. 2016). The only available information regard- ing the microphytobenthic community of the Ligurian Sea refers exclusively to epibiontic species associated with marine hydroids (Romagnoli et al. 2007, 2014), showing that diatoms living on this marine invertebrate derive an advan- tage from both the host and the environmental conditions due to the availability of nutrients and organic compounds (Cibic and Facca 2010). Diatom assemblages from the Tyrrhenian Sea have been used to study the effects of heavy metal contamination, where this group showed the presence of teratogenic forms in response to this kind of pollution (Rogelja et al. 2016). The epiphytic diatom communities of the endemic Med- iterranean seagrass P. oceanica are among the most thor- oughly studied (Mazzella et al. 1994, De Stefano et al. 2000). Mazzella et al. (1994) identified Cocconeis Ehrenberg species as the most abundant and frequent diatoms on the leaves in all seasons and throughout the depth range of the seagrass distribution. De Stefano et al. (2000) analysed eight species of Cocconeis on the leaves of P. oceanica using SEM on sam- ples collected around the island of Ischia (Gulf of Naples, Italy). Their results showed that certain Cocconeis species such as C. scutellum var. posidoniae M.De Stefano, D.Marino & L.Mazzella and C. neothumensis var. marina M.De Stefano, D.Marino & L.Mazzella exhibit high abundance, forming a continuous, almost monospecific layer on the colonised segments of the leaves (Mazzella et al. 1994). In a comparative study, De Stefano et al. (2008) provided addi- tional insights into C. scutellum Ehrenberg and its varieties and presented information on the geographical distribution of all the analysed C. scutellum taxa, including from the leaves of Posidonia in the Mediterranean region collected in spring 2005 and 2006 (the season when the density of epiphytic macroalgae is low) and covering almost the entire Mediter- ranean Sea (including sampling sites along the coasts of Spain, Italy, Slovenia, Croatia, Greece and Türkiye). A comprehensive review of the existing literature reveals that there is only a limited number of studies focused on the coastal waters of Türkiye (Kaleli and Akçaalan 2021). The Sea of Marmara and the Aegean Sea were the most inten- sively studied areas, with 20 and 19 studies, respectively. Studies on benthic and planktonic species composition and biogeographic distribution in the Mediterranean Sea and the Black Sea were rather sparse with only 11 and 14 stud- ies, respectively (Kaleli and Akçaalan 2021). On the Turkish coast of the Aegean Sea, Altuğ et al. (2011) and Aslan et al. (2018) investigated the composition of phytoplankton in the northeastern Aegean Sea. Koray (2001) published a check- list of phytoplankton in the Turkish seas. These studies fo- cused mainly on the planktonic forms of diatoms. Several studies have also been carried out on the benthic diatoms of the shores of Greece (Economou-Amilli 1980, Belegratis 2002, Belegratis and Economou-Amilli 2002, Louvrou 2007). Foged (1985a, b) published monographs on the dia- tom communities of the islands of Samos, Kos and Kalym- nos. In the southern Aegean Sea, periphyton colonisation in hydrothermal marine areas of the island of Milos has been studied (Louvrou 2007). The author later described Detonia dobrinae Louvrou, Danielidis & Economou-Amilli (Louvrou et al. 2006) and several taxa of Meloneis I.Louvrou, D.B.Danielidis & A.Economou-Amilli (Louvrou et al. 2012). Loir (2010-2014) published micrographs of diatoms from the Mediterranean Sea, including on the island of Crete. Tas (2013) observed that diatoms and dinoflagellates were the dominant groups of phytoplankton groups on the Datça peninsula. Cocconeis and Amphicocconeis M.De Stefano & D.Marino have been studied extensively in the Aegean Sea (De Stefano and Marino 2003, De Stefano et al. 2006, 2008, Majewska et al. 2014). These studies introduced several new species and expanded knowledge of these genera using both LM and SEM. Recently, Konucu and Develi (2021) reported two new monoraphid diatom species, Cocconeis sigillata Riaux-Gobin et Al-Handal and Amphicocconeis rodriguensis Riaux-Gobin et Al-Handal, from samples collected in the northeastern Mediterranean Sea. Morphological studies by Pennesi et al. (2011, 2012) in- vestigated the ultrastructure of some marine Mastogloia Thwaites ex W.Smith species collected during opportunis- tic sampling of benthic diatoms from diverse geographical regions, including tropical, subtropical including Indonesia and the Red Sea, and temperate zones, such as Patmos Is- land in the Aegean Sea, Greece, where epiphytic diatom samples were collected from seagrasses and macroalgae in September 2000. The genus Mastogloia has been reported in Turkish temperate inland waters, exhibiting a presence in both pelagic and benthic diatom flora (Elmacı and Obalı 1998, Akbulut and Yıldız 2002, Çelekli and Külköylüoğlu 2006, Sıvacı et al. 2007, Çolak Sabancı 2013). Recently, several papers have been published on diatom communities in the Aegean Sea; Li et al. (2018) described Gedaniella flavovirens (H.Takano) Chunlian Li, A.Witkowski & M.P. Ashworth in the Fragilariaceae, and several taxa were introduced from the Aegean Turkish coasts in the Iztuzu coast of Muğla (Kaleli et al. 2020). Additionally, diatoms from Iztuzu beach and Iztuzu coastal lake (Kaleli 2019, Kaleli 2022), as well as BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 119 the benthic diatom composition in Homa Lagoon (Izmir) (Çolak Sabancı 2010, 2011, 2012a, b, 2013, Çolak Sabancı and Koray 2010), have been studied. Some of these studies introduced new records for the Turkish flora, while others documented diverse compositions. In the African Mediterranean region, on the other hand, there have been very few studies. Aleem (1950) contributed new taxa from the Levantine Basin of Egypt. Voigt (1963) worked on Mastogloia species in the Mediterranean. Zalat (2001, 2002) and Zalat et al. (2019) investigated the commu- nity structure of the Suez Canal and Egyptian coasts, de- scribing new species and showing their spatial distribution, focussing on habitat preferences and life forms. Their stud- ies revealed that most species are cosmopolitan, with a sig- nificant presence of freshwater forms. More recently, a study on the distribution of diatoms on the Algerian coasts has been published (Kaddeche et al. 2022). Hydrographic and oceanographic characteristics of the Adriatic Sea The Adriatic Sea, the northernmost embayment of the Mediterranean Sea, 800 km long and 150 km wide, divided into three sections (Northern, Middle and Southern), is un- der considerable continental influence and characterised by diverse water masses (Viličić et al. 2002, Vilibić et al. 2023, Verri et al. 2024). Water exchange with the Mediterranean Sea takes place through the 800 m deep Strait of Otranto, with significant inflows into the Southern Adriatic Sea in winter, driven by the "Bimodal Oscillating System" (BiOS). This internal mechanism influences the biogeochemistry and biology of both the Adriatic and Ionian Seas (Civitarese et al. 2010, 2023, Ljubimir et al. 2017). The BiOS connects the deep thermohaline cell of the eastern Mediterranean, which originates in the Southern Adriatic, with the North- ern Ionian Gyre (NIG) and causes decadal shifts in circula- tion and redistribution of salinity (Civitarese et al. 2023). The eastern Adriatic coast is influenced by high-salinity and nutrient-poor inflows from the Ionian Sea and freshwater discharges from oligotrophic karstic rivers (Batistić et al. 2014, Ljubimir et al. 2017). Saline waters enter the Adriatic Sea in intermediate layers on the eastern side of the strait, while the Adriatic water mainly exits via the western shelf and the deep section of the strait (Gačić et al. 1996, Vilibić et al. 2023). The nutrient levels in the Ionian Sea and the Adriatic Sea show similar variability, although the Adriatic Sea generally has a lower nutrient content. By contrast, an- ticyclonic NIG enhances the inflow of fresher, nutrient-rich Modified Atlantic Water (MAW) originating in the Western Mediterranean, causing nutricline upwelling at the NIG border, decreasing Southern Adriatic temperature, salinity, and density, and reducing winter convection, thereby in- creasing ecosystem productivity (Civitarese et al. 2010). An overview of productivity across the geographical regions of the Adriatic Sea shows that the Northern Adriatic, particu- larly the Gulf of Trieste, Po River delta, and Rovinj area, has been the focus of most studies on primary production (N=46). The Middle Adriatic has been notable for in situ primary production measurements since the 1960s. Annu- al primary production ranges from 87.4–260.0 g C m–² y–¹ in the Northern Adriatic and 70.0–177.4 g C m–² y–¹ in the Middle Adriatic. The Southern Adriatic is the least studied, with only daily estimates available (236–374 mg C m–² d–¹) (Matek and Ljubešić 2024). The occurrence of organisms from the Atlantic/Western Mediterranean and the Eastern Mediterranean/temperate zones in the Adriatic correlates with the decadal circulation changes (the anticyclonic and cyclonic circulations) of the NIG (Civitarese et al. 2010, 2023). Review of diatom investigations in the Adriatic Sea The study of diatoms in the Adriatic Sea began in the late 1800s and several species were included in the diatom flora like Bacillaria adriatica Lobarzewski (Lobarzewski 1840). One of the pioneering studies in the Adriatic region was conducted by Kützing, who described Berkeleya adriatica Kützing (Kützing 1844) and Amphitetras adriatica Kützing (Kützing 1845). This was followed by the work of Grunow (1860), Cleve and Grunow (1880), and Van Heurck (1881), who discovered additional new species. In the 1900s, Mastogloia adriatica Voigt was described by Voigt (1963), and these records were followed by local studies in the 2000s. In recent decades, several authors have published stud- ies on phytoplankton composition, such as Viličić et al. (2002) with a checklist of phytoplankton in the eastern Adriatic Sea, Viličić (2014) with a book on the ecology and composition of phytoplankton in the Adriatic, Neri et al. (2024) with a comparative analysis of phytoplankton diver- sity in the Northern Adriatic Sea using microscopy and me- tabarcoding etc. According to Viličić et al. (2002), marine diatoms, comprising 518 species (330 pennates and 174 cen- tric), were the predominant phytoplankton group in the eastern Adriatic. The study highlighted a regional distribu- tion, with pennate diatoms more abundant in the Northern Adriatic and centric diatoms dominating the offshore Southern Adriatic. In the Adriatic Sea, benthic diatoms have been reported from several areas, including: the Gulf of Trieste (Bartole et al. 1991-94, Sdrigotti et al. 1999, Munda 2005, Cibic et al. 2007a, b, Cibic and Blasutto 2011, Franzo et al. 2014, Rogelja et al. 2018), Venice Lagoon (Tolomio and Andreoli 1989, Tolomio et al. 1999, Facca et al. 2002a, b, 2003, 2004, Tolomio et al. 2002, Facca and Sfriso 2007, Baldassarre et al. 2023), the northwestern Adriatic coast (Totti 2003, Totti et al. 2007, Franzo et al. 2015, Accoroni et al. 2016, Pennesi and Danovaro 2017, Cibic et al. 2019, Natali et al. 2023) and the eastern Adriatic coast (Burić et al. 2004, Miho and Witkowski 2005, Caput Mihalić et al. 2008, Levkov et al. 2010, Car et al. 2012, 2019a, b, 2020, 2021, Mejdandžić et al. 2015, Nenadović et al. 2015, Hafner et al. 2018a, b, Kanjer et al. 2019, Seveno et al. 2023, 2024). CAR A., KALELI A. 120 ACTA BOT. CROAT. 84 (2), 2025 The Northern Adriatic Sea, the most intensively studied area (e.g. Sdrigotti et al. 1999, Totti 2003, Munda 2005, Totti et al. 2007, Facca and Sfriso 2007, Mejdandžić et al. 2015), is a shallow sub-basin of the Adriatic Sea characterized by a morphologically complex coastline leading to the forma- tion of variable hydrodynamic and sedimentary environ- ments. The hydrology in this region is influenced by various factors such as winds and river discharge, and the general circulation is cyclonic. Freshwater (mean flow of 1496 m3 s-1 in the period 1917-2008, Cozzi and Giani 2011) originates from major rivers along the northern and northwestern coasts, with the Po River contributing nutrient-rich water, constituting approximately one-third of the total freshwater input into the Adriatic Sea. The Po's freshwater flow, char- acterized by relatively fresh and mesotrophic water, might form a thin surface layer over the northern sub-basin in summer, while it is reduced in winter and flows directly south along the Italian coast (Poulain 2001 and references therein). An analysis of the most recent decade (2013–2022) shows a pronounced negative trend across the entire Adri- atic Sea, indicating a consistent annual reduction of −4.2% in freshwater input throughout the river basin (Aragão et al. 2024). The sedimentation pattern is consistent with the hydrodynamic circulation (Ravaioli et al. 2003) and shows a narrow strip of recent sand along the coast, followed by a broad belt of muddy sediments. Offshore, there is a muddy transition zone characterized by a gradual increase in sand content, dominating a wide-open sandy shelf area with minimal recent sedimentation (known as relict sands) (Franzo et al. 2015 and references therein). Although the Northern Adriatic shelf is a relatively low-energy environ- ment with low tidal ranges and wave heights, ephemeral de- position of sediments occurs after high-tide events and is subsequently remobilized by waves in dense currents (Traykovski et al. 2007). Sediments migrate southward in a series of wind-induced resuspension events promoted by the Bora and Scirocco winds (Fain et al. 2007). The main sourc- es of biogenic elements are both autochthonous (plankton) and allochthonous (atmospheric inputs and soil organic matter transported by rivers) factors (Franzo et al. 2015). In line with the findings of Totti (2003), Franzo et al. (2015) also reported the presence of the centric diatom Paralia sulcata (Ehrenberg) Cleve in sediments off the coast of Emilia- - Romagna (Northern Adriatic) in a study aimed at improv- ing the understanding of offshore benthic communities in the Adriatic Sea. Cibic et al. (2012) studied benthic diatom community dynamics in relation to temperature, salinity, nutrient concentrations, freshwater inflow, and mucilage in the Gulf of Trieste (Northern Adriatic Sea, Italy) over seven years (1999–2005) at two sublittoral stations. They found that Nitzschia and Navicula showed a positive trend with increasing temperature, while Pleurosigma exhibited a neg- ative trend. Navicula and Nitzschia appeared to be negative- ly affected by mucilage events in the summers of 2000 and 2004, while Diploneis occupied the ecological niche tempo- rarily vacated by Navicula and Nitzschia. Diploneis had a negative relationship with temperature at the shallower site. Although no significant correlations between bin-averaged salinity and diatom abundance were found, Cylindrotheca was observed at high salinity levels. Despite the general sea- sonal variability observed in benthic diatom communities, characterized by distinct summer and winter assemblages (Cibic et al. 2012), Franzo et al. (2015) did not observe sig- nificant temporal variability in microalgal composition. In the Venice Lagoon, benthic diatoms from the surface sediment layer were studied to investigate possible relation- ships between epiphytic diatoms and water quality in shal- low coastal areas characterised by marked physical and chemical gradients and significant anthropogenic impacts (Facca and Sfriso 2007). In addition, Cibic and Blasutto (2011) investigated the response of diatoms to different nu- trient concentrations under nutrient-rich conditions in a benthic diatom study at three sublittoral sites in the Gulf of Trieste (Italy). It can be observed that the dominance of a single species leads to a decline in diversity. Under oligotro- phic conditions, without providing a competitive advantage for a single species when diatoms faced limitations by more than one nutrient, diversity remained high (Cibic and Blasutto, 2011). This study suggests that benthic diatoms, like macrobenthos, can serve as useful indicators of nutrient enrichment and represent a potential and innovative tool for biomonitoring. The most interesting environments along the Middle Adriatic are highly stratified estuaries formed by several small karstic rivers (Zrmanja, Krka), which are maintained by sufficient river discharge and low tides (Viličić et al. 2002). In the oligotrophic and highly stratified Zrmanja es- tuary, the composition and abundance of diatoms in biofilm formed on artificial substrates exposed at different depths were studied in July 2000 (Burić et al. 2004, Caput Mihalić et al. 2008). The study conducted by Caput Mihalić et al. (2008) focused on the composition and abundance of dia- toms within biofilms on artificial substrates exposed at dif- ferent depths in the Zrmanja estuary in July 2000. The pre- dominant periphyton species identified were Amphora coffeaeformis (C.Agardh) Kützing and Navicula veneta Kützing. Shannon-Wiener index ranged from 0.87 to 2.08 at the upper estuary station and from 1.1 to 2.7 at the other station. Additionally, Burić et al. (2004) investigated the abundance of periphytic pennate diatoms, in particular Cocconeis scutellum, attached to artificial substrates in the karstic Zrmanja estuary in the summer of 2000. The authors focused on comparing the abundance of attached Cocconeis cells with those suspended in the planktonic phase. Previous research focused mainly on the Northern Adriatic and estuaries (e.g. Burić et al. 2004, Caput Mihalić et al. 2008, Levkov et al. 2010), resulting in a significant knowledge gap regarding the diatom flora of the Southern Adriatic, especially before 2008. Although part of the mate- rial for the taxonomic study by Levkov et al. (2010) came from the Ombla River estuary near Dubrovnik (the South- ern Adriatic), it focused mainly on 15 species of the genus Rhoicosphenia based on LM and SEM. A literature review revealed not only a lack of studies in the coastal waters of BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 121 the Southern Adriatic before 2008, but also the absence of continuous long-term observations of benthic diatoms at a consistent location. There was also a shortage of informa- tion on the seasonal distribution and succession of epiphyt- ic diatoms on macroalgae. The most comprehensive study of diatom community structure in the Southern Adriatic was conducted between autumn 2008 and autumn 2010 in areas inhabited by invasive Caulerpa species (Car et al. 2012, 2019a, b). Prior to the studies by Car et al. (2012, 2019a, b), there were no studies on diatom communities associated with the marine green macroalga Caulerpa taxifolia (M.Vahl) C.Agardh (“killer seaweed", Bryopsidales, Chlorophyta), which outcompetes native seaweeds and seagrasses in the Mediterranean Sea by forming dense carpets, leading to biodiversity loss. The study by Kanjer et al. (2019) provided valuable insights into the biodiversity of threatened P. oceanica meadows, affected by human activities and invasive species, while also providing crucial information for the identification of taxa-specific epiphytic diatom communi- ties in the eastern Adriatic seagrass meadows. This study describes the epiphytic diatom community living on the blades of the P. oceanica in the coastal region of the eastern Adriatic Sea (Dugi Otok island). LM analysis of 21 samples revealed a diverse epiphytic diatom community with 68 taxa belonging to 30 genera across all depths (43 from 10 m depth, 41 from 15 m depth and 39 from 20 m depth), dom- inated by the genus Cocconeis, which is typical of the epi- phytic growth on the leaves of P. oceanica. All identified species are common in other marine benthic periphytic and epiphytic habitats. Although there was no positional influ- ence on community structure, ANOSIM tests (P < 0.05) in- dicated significant depth-related differences. In contrast to the study of Kanjer et al. (2019), the study by Car et al. (2019a) focused on the taxonomy of epiphytic diatoms associated with invasive macroalgae of the genus Caulerpa. These macroalgae are characterised by secondary metabolites such as caulerpenyne (CYN), which act as chemical defence mechanisms against herbivores and epi- phytes (Box et al. 2008, 2010). It is interesting to note that the CYN content varies between the different Caulerpa spe- cies, e.g. Caulerpa prolifera has higher CYN concentrations than invasive Caulerpa species under similar climatic con- ditions (Box et al. 2010). Epiphytic diatoms were studied in the summer and autumn of 2010 on the eastern Adriatic coast (Hvar Island) on fronds of C. taxifolia and, for com- parison, on the autochthonous macroalgae Padina sp. and Halimeda tuna (J.Ellis & Solander) J.V.Lamouroux. Quali- tative analysis was performed using LM and SEM. The Shannon-Wiener diversity index determined for C. taxifolia showed a wide range of values (3.11-4.88), with a maximum in August and a minimum in October. While the number of taxa on the fronds of C. taxifolia increased from June (41) to August (88), it decreased in autumn due to the high rela- tive abundance of Cocconeis caulerpacola Witkowski, Car and Dobosz, a diatom typical of Caulerpa. On the other hand, the highest number of taxa on Padina sp. was ob- served in September (82). The detailed composition of the epiphytic diatoms and the seasonal dynamics in the area af- fected by the invasive macroalga Caulerpa taxifolia were determined for the first time. In a study by Car et al. (2012), a new diatom species, C. caulerpacola, was observed on the green alga C. taxifolia over a wide geographical range; from the Adriatic (Stari Grad Bay, Hvar Island, Middle Adriatic, Croatia), on the Mediterranean coasts (Saint Raphaël, west of Cannes, France), and on the east coast of Australia (Moreton Bay, southeast Queensland). It has also been observed on Caulerpa cylindracea Sonder (syn. C. racemosa var. cylindracea (Sonder) Verlaque, Huisman & Boudouresque), another Mediterranean invasive alga. LM, SEM and TEM (transmis- sion electron microscopy) were used for morphological and fine structure analyses. The new species was compared with closely related species (C. borbonica, C. diruptoides and C. pseudodiruptoides). Surprisingly, C. caulerpacola was com- monly found on C. taxifolia despite its reputation as a 'killer seaweed' in very high abundance, but its occurrence seems to be very patchy. Indeed, C. taxifolia seems to be a suitable host for epiphytic diatoms, especially for the tiny Cocconeis caulerpacola. Furthermore, a study by Car et al. (2019b) compared the epilithic diatom community structures at three sites on the eastern Adriatic coast (Croatia) in the presence of two differ- ent invasive Caulerpa species, C. taxifolia and C. cylindracea, which have spread rapidly in the Mediterranean in recent decades. Between November 2008 and October 2010, 40 samples were collected seasonally at three sites (Hvar Island, Mljet Island and Dubrovnik). Qualitative analyses using LM and EM identified 310 epilithic taxa from 65 genera. The predominant genera were Mastogloia (48), Amphora (32), Diploneis (24), Nitzschia (23), Navicula (22) and Cocconeis (19). A similar seasonal shift in growth forms was observed at all sampling sites, with a doubling (Hvar, Mljet) or tripling (Dubrovnik) of the number of erect diatoms in spring. Apart from erect forms, Dubrovnik and Mljet were charac- terised by adnate forms, while Hvar was characterised by tube-dwelling forms. The highest values of the Shannon- Wiener diversity index occurred in autumn and ranged from 5.26 to 5.34. Significant differences in diatom commu- nities among sampling sites were correlated with the presence of invasive macroalgae (Hvar – C. taxifolia; Mljet and Dubrovnik - C. cylindracea). The main taxa contributing to the community variance were Cocconeis scutellum var. scutellum, Rhopalodia pacifica Krammer, Navicula ramosissima (C.Agardh) Cleve, and Berkeleya rutilans (Trentepohl ex Roth) Grunow. While the spatial variation may reflect unmeasured environmental factors, the results suggest the possible influence of invasive Caulerpa algae affecting the habitat through competition with native algae. This study improves the understanding of diatom diversity in chal- lenging environments at both regional and broader scales. A diatom community similar to the epilithic diatom in a study of Car et al. (2019b) has also been reported for vari- ous hard substrates in the Adriatic Sea (Munda 2005, Totti CAR A., KALELI A. 122 ACTA BOT. CROAT. 84 (2), 2025 et al. 2007). In a study examining seasonal fouling of arti- ficial substrates by diatoms in the heavily polluted and eu- trophic area near Piran in the Gulf of Trieste, with concrete plates (50 × 50 cm) used to observe the changes in diatom assemblages, Munda (2005) reported that the genera Berkeleya, Navicula and Licmophora dominated and covered most of the experimental surfaces. Achnanthes taxa were among the initial colonizers, while those belonging to Nitzschia and some epipelic taxa settled in autumn. Regarding the depth distribution, the highest taxa richness was observed in spring at 3 m depth and in autumn at 7 m depth (Munda 2005). Additionally, the community structure of epilithic dia- toms reported in Car et al. (2019b) study is in agreement with the results from the Northern Adriatic Sea (Ancona region) in the study by Totti et al. (2007), which was con- ducted on artificial hard substrates (marble, quartzite and slate) to evaluate substrate-dependent differences in coloni- sation and to characterise the seasonal variation of micro- epilithic communities in terms of abundance, biomass and community structure from April 2003 to April 2004 on a seasonal basis, and showed a dominance of motile taxa ( Navicula spp., Nitzchia spp.), followed by erect (Grammatophora marina, G. oceanica,), adnate (Halamphora coffeiformis and C. scutellum) and tube-dwelling diatoms (B. rutilans). How- ever, no significant differences in abundance and biomass were observed in the three substrates. Although Totti et al. (2007) reported high seasonal variability in epilithic micro- algal communities, with lower cell densities in winter, the Car et al. (2019b) data suggested that the variation in com- munity structure in terms of relative abundances of diatom taxa was related to sampling sites and possibly to the pres- ence of the two invasive macroalgae considered, rather than to seasonal effects. In addition, studies on benthic diatoms in Neum Bay (Bosnia and Herzegovina) in 2010 and 2011 (Hafner et al. 2018a, b) contributed significantly to the knowledge of dia- toms in the southeastern Adriatic. The study by Hafner et al. (2018a) aimed to expand the knowledge of the MPB in the Adriatic Sea by investigating, for the first time the ben- thic diatom communities in the small semi-enclosed oligo- trophic Neum Bay (Bosnia and Herzegovina), where benthic diatoms growing naturally on rocks and macroalgae were identified at a single station in 2010 and 2011. A total of 24 samples were taken at two different depths, 0.5 m and 8 m, and analysed by LM and SEM. A total of 425 pennate and 58 centric taxa (species and infraspecific taxa) belonging to 60 families and 115 genera were identified. The genera with the largest number of taxa were: Mastogloia (46 taxa), Navicula (36), Diploneis (35), Nitzschia (34), Amphora (31), Cocconeis (27), Achnanthes (14), Halamphora (12), Lyrella (11), and Surirella and Licmophora (10 each). Amphora bigibba var. interrupta and Cocconeis scutellum were the most common taxa, occurring in 87.5% of the samples. A total of 142 taxa were found only once (sporadic taxa). Although the benthic diatom richness in the bay was high, the taxa were not evenly distributed in time. Consistent quantitative and qualitative data are still needed to better determine the seasonal and spatial changes of the epilithic assemblages in the region. Two recent studies were conducted to understand the development of diatom communities in relation to physico- chemical parameters (Hafner et al. 2018b, Car et al. 2020). In the study by Hafner et al. (2018b) in Neum Bay (Bosnia and Herzegovina), the taxonomic composition and struc- ture of a marine epilithic diatom community was sampled from the bottom of the two sites at monthly intervals from January to December 2011. While the complete list of spe- cies and infraspecific taxa of benthic diatoms in Neum Bay from 2010 and 2011 (Hafner et al. 2018a) included 483 dia- tom taxa within 115 genera, the list of marine benthic dia- toms found in Neum Bay in 2011 included 264 taxa (species and infraspecific taxa) belonging to 69 genera (Hafner et al. 2018b). Of these, 149 and 203 taxa occurred in the samples from shallow (0.5 m depth) and deep (8 m depth) sites, re- spectively. The monthly distribution of most diatoms was irregular, and many sporadic taxa were found. The differ- ence between the shallow and the deep sites was mainly due to the diatom taxa that were frequently found and whose percentage abundance was high. These were Halamphora coffeiformis, Caloneis excentrica, Cocconeis scutellum var. scutellum, Licmophora f labellata, Licmophora gracilis, Licmophora sp., Navicula abunda, Rhabdonema adriaticum, and Striatella unipunctata. Canonical correspondence anal- ysis (CCA) showed that temperature, oxygen saturation (O2/O2'), silicate concentration (SiO4) and salinity were the most important factors influencing diatom community structure in the bay. While recent studies have focused on benthic diatoms from natural substrates in the Middle and Southern Adri- atic (Car et al, 2012, 2019a, b, 2021; Hafner et al. 2018a, b, Kanjer et al. 2019, Seveno et al. 2024), some earlier studies on benthic diatoms in the Adriatic Sea were conducted on artificial substrates (e.g. Tolomio and Andreoli 1989, Tolomio et al. 1991, Bartole et al. 1991-1994, Burić et al. 2004, Munda 2005, Totti et al. 2007, Caput Mihalić et al. 2008, Mejdandžić et al. 2015). Mejdandžić et al. (2015) investigated biofilm formation on an artificial substrate made of Plexiglas (poly- mer of methyl methacrylate) in the Northern Adriatic Sea (Rovinj). The study, conducted in autumn 2013, at a depth of 5 m above the sea floor, analyzed diatom and bacterial succession over a one-month exposure. Samples for algo- logical analysis were taken after 1 hour, half a day, a day or a few days, a week to one month. Selective agar plates, epifluorescence, LM, electron microscopy (EM), and high- performance liquid chromatography (HPLC) were used for pigment analysis. During the exposure, all biofilm compo- nents increased in abundance; the bacterial community was dominated by heterotrophic marine bacteria with 1.96 ± 0.79 × 104 colony forming units (CFU) cm–2, the photo- trophic component was dominated by diatoms (6.10 × 105 cells cm–2), and fucoxanthin was the dominant pigment (up to 110 ng cm–2). The diatom community, led by Cylindrotheca closterium (Ehrenberg) Reimann & J.C.Lewin and other BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 123 pennate benthic diatoms, comprised 30 different taxa, as revealed by detailed EM analysis. The study confirmed that a Plexiglas surface in a marine environment was susceptible to biofouling within 30 days of contact. Bacteria and cya- nobacteria were initially involved in the colonisation pro- cesses, followed by diatoms, which formed a primary bio- film in the sea. The use of artificial substrates in diatom studies offers advantages such as reduced effort and cost for sampling and processing, less habitat disturbance and higher sampling accuracy (Lamberti and Resh 1985, Lane et al. 2003). The main advantage of using artificial substrates over sampling natural habitats is the consistent standardisation between replicates. In addition, the use of artificial substrates for monitoring purposes has no impact on algal colonisation and allows global applicability without restrictions due to the natural life cycle and distribution range of macroalgae (Carreira-Flores et al. 2020, Natali et al. 2023). Although ar- tificial substrates have been used for diatom studies for al- most 100 years (Naumann 1915; cited in Tuchman and Stevenson 1980, Hoagland et al. 1986, Barbiero 2000), there is still concern about whether diatom communities that develop on artificial substrates accurately represent communities that develop on natural substrates (Lane et al. 2003). Ideally, artificial substrates should have community composition and abundance representative of natural substrates at the same site (Tuchman and Stevenson 1980, Lamberti and Resh 1985, Lane et al. 2003). But usually, the colonization patterns of artificial substrates differ from those observed on natural substrates (Mejdandžić et al. 2015). For example, diatom communities that develop on artificial substrates may better represent the diatom community of a particular natural substrate (Lane et al. 2003). Therefore, further comparative studies of diatom community structure on different artificial and natural substrates are needed. Since 2015, only two studies, conducted by Nenadović et al. (2015) in the Middle Adriatic, and Car et al. (2021) in the Southern Adriatic, have explored the differences in com- position between artificial and natural substrates. The study by Nenadović et al. (2015) focused on the initial coloniza- tion of diatoms in the periphytic community on various submerged artificial substrates, characterized by diverse physico-chemical properties, was situated in the coastal re- gion of the Middle Adriatic Sea (Zadar region). The study analysed the abundance and genus composition of diatoms in newly formed biofilms. Additionally, comparisons were made with the diatom community on a natural substrate, specifically the seagrass P. oceanica sampled from a nearby meadow, to evaluate differences in abundance and compo- sition on artificial substrates compared to an already estab- lished community on a natural substrate. This study aimed to determine the differences in abundance and composition of the diatom community on diverse artificial substrates and to discuss these differences by analysing the affinity of diatoms for specific artificial materials. Comparative inves- tigations indicate that, while living organisms (macro- phytes) and organic materials (wood, leaves) serve as supple- mentary nutrient sources for attached communities, the introduction of newly deployed inorganic artificial sub- strates (such as glass and plastic) into marine environments provides an opportunity to study the initial development and the succession of diatoms in periphyton communities (Nenadović et al. 2015). Furthermore, although Mejdandžić et al. (2015) and Nenadović et al. (2015) studied the devel- opment of periphytic diatoms on various artificial sub- strates (plexiglass, asbestos, painted iron, wood, concrete, glass, plastic, etc.), their results referred mostly to the ge- neric level. The relevance of studying diatom affinity as an important fouling community on different artificial mate- rials is to gain a comprehensive understanding of the impact of marine debris. This understanding is essential for sys- tematic work on alleviating the negative impacts of litter on the marine environment. In addition, to understand the differences in composi- tion on artificial and natural substrates, three habitats (epi- lithon, epiphyton and artificial substrate) were compared in a recent diatom study by Car et al. (2021). Weekly samples were taken in August and September 2016 from the marine lake Mrtvo More in the Southern Adriatic. The study in- volved detailed LM and, for the first time, SEM analyses of benthic diatoms from a marine lake. A total of 97 taxa with- in 42 genera were identified, with Cocconeis scutellum and Halamphora coffeiformis being the most frequent. Shannon- Wiener diversity index (H') values varied from 1.78 to 4.52 throughout the year. Two groups were recognized: (1) epilithon and artificial glass substrate, (2) macroalgae. The analysis indicated that diatom communities on artificial substrates corresponded exactly to those on rock substrates, suggesting their utility as representative alternatives for studying epilithic diatoms in future experiments. The influence of physico-chemical parameters on the initial colonisation of bacteria and diatoms on a submerged artificial substrate and the development of diatom commu- nities was studied weekly from April to October 2016 at a station in the marine lake Mrtvo More in southern Croatia (Car et al. 2020). The physico-chemical parameters (tem- perature, salinity, oxygen saturation, Chl-a concentration, nutrients: silicate, phosphate, nitrate, nitrite, ammonium) varied significantly according to month and season. Ac- cording to the TRIX trophic index, the lake showed differ- ent trophic characteristics: (i) oligotrophic (at the beginning and end of the study), (ii) mesotrophic (end of June to mid- July), (iii) eutrophic (end of July to mid-September). Hetero- trophic bacteria peaked (69,268 cells cm–2) in early June when the diatom abundance began to increase. Among the diatoms, adnate forms were the primary colonizers, partic- ularly Cocconeis dirupta W.Gregory var. flexella (Janisch and Rabenhorst) Grunow and Cocconeis scutellum Ehrenberg var. scutellum, while motile taxa joined the fouling communities from July to September. The lake has a high diatom species richness, with the species diversity index being highest in August. The total number of diatom taxa (285 within 72 genera detected on an artificial glass substrate is comparable to some studies of epilithic diatoms in the CAR A., KALELI A. 124 ACTA BOT. CROAT. 84 (2), 2025 Southern Adriatic (Hafner et al. 2018b, Car et al. 2019b), but higher than in previous studies of periphytic diatoms grow- ing on artificial substrates in the Northern Adriatic (Mejdandžić et al. 2015, Nenadović et al. 2015) or in a study of the surface sediment layer in the Venice Lagoon (Facca and Sfriso 2007). However, we believe that this could be due, at least in part, to differences in the methodology used. This study has shown that, among the physico-chemical param- eters, temperature, salinity and nitrate concentration have the greatest influence on the abundance of diatom species and that species diversity increases with nutrient enrich- ment. Strong correlations between environmental variables and diatoms were found and shifts in dominance at the spe- cies level were observed. Two of the most recent studies on benthic diatoms in the Adriatic Sea (Seveno et al. 2023, 2024) focused on blue Haslea species, known for the synthesis of blue-green water- soluble pigments such as marennine, which have allelopath- ic, antioxidant, antiviral and antibacterial properties. Re- cent descriptions of new species (H. karadagensis, H. nusantara, H. provincialis and H. silbo) have deepened our understanding of these pigments. While their beneficial properties have been demonstrated under laboratory condi- tions, the dynamics of blooms of blue Haslea spp. in its nat- ural habitat have been limited to oyster ponds. The study by Seveno et al. (2023) is the first documentation of benthic blooms of Haslea spp. in open environments, particularly on periphyton covering turf and macroalgae-like Padina. Two blue Haslea species – H. ostrearia and H. provincialis – were recorded at monitoring sites in the Mediterranean (Corsica, France and Croatia). The blooms followed the spring phytoplankton bloom and occurred in shallow, calm waters, suggesting that light plays an important role in their formation. The end of the blooms, marked by a warming of the upper water masses, coincided with shifts in the biofilm community, highlighting the importance of environmental conditions in regulating these unique benthic phenomena (Seveno et al. 2023). Most work on diatoms in the Adriatic Sea is based on LM and EM, while molecular biology techniques are only spo- radically used to analyse benthic diatom samples (Lobban et al. 2015, Witkowski et al. 2016, Li et al. 2019, Baldassarre et al., 2023). New molecular techniques can expand our knowledge of a species and enhance the understanding of its morphological range, biogeography and reproductive iso- lation (Medlin 2018). DNA sequencing has had a significant impact on the phylogeny, evolution and systematics of diatoms at the species and population level (Medlin and Kaczmarska 2004, Sorhannus 2007, Souffreau et al. 2011, Theriot et al. 2010, Witkowski et al. 2016). DNA metabar- coding has established itself as an alternative to LM-based identifications due to its speed, reproducibility and cost (Zimmermann et al. 2015). However, it is well known that a species can produce different diatom morphologies depend- ing on the season or habitat (see references in Cox 2014), and it has been proposed that the taxonomic level of forma be used to reflect morphologies that change with specific environmen- tal conditions (Cox 2014, Medlin 2018). Morphological changes are particularly striking in pennate diatoms, but they may be more subtle in centric species (Medlin 2018). Conflicts may arise between phylogenetic/molecular specia- tion and morphological speciation. It is recommended that a combination of species-level methods (both molecular and traditional) be used to resolve the species boundary issues. It cannot be assumed that the barcode of a single individual of a species is representative of the species, as different indi- viduals in a population and individuals in different geo- graphic populations may have slightly different barcode se- quences (Medlin 2018). To get a semi-quantitative overview of the diversity of an environmental sample, we would theo- retically first need to know the barcodes of all possible or- ganisms in the biosphere. Unfortunately, we are still very far away from such knowledge and it is questionable whether we will ever know them. GenBank provides sequences for perhaps five hundred identified species of marine and fresh- water diatoms (out of an estimated 100,000 species). The problem is that these sequences are underpinned with few voucher images and environmental data. A larger dataset of voucher sequences would improve DNA barcoding and al- low easier and faster identification of diatoms. This is oth- erwise a time-consuming process, requiring diatom taxon- omists to spend considerable time identifying and characterizing assemblages under the microscope. Current- ly, the traditional LM method for identifying diatoms in a mixed sample is still faster and more reliable than DNA bar- coding of environmental samples for an experienced diatom researcher (Jahn et al. 2007). However, the combination of LM and DNA approaches provides stronger support for eco- logical studies of benthic microalgal communities in shallow coastal environments than either approach alone (Pérez- - Burillo et al. 2022). In the study on the impact of MOSE (Experimental Electromechanical Module) flood barriers on the microphytobenthic community of the Venice Lagoon in 2019 and 2020, the integration of classical taxonomy with 18S rRNA gene metabarcoding enabled a more comprehen- sive understanding of the potential of the whole communi- ty and underlines the complementarity of these methods in ecological research (Baldassarre et al. 2023). Generally, SEM has revolutionised the systematics of di- atoms by revealing ultrastructural features visible that are invisible to LM, becoming essential for the delimitation of taxa; it is an essential aid to diatom classification (Round et al. 1990). The introduction of multivariate statistical meth- ods has further advanced diatom studies by revealing com- plex morphometric features. More recently, molecular biol- ogy techniques have provided even more insight into variation and greatly facilitated species discovery and de- limitation (Alverson 2008). Overall, these tools have im- proved our ability to recognise subtle differences. In addi- tion, collections support the ongoing discovery, classification and revision of diatom species and genera, contributing to the complex and dynamic taxonomy of diatoms. The first institutional collection of permanent microscopic speci- mens of diatoms in Croatia, the Croatian National Diatom BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 125 Collection, was established in March 2018 at the University of Zagreb, Faculty of Science. The collection contains, among other things, type specimens of newly discovered species in Croatia, which are no longer sent to other Euro- pean collections, but are kept in Croatia as national biodi- versity treasures. Collections such as this represent an in- valuable wealth of information for biological research, providing insights into the diversity of diatom taxonomy, the biodiversity of specific areas, species distribution and temporal and spatial changes. Taken together, this informa- tion facilitates the effective protection of water ecosystems in Croatia (Gligora Udovič and Ljubešić 2021). Since studies on the distribution of benthic diatom spe- cies along the Adriatic coast are scattered in space and time and provide rather heterogeneous information in terms of methodology, sample size and organisms studied, this re- view aims to provide an updated and integrated picture of Mediterranean benthic diatoms in the coastal waters of the Adriatic Sea, based on the studies conducted over the last 35 years. This paper aimed to compile a list of benthic diatom species detected in the Adriatic coasts based on the results of the most important studies carried out so far on marine diatoms and includes an updated nomenclature. The list was created to consolidate all previously scattered data for future research on benthic diatoms in Croatia, other coastal areas and beyond. This list aims to expand the knowledge of ma- rine diatoms in the Adriatic Sea by reviewing previous stud- ies and including an updated nomenclature. Methodological approach This study is based on literature records of marine dia- toms recorded in the Adriatic Sea. The literature search re- sulted in 52 references from 1989 to 2024 (On-line Suppl. Tab. 1), including planktonic diatoms and marine benthic diatoms in the coasts, coastal lakes and lagoons of the Adri- atic Sea. References included articles, reviews, checklists, and PhD theses. The diatom checklist was compiled using information from publications dating from 1999 to 2021. The old pub- lished data of diatoms from the Adriatic Sea by Viličić et al. (2002) were supplemented with more recent information. The main sources for the diatom checklist, besides Viličić et al. (2002), are Sdrigotti et al. 1999, Cibic et al. 2007a, Hafner et al. (2018a, b), Rogelja et al. 2018, and Car et al. (2019b, 2020, 2021). The results of this study depend on the correct identifi- cation by the authors of each reference. In most cases, there are no photographs or illustrations of the taxa, so verifica- tion of the records is difficult and some identifications may be incorrect. Despite efforts to avoid synonyms or the changed names of longer established taxa, some taxa may be listed here twice under different names. For each species, information on its study areas was pro- vided (Tab. 1, Fig. 1). The freshwater species reported in some studies were not excluded due to river pressure and altered salinity at different sites. Achnanthes apiculata (Greville) Riaux-Gobin, Compère, Hinz & Ector [M] Achnanthes brevipes var. brevipes Agardh* Achnanthes brevipes var. intermedia (Kützing) Cleve [M, S] Achnanthes brockmannii Hustedt [M, S] Achnanthes cf. ceramii Hendey [S] Achnanthes cf. javanica Grunow [M] Achnanthes cuneata Grunow [S] Achnanthes curvirostrum J.Brun [S] Achnanthes danica (Flögel) Grunow [N, S] Achnanthes fimbriata (Grunow) R.Ross [M] Achnanthes gracilis Lee [M] Achnanthes groenlandica (Cleve) Grunow [M, S] Achnanthes hyperboreoides A.Witkowski, Metzeltin & Lange-Bertalot [S] Achnanthes javanica Grunow [M, S] Achnanthes kuwaitensis Hendey [M, S] Achnanthes lemmermannii Hustedt [M] Achnanthes longipes C. Agardh* Achnanthes marginalis Hendey [M] Achnanthes parvula Kützing [M] Achnanthes pseudogroenlandica Hendey [M, S] Achnanthes separata Hustedt ex Simonsen [M, S] Achnanthes septata var. incurvata (Østrup) A. Cleve [M] Actinocyclus alienus Grunow [N] Actinocyclus gallicus F.Meister [M] Actinocyclus ingens var. irregularis (Van Heurck) D.M.Harwood & T.Maruyama [M] Actinocyclus normanii (W.Gregory ex Greville) Hustedt [M] Actinocyclus ochotensis A.P.Jousé [M] Actinocyclus octonarius Ehrenberg [N, S] Actinocyclus roperi (Brébisson) Grunow ex Van Heurck [S] Actinocyclus splendens J.Rattray [M] Actinocyclus subtilis (W. Gregory) Ralfs [M, S] Actinoneis lorenziana (Grunow) Mereschkowsky [N] Actinoptychus adriaticus Grunow [N, M] Actinoptychus heliopelta Grunow [M] Actinoptychus senarius (Ehrenberg) Ehrenberg [N, M] Actinoptychus splendens (Shadbolt) Ralfs [N, M] Actinoptychus vulgaris Schumann [M] Amphicocconeis disculoides (Hustedt) Stefano & Marino [S] Amphiprora pseudoduplex (Osada & Kobayasi) Hällfors [M] Amphitetras antediluviana Ehrenberg [N, S] Amphitetras subrotundata Janisch [S] Amphora abludens Simonsen [M, S] Amphora acuta Gregory [N] Amphora amoena Hustedt [M] Amphora angusta Gregory [N] Amphora arcus Gregory [N] Amphora arenaria Donkin* Tab. 1. Checklist of diatoms found in the Northern (N), Middle (M) and Southern (S) Adriatic Sea. Taxa found in all regions of the Adriatic Sea are indicated with asterisk (*). CAR A., KALELI A. 126 ACTA BOT. CROAT. 84 (2), 2025 Amphora arenicola Grunow ex Cleve [M] Amphora beaufortiana Hustedt [M] Amphora bigibba var. interrupta (Grunow) Cleve [M, S] Amphora binodis Gregory [N] Amphora cf. amoena Hustedt [M, S] Amphora cf. gacialis W.Smith [M] Amphora cf. ramsbottomi M.M.Salah [M] Amphora cingulata Cleve* Amphora constricta (Ehrenberg) W. Carruthers [M] Amphora copulata (Kützing) Schoeman & R.E.M.Archibald [M] Amphora corpulenta var. capitata Tempère & Peragallo [M] Amphora crassa W.Gregory* Amphora delicatissima Krasske [M, S] Amphora dubia Gregory [N] Amphora exilitata M.H.Giffen [S] Amphora gracialis W. Smith [M, S] Amphora gracilis Ehrenberg [M, S] Amphora graeffeana Hendey [M] Amphora graeffii Grunow [N] Amphora granulifera Cleve [N] Amphora grevilleana Gregory [M] Amphora helenensis Giffen [M, S] Amphora holsaticoides T.Nagumo & H.Kobayasi [M, S] Amphora immarginata Nagumo [M, S] Amphora insecta Grunow ex Schmidt* Amphora janischii A. Schmidt [N] Amphora jostesorum Witkowski. Lange-Bertalot & Metzeltin [S] Amphora laevis W.Gregory [N, M] Amphora laevissima W.Gregory* Amphora lineolata (Ehrenberg) Ehrenberg* Amphora lunata Østrup [M, S] Amphora marina W.Smith [N] Amphora mexicana A.W.F.Schmidt [N] Amphora obtusa W.Gregory [N, M] Amphora ocellata Donkin [S] Amphora ovalis (Kützing) Kützing [N, M] Amphora pannucea Giffen [M, S] Amphora pediculus (Kützing) Grunow [M] Amphora piper Cholnoky [S] Amphora praelata Hendey [S] Amphora proteus W.Gregory* Amphora pusio Cleve [N] Amphora robusta W.Gregory [N] Amphora staurophora (Castracane) Cleve [N] Amphora turgida var. parallela Heiden [S] Amphora weissflogii A. Schmidt [N] Aneumastus maculosus (Donkin) Lange-Bertalot. [N] Anorthoneis hyalina Hustedt [M] Ardissonea formosa (Hantzsch) Grunow* Ardissonea robusta (Ralfs ex Pritchard) De Notaris* Astartiella stroemii (Hustedt) Bukhtiyarova [N] Asterolampra marylandica Ehrenberg [M, S] Asteromphalus flabellatus (Brébisson ) Greville [N] Asteromphalus heptactis (Brébisson) Ralfs [N, M] Asteromphalus hookeri Ehrenberg [M] Asteromphalus robustus Castracane [N, M] Aulacoseira granulata (Ehrenberg) Simonsen [M, S] Auliscus sculptus (W.Smith) Brightwell [N, M] Auricula adriatica H. Peragallo & M. Peragallo [N, S] Auricula amphitritis Castracane [N, M] Auricula complexa (Gregory) Cleve [N] Auricula decipiens (Grunow) & Cleve [N] Auricula intermedia (F.W.Lewis) Cleve [S] Azpeitia nodulifera (A.W.F.Schmidt) G.A.Fryxell & P.A.Sims [N, M] Azpeitia obscura (Greville) P.A. Sims [N] Azpeitia tabularis (Grunow) Fryxell et P.A. Sims [N] Bacillaria paxillifera (O.F.Müller) T.Marsson* Bacillaria socialis (Gregory) Ralfs [M, S] Berkeleya micans (Lyngbye) Grunow [N] Berkeleya rutilans (Trentepohl ex Roth) Grunow [M, S] Berkeleya scopulorum (Brébisson ex Kützing) E.J.Cox [N, M] Biddulphia biddulphiana (J.E.Smith) Boyer* Biddulphia catenata Schussnig [N, M] Biddulphia spinosa (Bailey) Boyer [N] Biddulphia titiana Grunow [N, S] Biddulphiella tridens (Ehrenberg) P.A.Sims & M.P.Ashworth [N, M] Biremis ambigua (Cleve) D.G.Mann [M, S] Bleakeleya notata (Grunow) Round [S] Brachysira aponina Kützing [M] Brachysira estonarium Witkowski, Lange-Bertalot & Metzeltin [M] Brebissonia lanceolata (C.Agardh) R.K.Mahoney & Reimer [M, S] Caloneis bicuneata (Grunow) C.S.Boyer [M, S] Caloneis bioculata (Grunow) Hustedt [N] Caloneis clavicula Z.Levkov & Metzeltin [M] Caloneis egena (A.F.W.Schmidt) Cleve [N, M] Caloneis elongata (Grunow) Boyer [M, S] Caloneis excentrica (Grunow) Boyer [M, S] Caloneis fusioides (Grunow) Heiden & Kolbe [N, S] Caloneis latefasciata Grunow [N] Caloneis liber (W.Smith) Cleve* Caloneis linearis (Cleve) Boyer [N] Campylodiscus adriaticus Grunow* Campylodiscus biangulatus Greville* Campylodiscus clevei Leuduger-Fortmorel* Campylodiscus clypeus (Ehrenberg) Ehrenberg ex Kützing [M] Campylodiscus echeneis Ehrenberg ex Kützing [N, M] Campylodiscus fastuosus Ehrenberg [N, M] Campylodiscus giffenii Lobban & J.S.Park [M, S] Campylodiscus hodgsonii W. Smith [N] Campylodiscus imperialis Greville [N] Campylodiscus innominatus R.Ross & Abdin [M, S] Campylodiscus neofastuosus Ruck & Nakov* Campylodiscus punctulatoides M.Ricard [M] Campylodiscus thuretii Brébisson* Carinasigma latum (E.J.Cox) G.Reid [M] Carinasigma rectum (Donkin) G.Reid [N, S] Catacombas obtusa (Pantocsek) Snoeijs [S] Chamaepinnularia truncata (D.König) A.Witkowski, Lange-Bertalot & Metzeltin [M] Cistula lorenziana (Grunow) Cleve [M] Climacosphenia moniligera Ehrenberg* Cocconeiopsis orthoneoides (Hustedt) Witkowski, Lange-Bertalot & Metzeltin [M] Cocconeis britannica Naegeli in Kützing [N, M] Cocconeis californica Grunow [N, M] Cocconeis californica var. kerguelensis H.Heiden [M] Cocconeis capensis (Cholnoky) Witkowski [M] Tab. 1. Continued BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 127 Cocconeis cf. distantula Giffen [M, S] Cocconeis cf. irregularis (Schulz) Witkowski [S] Cocconeis convexa M.H.Giffen [S] Cocconeis costata var. hexagona Grunow [M, S] Cocconeis costata W.Gregory [M, S] Cocconeis dirupta var. flexella (Janisch & Rabenhorst) Grunow [M, S] Cocconeis dirupta W.Gregory [M, S] Cocconeis diruptoides Hustedt [S] Cocconeis discrepans A.W.F.Schmidt [M] Cocconeis distans W.Gregory [N, M] Cocconeis fasciolata (Ehrenberg) N.E.Brown [M] Cocconeis fimbriata Ehrenberg [N] Cocconeis fluminensis (Grunow) H. Peragallo & M. Pergallo [N] Cocconeis guttata Hustedt in Aleem & Hustedt [M, S] Cocconeis irregularis (P.Schulz) A.Witkowski in Witkowski [S] Cocconeis krammeri Lange-Bertalot & Metzeltin [M] Cocconeis latecostata Hustedt [M] Cocconeis maxima (Grunow) H. Peragallo & M. Peragallo [N, M] Cocconeis molesta Kützing [N, M] Cocconeis molesta var. crucifera Grunow [M] Cocconeis notabilis Pantocsek [M] Cocconeis nugalas M.H.Hohn & J.Hellerman [M] Cocconeis pelta A. Schmidt [M] Cocconeis peltoides Hustedt [M, S] Cocconeis pinnata Gregory ex Greville [N] Cocconeis placentula var. euglypta (Ehrenberg) Cleve [M] Cocconeis pseudodiruptoides Foged [S] Cocconeis pseudomarginata W.Gregory [N, S] Cocconeis schmidtii Heiden [M, S] Cocconeis scutellum Ehrenberg* Cocconeis speciosa W.Gregory [M, S] Cocconeis stauroneiformis H.Okuno [S] Cocconeis woodii Reyes-Vasquez [M, S] Coronia decora (Brébisson) Ruck & Guiry* Coronia impressa (Grunow) Ruck & Guiry [N] Coronia limbata (Brébisson) Ruck & Guiry [N, M] Coronia samoensis (Grunow) Ruck & Guiry [N] Coscinodiscus asteromphalus Ehrenberg [N] Coscinodiscus centralis Ehrenberg* Coscinodiscus cinctus Kützing [N] Coscinodiscus curvatulus Grunow [N, S] Coscinodiscus fimbriatus Ehrenberg [N] Coscinodiscus gigas Ehrenberg [S] Coscinodiscus granii Gough [S] Coscinodiscus hauckii var. mesoleius Cleve [M] Coscinodiscus janischii Schmidt [S] Coscinodiscus marginatus Ehrenberg [N] Coscinodiscus obscurus A. Schmidt [N] Coscinodiscus oculus iridis (Ehrenberg) Ehrenberg [N] Coscinodiscus perforatus Ehrenberg [N] Coscinodiscus radiatus Ehrenberg [N] Coscinodiscus thorii Pavillard [S] Craspedostauros decipiens (Hustedt) E.J.Cox [S] Craspedostauros indubitabilis (Lange-Bertalot & S.I.Genkal) E.J.Cox [S] Craticula cuspidata (Kutzing) D.G.Mann [N] Craticula halophila (Grunow) D.G.Mann [N] Ctenophora pulchella var. lanceolata (O'Meara) L.Bukhtiyarova [M] Cyclophora tenuis Castracane [M, S] Cyclotella kützingiana var. pelagica Grunow [N] Cyclotella striata (Kützing) Grunow [N] Cylindrotheca closterium (Ehrenberg) Reimann & J.C.Lewin [N, M] Cymatoneis sulcata (Greville) Cleve [M] Cymatosira belgica Grunow [N] Cymatosira lorenziana Grunow [M] Delphineis minutissima (Hustedt) Simonsen [M] Delphineis surirella (Ehrenberg) G.W.Andrews [N] Diatoma elongata (Lyngbye) C.Agardh [M] Diatoma vulgaris Bory [M] Dickieia subinflata (Grunow) D.G.Mann [M] Dimeregramma marinum (W.Gregory) Ralfs [N, M] Diploneis aestuarii Hustedt [N] Diploneis adonis (Brun) Cleve [N] Diploneis advena (A. Schmidt) Cleve [M] Diploneis bombus (Ehrenberg) Ehrenberg* Diploneis caffra (M.H.Giffen) Witkowski, Lange-Bertalot & Metzeltin [M] Diploneis campylodiscus (Grunow) Cleve [N] Diploneis cf. parca (A.W.F.Schmidt) Boyer [S] Diploneis cf. smithii (Brébisson) Cleve [N, M] Diploneis chersonensis (Grunow) Cleve [M, S] Diploneis coffaeiformis (A.W.F.Schmidt) Cleve [M] Diploneis crabro (Ehrenberg) Ehrenberg* Diploneis decipiens var. parallela A.Cleve [N] Diploneis dalmatica (Grunow) Cleve [M, S] Diploneis didyma (Ehrenberg) Ehrenberg* Diploneis dilatata (M.Peragallo) Lange-Bertalot & Fuhrmann [M] Diploneis distortum W. Smith [N] Diploneis divergens (A.W.F.Schmidt) Cleve [N, M] Diploneis fusca (W.Gregory) Cleve [N, M] Diploneis incurvata var. dubia Hustedt [M, S] Diploneis incurvata (W.Gregory) Cleve [M] Diploneis littoralis (Donkin) Cleve [M] Diploneis littoralis var. clathrata (Østrup) Cleve [M] Diploneis mediterranea (Grunow) Cleve [N] Diploneis mirabilis D.König [M] Diploneis nitescens (W.Gregory) Cleve [M, S] Diploneis notabilis (Greville) Cleve* Diploneis novaeseelandiae (A.Schmidt) Hustedt [M] Diploneis papula (A.W.F.Schmidt) Cleve [N, M] Diploneis parca (A. Schmidt) Boyer [M] Diploneis rex S.J.M.Droop [M] Diploneis sejuncta f. constricta Hustedt [M] Diploneis smithii (Brébisson) Cleve* Diploneis smithii var. dilatata (Peragallo) Lange-Bertalot & Fuhrmann [M] Diploneis smithii var. recta Peragallo [M, S] Diploneis splendida Cleve* Diploneis stroemii Hustedt [M, S] Diploneis subovalis Cleve [N] Diploneis subcincta (A. Schmidt) Cleve [N] Diploneis suborbicularis (Gregory) Cleve [M] Diploneis vacillans (A.W.F.Schmidt) Cleve* Diploneis vacillans var. renitens (A.Schmidt) Cleve [M, S] Diploneis vetula (A.W.F.Schmidt) Cleve [S] Diploneis weissflogii (A.W.F.Schmidt) Cleve [M] Divergita toxoneides (Castracane) Theriot [S] Ehrenbergiulva granulosa (Grunow) Witkowski, Lange-Bertalot & Metzeltin [N, M] Tab. 1. Continued CAR A., KALELI A. 128 ACTA BOT. CROAT. 84 (2), 2025 Ehrenbergiulva hauckii (Grunow) Witkowski, Lange-Bertalot & Metzeltin [N, M] Encyonema ventricosum (C.Agardh) Grunow [M] Entomoneis alata (Ehrenberg) Ehrenberg [N, S] Entomoneis corrugata (Giffen) Witkowski, Lange-Bertalot & Metzeltin [M] Entomoneis decussata (Grunow) K.Osada & H.Kobayasi [S] Entomoneis gigantea (Grunow) Nizamuddin [N] Entomoneis gigantea var. sulcata (O'Meara) Moreno, Licea & Santoyo [N, S] Entomoneis paludosa (W. Smith) Reimer* Entomoneis pulchra (Bailey) Reimer [S] Epithemia gibba (Ehrenberg) Kützing [N] Epithemia pacifica (Krammer) Lobban & J.S.Park [M, S] Fallacia aequorea (Hustedt) D.G.Mann [M] Fallacia bioculata (Grunow ex A.W.F.Schmidt) D.G.Mann [N] Fallacia cf. vittata (Cleve) D.G. Mann [M, S] Fallacia floriniae (M.Møller) Witkowski [M, S] Fallacia forcipata (Greville) Stickle & D.G.Mann* Fallacia hyalinula (De Toni) Stickle & D.G.Mann [N] Fallacia litoricola (Hustedt) D.G.Man [M] Fallacia nummularia (Greville) D.G.Mann [N, M] Fallacia ny (Cleve) D.G.Mann [M, S] Fallacia nyella (Hustedt) D.G.Mann [M, S] Fallacia pygmaea (Kützing) Stickle & D.G.Mann* Fallacia reichardtii (Grunow) Witkowski, Lange-Bertalot & Metzeltin [N] Fallacia rogalli Voigt [M] Fallacia spatiata (Østrup) Witkowski & Lange-Bertalot & Metzeltin [M] Fallacia versicolor (Grunow) D.G.Mann [N] Fogedia acuta (Salah) Witkowski, Lange-Bertalot & Metzeltin [S] Fogedia christensenii A.Witkowski, Metzeltin & Lange-Bertalot [S] Fogedia finmarchica (Cleve & Grunow) A.Witkowski, Metzeltin & Lange-Bertalot [S] Fragilaria barbatula (Kützing) Lange-Bertalot [N] Fragilaria capensis Grunow [M, S] Fragilaria improbula Witkowski & Lange-Bertalot [M] Fragilaria quebecensis (Poulin. Hudon & Cardinal) A.Witkowski. Lange-Bertalot & Metzeltin [M] Glyphodesmis distans (Gregory) Grunow in Van Heurck [N] Glyphodesmis williamsonii (W. Smith) Grunow [N] Gomphonella olivacea (Hornemann) Rabenhorst [M] Gomphonema apicatum Ehrenberg [M] Gomphonema coronatum Ehrenberg [M] Grammatophora angulosa Ehrenberg [N, S] Grammatophora angulosa var. islandica (Ehrenberg) Grunow [S] Grammatophora arcuata Ehrenberg [M, S] Grammatophora gibberula Kützing [N, M] Grammatophora hamulifera Kützing [N, M] Grammatophora longissima Petit [N] Grammatophora macilenta W. Smith* Grammatophora marina (Lyngbye) Kützing* Grammatophora maxima Grunow [S] Grammatophora oceanica Ehrenberg* Grammatophora oceanica var. subtilissima (Bailey) De Toni [M, S] Grammatophora serpentina Ehrenberg [M, S] Grammatophora undulata Ehrenberg* Grunowago bacillaris (Grunow) Lobban & Ashworth [N] Gyrosigma acuminatum (Kützing) Rabenhorst [N] Gyrosigma attenuatum (Kützing) Rabenhorst [N, S] Gyrosigma balticum (Ehrenberg) Rabenhorst [N, S] Gyrosigma balticum var. diminutum (Grunow) A.Cardinal, Poulin & Bérard-Therriault [N] Gyrosigma fasciola (Ehrenberg) J. W. Griffith & Henfrey [N] Gyrosigma lineare (Grunow) Cleve [N] Gyrosigma macrum (W.Smith) J.W.Griffith & Henfrey [N, S] Gyrosigma prolongatum (W.Smith) J.W.Griffith & Henfrey [N] Gyrosigma scalproides (Rabenhorst) Cleve [M] Gyrosigma strigilis (W.Smith) J.W.Griffin & Henfrey [N] Gyrosigma wansbeckii (Donkin) Cleve [N] Halamphora acutiuscula (Kützing) Levkov* Halamphora borealis (Kützing) Levkov [M] Halamphora coffeiformis (C. Agardh) Levkov* Halamphora costata (W.Smith) Levkov* Halamphora cuneata (Cleve) Levkov [M, S] Halamphora cymbifera (Gregory) Levkov [M] Halamphora exigua (W.Gregory) Levkov [M, S] Halamphora granulata (Gregory) Levkov [M, S] Halamphora holsatica (Hustedt) Levkov [M] Halamphora hyalina (Kützing) Rimet & R.Jahn* Halamphora hybrida (Grunow) Levkov [M] Halamphora interrupta (Heiden) Levkov [M] Halamphora kolbei (Aleem) Álvarez-Blanco & S.Blanco [M, S] Halamphora lineata (Gregory) Levkov [M] Halamphora luciae (Cholnoky) Levkov [M, S] Halamphora pseudoholsatica (T.Nagumo & H.Kobayashi) J.G.Stepanek & Kociolek [M, S] Halamphora pseudohyalina (Simonsen) J.G.Stepanek & Kociolek [M, S] Halamphora staurophora (Juhlin-Dannfelt) Álvarez-Blanco & S.Blanco [M, S] Halamphora subacutiuscula (Schoeman) J.G.Stepanek & Kociolek [M, S] Halamphora subangularis (Hustedt) Levkov [M, S] Halamphora subholsatica (Krammer) Levkov [M, S] Halamphora terroris (Ehrenberg) P.Wang [N] Halamphora turgida (Gregory) Levkov [S] Halamphora veneta (Kützing) Levkov [N] Hantzschia cf. distinctepunctata Hustedt [S] Hantzschia cf. marina (Donkin) Grunow [S] Hantzschia elongata (Hantzsch) Grunow [M] Hantzschia leptocephala (Østrup) Lange-Bertalot & Metzeltin [S] Hantzschia marina (Donkin) Grunow [N] Hantzschia virgata (Roper) Grunow [S] Haslea britannica (Hustedt & Aleem) Witkowski, Lange-Bertalot & Metzeltin [S] Haslea cf. balearica Witkowski [M] Haslea crucigera (W.Smith) Simonsen [S] Haslea duerrenbergiana (Hustedt) F.A.S.Sterrenburg [M, S] Haslea spicula (Hickie) Bukhtiyarova [M, S] Haslea stundlii (Hustedt) Blanco, Borrego-Ramos & Olenici [M] Haslea vitrea (Cleve) Simonsen [N] Hendeyella dubia (Grunow) Li, Witkowski & Ashworth* Hemiaulus hauckii Grunow ex Van Heurck [N] Hippodonta caotica Witkowski, Lange-Bertalot & Witkowski [S] Homoeocladia angularis (W.Smith) Kuntze [N, M] Homoeocladia distans (W.Gregory) Kuntze [N, M] Tab. 1. Continued BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 129 Hyalodiscus radiatus (O'Meara) Grunow [M, S] Hyalodiscus scoticus (Kützing) Grunow [M, S] Hyalodiscus subtilis Bailey [N] Hyalosira delicatula Kützing [N] Hyalosynedra laevigata (Grunow) D. M. Williams & Round* Iconella linearis (W.Smith) Ruck & Nakov [M] Iconella spiralis (Kützing) E.C.Ruck & T.Nakov [M] Karayevia submarina (Hustedt) Bukhtiyarova [M] Lampriscus shadboltianus (Greville) Peragallo & Peragallo [M] Licmophora abbreviata C.Agardh [M, S] Licmophora communis (Heiberg) Grunow [N] Licmophora dalmatica (Kützing) Grunow [N] Licmophora debilis (Kützing) Grunow [N, M] Licmophora ehrenbergii (Kützing) Grunow* Licmophora flabellata (Greville) C. Agardh* Licmophora gigantea Mereschkowsky [N] Licmophora gracilis (Ehrenberg) Grunow* Licmophora grandis (Kützing) Grunow [N] Licmophora hastata Mereschkowsky [N, M] Licmophora hyalina (Kützing) Grunow [N] Licmophora lyngbyei (Kützing) Grunow [N] Licmophora juergensii Agardh [N, S] Licmophora mediterranea Mereschkowsky [N] Licmophora paradoxa (Lyngbye) C. Agardh* Licmophora parasitica Mereschowsky [N] Licmophora pfannkuckae Giffen [M, S] Licmophora reichardtii Grunow [N, S] Licmophora remulus (Grunow) Grunow* Licmophora robusta H.Peragallo & M.Peragallo [N, S] Licmophora tenuis (Kützing) Grunow [N] Licmophora tincta (C.Agardh) Grunow [S] Lindavia glomerata (H.Bachmann) Adesalu & Julius [N] Luticola ventricosa (Kützing) D.G.Mann [M] Lyrella abrupta (W.Gregory) D.G.Mann [N, M] Lyrella amphoroides D.G.Mann [M] Lyrella cf. lyroides (Hendey) D.G.Mann [M] Lyrella clavata (W.Gregory) D.G.Mann [N, M] Lyrella david-mannii Witkowski, Lange-Bertalot & Metzeltin [M] Lyrella fogedii Witkowski, Lange-Bertalot & Metzeltin [M] Lyrella hennedyi (W.Smith) Stickle & D.G.Mann* Lyrella lyra (Ehrenberg) Karayeva [M, S] Lyrella lyroides (Hendey) D.G.Mann [N, M] Lyrella spectabilis (Gregory) D. G. Mann [N, M] Lyrella subforcipata (Hustedt) Gusliakov & Karayeva [M] Lyrella sulcifera (Hustedt) Witkowski [M] Martyana schulzii (C.Brockmann) Snoeijs [M, S] Mastogloia adriatica Voigt [M, S] Mastogloia affinis Cleve [M] Mastogloia angulata F.W.Lewis [N, S] Mastogloia apiculata W. Smith [N] Mastogloia baldjikiana Grunow [M] Mastogloia beaufortiana Hustedt [M] Mastogloia belaensis Voigt [S] Mastogloia binotata (Grunow) Cleve* Mastogloia biocellata (Grunow) G.Novarino & A.R.Muftah [S] Mastogloia borneensis Hustedt [M, S] Mastogloia braunii Grunow* Mastogloia cf. affirmata (Leudiger-Fortmorel) Cleve [S] Mastogloia corsicana (Grunow) H.Peragallo & M.Peragallo* Mastogloia crucicula var. alternans Zanon [M, S] Mastogloia crucicula (Grunow) Cleve* Mastogloia cuneata (F.Meister) Simonsen [M, S] Mastogloia cyclops Voigt [M, S] Mastogloia decipiens Hustedt [M, S] Mastogloia decussata Grunow [M, S] Mastogloia emarginata Hustedt [M, S] Mastogloia erythraea Grunow* Mastogloia erythraea var. grunowii Foged [M] Mastogloia exigua F.W.Lewis [M, S] Mastogloia exilis Hustedt [M, S] Mastogloia fallax Cleve [M, S] Mastogloia fimbriata (T.Brightwell) Grunow* Mastogloia foliolum Brun [M] Mastogloia grunowii A.W.F.Schmidt [M, S] Mastogloia horvathiana Grunow [M, S] Mastogloia jelineckii (Grunow) Grunow [M, S] Mastogloia ignorata Hustedt [M] Mastogloia inaequalis Cleve [N] Mastogloia laminaris (Ehrenberg) Grunow [S] Mastogloia lanceolata Thwaites ex W. Smith [N, S] Mastogloia latecostata Hustedt [M] Mastogloia linearis Simonsen [M] Mastogloia macdonaldii Greville* Mastogloia manokwariensis Cholnoky [M, S] Mastogloia marginulata Grunow [N] Mastogloia mauritiana Brun* Mastogloia mediterranea Hustedt [M] Mastogloia obliqua Hagelstein [S] Mastogloia ovalis A.W.F.Schmidt [M, S] Mastogloia ovata Grunow [N, M] Mastogloia ovulum Hustedt* Mastogloia ovum-paschale (A. Schmidt) Mann [N] Mastogloia paradoxa Grunow [N, M] Mastogloia parva Hustedt [M, S] Mastogloia peragalloi Cleve* Mastogloia pseudoexigua Cholnoky [M] Mastogloia pseudolatecostata T.A.Yohn & R.A.Gibson [M, S] Mastogloia pumila (Grunow) Cleve [N, M] Mastogloia punctifera Brun [N] Mastogloia pusilla Grunow* Mastogloia quinquecostata Grunow [N, M] Mastogloia recta Hustedt [M] Mastogloia regula Hustedt [M, S] Mastogloia robusta Hustedt [S] Mastogloia similis Hustedt [M, S] Mastogloia smithii Thwaites ex W.Smith [N, M] Mastogloia splendida (Gregory) H.Pergallo* Mastogloia subaspera Hustedt [N] Mastogloia undulata Grunow [N] Mastogloia varians Hustedt [M, S] Mastogloia vasta Hustedt [M, S] Melosira lineata (Dillwyn) C.Agardh [N] Melosira moniliformis var. octogona (Grunow) Hustedt [M] Melosira moniliformis C.Agardh* Melosira nummuloides C.Agardh* Melosira varians C.Agardh [M] Meridion circulare (Greville) C.Agardh [M] Metascolioneis tumida (Brébisson ex Kützing) Blanco & Wetzel [N] Tab. 1. Continued CAR A., KALELI A. 130 ACTA BOT. CROAT. 84 (2), 2025 Microtabella interrupta (Ehrenberg) Round* Nanofrustulum cf. sopotense (Witkowski & Lange-Bertalot) E.Morales, C.E.Wetzel & L.Ector [S] Nanofrustulum krumbeinii (A.Witkowski, Witak & K.Stachura) E. Morales [M] Nanofrustulum shiloi (J.J.Lee, Reimer & McEnery) Round, Hallsteinsen & Paasche [S] Nanofrustulum sopotense (Witkowski & Lange-Bertalot) E.Morales, C.E.Wetzel & Ector [M, S] Navicula abunda Hustedt [M] Navicula agnita Hustedt [M, S] Navicula ammophila Grunow [N] Navicula apiculata Brébisson [M] Navicula aproximata Greville [N] Navicula arenaria Donkin [N, M] Navicula arenaria var. rostellata Lange-Bertalot [M] Navicula avenacea (Rabenhorst) Brébisson ex Grunow [N] Navicula bipustulata A.Mann [M, S] Navicula borneoensis Hustedt [M, S] Navicula cancellata Donkin* Navicula cf. fauta Hustedt [M] Navicula cincta (Ehrenberg) Ralfs [M, S] Navicula cluthensis f. rostrata (Simonsen) Hustedt [M] Navicula cluthensis var. cluthensis Gregory [M] Navicula crucifera Grunow [M] Navicula dehissa Giffen [S] Navicula digitoradiata (W.Gregory) Ralfs [N, M] Navicula directa (W.Smith) Brébisson* Navicula distans (W.Smith) Brébisson [N, S] Navicula eidrigiana J.R.Carter [M, S] Navicula erifuga Lange-Bertalot [S] Navicula flagellifera Hustedt [M, S] Navicula frigida Grunow [N] Navicula formentere Cleve [S] Navicula gregaria Donkin [S] Navicula grevilleana Hendey [N] Navicula grippii Simonsen [S] Navicula hauckii Cleve [N] Navicula johanrossii Giffen [S] Navicula kariana Grunow [S] Navicula libellus Gregory [N] Navicula longa (W.Gregory) Ralfs [M] Navicula longa var. irregularis Hustedt [M] Navicula luciae A.Witkowski & Lange-Bertalot [M] Navicula lusoria M.F.Giffen [M, S] Navicula mediterranea Cleve et Brun [N] Navicula meniscus Schumann [N] Navicula nortumbrica Donkin [N] Navicula palpebralis Brébisson ex W.Smith* Navicula palpebralis var. angulosa (W.Gregory) Van Heurck [M] Navicula palpebralis var. minor Grunow [S] Navicula pavillardii Hustedt [M] Navicula pennata f. maxima (Cleve) Frenguelli [M] Navicula pennata A.W.F.Schmidt [N, M] Navicula peregrina (Ehrenberg) Kützing [N] Navicula phyllepta Kützing [N] Navicula pinnata Pantocsek [M] Navicula quarnerensis Grunow [S] Navicula ramosissima (C.Agardh) Cleve [N, M] Navicula recurvata Gran [S] Navicula rolandii S.Wunsam, A.Witkowski, & Lange-Bertalot [M] Navicula rostellata Kützing [S] Navicula salinarum Grunow [M] Navicula salinarum var. rostrata (Hustedt) Lange-Bertalot [S] Navicula salinicola Hustedt [S] Navicula subagnita Proshkina-Lavrenko [M, S] Navicula subrostellata Hustedt [M] Navicula superimposita A. Schmidt [N] Navicula vara Hustedt [M] Navicula viridula (Kützing) Ehrenberg [N] Navicula zostereti Grunow [N, M] Neidium trigonocephalum (Ehrenberg) A.Tuji & Y.Tanimura [M] Neobrightwellia alternans (Bailey) M.P.Ashworth & Sims [N, M] Neofragilaria burchardtiae (Witkowski, Metzeltin & Lange-Bertalot) E.Morales [M, S] Neofragilaria nicobarica Desikachary, Prasad & Prema [M] Neohuttonia reichardtii (Grunow) Hustedt [S] Nitzschia acuminata var. subconstricta Grunow [M] Nitzschia agnewii Cholnoky [S] Nitzschia agnita Hustedt [M, S] Nitzschia bartholomei (Grunow) Lange-Bertalot & Krammer [M] Nitzschia behrei Hustedt [M] Nitzschia bilobata W.Smith [N, M] Nitzschia bulnheimiana (Rabenhorst) H.L.Smith [S] Nitzschia capitellata Hustedt [M, S] Nitzschia carnicobarica Desikachary & Prema [S] Nitzschia cf. bulnheimiana (Rabenhorst) H.L.Smith [M] Nitzschia cf. commutata Grunow [N] Nitzschia cf. sigma (Kützing) W. Smith [M, S] Nitzschia commutatoides Lange-Bertalot [M] Nitzschia distans var. tumescens Grunow [M] Nitzschia fasciculata (Grunow) Grunow [N] Nitzschia fluminensis Grunow [N, M] Nitzschia frustulum (Kützing) Grunow* Nitzschia fusiformis Grunow [M, S] Nitzschia gazellae Karsten [N] Nitzschia grossestriata Hustedt [M, S] Nitzschia hybrida Grunow [M, S] Nitzschia improvisa Simonsen [S] Nitzschia incerta (Grunow) M.Peragallo* Nitzschia incurva Grunow* Nitzschia incurvata var. lorenziana R.Ross [M, S] Nitzschia insignis W.Gregory* Nitzschia laevis Frenguelli [S] Nitzschia lanceola Grunow [M] Nitzschia lanceolata var. minor (Grunow) H.Peragallo & M.Peragallo [M, S] Nitzschia lanceolata W. Smith [M] Nitzschia liebethruthii Rabenhorst [M, S] Nitzschia longissima (Brébisson) Ralfs* Nitzschia macilenta W.Gregory* Nitzschia marginulata var. didyma Grunow [M, S] Nitzschia microcephala Grunow [N] Nitzschia normannii Grunow [M] Nitzschia obtusa W. Smith [N] Nitzschia ovalis H.J.Arnott [M, S] Nitzschia peisonis Pantocsek [M] Nitzschia pellucida Grunow [M, S] Tab. 1. Continued BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 131 Nitzschia plioveterana Lange-Bertalot [M] Nitzschia reversa W.Smith [S] Nitzschia scalpelliformis Grunow [M] Nitzschia sicula (Castracane) Hustedt [N, S] Nitzschia sigma (Kützing) W.Smith [N, M] Nitzschia sigma var. rigida Grunow ex Van Heurck* Nitzschia sigma var. rigidula (H.Peragallo & M.Peragallo) Grunow [N] Nitzschia sigma var. sigmatella Grunow [N] Nitzschia sigmaformis Hustedt [M] Nitzschia subconstricta Desikachary & Prema [M, S] Nitzschia valdestriata Aleem & Hustedt [S] Nitzschia ventricosa J.L.Palmer [M, S] Nitzschia vermicularis (Kützing) Hantzsch [N] Odontella aurita (Lyngbye) C.Agardh [N, M] Oestrupia ergadensis (W.Gregory) Witkowski, Lange-Bertalot & Metzeltin [M] Oestrupia kinkeriana (Truan) Heiden [N] Oestrupia powellii var. vidovichii (Grunow) Heiden [M] Oestrupia powellii (F.W.Lewis) Heiden [N, M] Okedenia inflexa (Brébisson ex Kützing) Eulenstein ex De Toni [N] Opephora marina (W.Gregory) Petit [N, M] Opephora mutabilis (Grunow) Sabbe & Wyverman [M, S] Opephora pacifica (Grunow) Petit* Opephora schwartzii (Grunow) Petit ex Pelletan [M] Pantocsekiella ocellata (Pantocsek) K.T.Kiss & E.Ács [M] Paralia sulcata (Ehrenberg) Cleve* Parlibellus bennikei Witkowski, Metzeltin & Lange-Bertalot [M, S] Parlibellus berkeleyi (Kützing) E.J.Cox [S] Parlibellus calvus A.Witkowski, Metzeltin & Lange-Bertalot [S] Parlibellus cf. cruciculoides (C.Brockmann) Witkowski, Lange-Bertalot & Metzeltin [S] Parlibellus delognei (Van Heurck) E.J.Cox [M, S] Parlibellus hamulifer (Grunow) E.J.Cox [N] Parlibellus plicatus (Donkin) E.J.Cox [N] Parlibellus rhombicula (Hustedt) Witkowski, Lange-Bertalot & Metzeltin [M, S] Parlibellus rhombicus (W.Gregory) E.J.Cox [N] Petrodictyon gemma (Ehrenberg) D.G.Mann [M, S] Petroneis besarensis (Giffin) Witkowski, Lange-Bertalot & Witkowski [M, S] Petroneis humerosa (Brébisson ex W.Smith) Stickle & D.G.Mann [M] Pinnularia claviculus Schulz [M, S] Pinnularia quadratarea var. cuneata Østrup [S] Pinnularia rectangulata (Gregory) Rabenhorst [N] Pinnularia trevelyana (Donkin) Rabenhorst [M] Placoneis flabellata (F.Meister) Kimura, H.Fukushima & Ts.Kobayashi [S] Plagiodiscus martensianus Grunow & Eulenstein [M] Plagiodiscus nervatus Grunow [M] Plagiogramma atomus Greville [M] Plagiogramma interruptum (W.Gregory) Ralfs [N, M] Plagiogramma laeve (Gregory) Ralfs [N] Plagiogramma minus (W.Gregory) Chunlian Li, Ashworth & Witkowski [N, M] Plagiogramma minus var. nanum (W.Gregory) Chunlian Li, Ashworth & Witkowski [N, M] Plagiogramma pulchellum Greville [N] Plagiogramma staurophorum (W.Gregory) Heiberg* Plagiogrammopsis vanheurckii (Grunow) Hasle, Stosch & Syvertsen [M] Plagiotropis elegans (W.Smith) Grunow [N] Plagiotropis gibberula Grunow [N] Plagiotropis lata (Cleve) Kuntze [N] Plagiotropis lepidoptera (W. Gregory) Kuntze* Plagiotropis tayrecta T.B.B.Paddock [M, S] Plagiotropis vitrea (W. Smith) Kuntze [N] Planothidium aff. engelbrechtii (Cholnoky) Round & L.Bukhtiyarova [M] Planothidium cf. campechianum (Hustedt) Witkowski, Lange-Bertalot & Metzeltin [M] Planothidium delicatulum (Kützing) Round & Bukhtiyarova [M] Planothidium dispar (Cleve) Witkowski, Lange-Bertalot & Metzeltin [N] Planothidium quarnerense (Grunow) Witkowski. Lange-Bertalot & Metzelin [N, M] Platessa conspicua (Ant.Mayer) Lange-Bertalot [M] Pleurosigma aestuari (Brébisson) W. Smith [N] Pleurosigma angulatum (J.T.Quekett) W.Smith* Pleurosigma decorum W.Smith [M, S] Pleurosigma elongatum W. Smith [N, S] Pleurosigma formosum W.Smith* Pleurosigma itium Ricard [M] Pleurosigma longum Cleve [N] Pleurosigma nicobaricum Grunow [N, M] Pleurosigma normanii Ralfs [N] Pleurosigma rigidum W. Smith [N] Pleurosigma speciosum W. Smith [N] Pleurosigma strigosum W. Smith [N] Pleurosira laevis (Ehrenberg) Compère [M] Podocystis adriatica (Kützing) Ralfs* Podosira dubia (Kützing) Grunow [N] Prestauroneis protracta (Grunow) Kulikovskiy & Glushchenko [M] Proschkinia complanata (Grunow) D.G.Mann [N] Protokeelia cholnokyi (Giffen) Round & Basson [M, S] Protoraphis atlantica R.A.Gibson [M] Psammodictyon constrictum (W.Gregory) D.G.Mann [M] Psammodictyon panduriforme (W. Gregory) D.G.Mann* Psammodictyon panduriforme var. continuum (Grunow) Snoeijis [M] Psammodictyon roridum (Giffen) D.G.Mann [M, S] Psammodictyon rudum (Cholnoky) D.G.Mann [M, S] Psammodiscus nitidus (W.Gregory) Round & D.G.Mann [N, M] Pseudostaurosira perminuta (Grunow) Sabbe & Wyverman [N] Pseudopodosira westii (W.Smith) Sheshukova-Poretskaya & Glezer [N] Pseudostaurosiropsis geocollegarum (Witkowski) E.A.Morales [M] Pseudostaurosiropsis punctiformis (Witkowski, Metzeltin & Lange- Bertalot) Witkowski, Seddon & Pliński [M] Rhabdonema adriaticum Kützing* Rhabdonema arcuatum (Lyngbye) Kützing [S] Rhabdonema minutum Kützing [N, M] Rhaphoneis amphiceros (Ehrenberg) Ehrenberg [N, M] Rhaphoneis nitida (Gregory) Grunow [N] Rhizosolenia styliformis T.Brightwell [N, M] Rhoicosigma compactum (Greville) Grunow [N] Rhoicosphenia abbreviata (C.Agardh) Lange-Bertalot [M, S] Rhoicosphenia linearis Østrup [M] Tab. 1. Continued CAR A., KALELI A. 132 ACTA BOT. CROAT. 84 (2), 2025 Rhoicosphenia marina (Kützing) M. Schmidt [M, S] Rhopalodia acuminata Krammer [M] Rhopalodia gibberula (Ehrenberg) Müller [N] Rhopalodia musculus (Kützing) Otto Müller* Rhopalodia sterrenburgii Krammer [M] Sarcophagodes mutabilis E.A.Morales [S] Seminavis barbarae Witkowski, Lange-Bertalot & Metzeltin [M] Seminavis obtusiuscula (Grunow) Danieledis & D.G.Mann [S] Shionodiscus oestrupii (Ostenfeld) A.J.Alverson, S.-H.Kang & E.C.Theriot [N] Skabitschewskia borealis (A.Cleve) Lange-Bertalot & Kulikovskiy [M] Stauroneis plicata C.Brockmann [M, S] Stauroneis smithii Grunow [M] Stauroneis undata Hustedt [S] Staurophora amphioxys (W.Gregory) D.G.Mann [N] Staurosirella guenter-grassii (Witkowski & Lange-Bertalot) E.A.Morales, C.E.Wetzel & Ector [M, S] Staurosirella martyi (Héribaud) Morales & Manoylov [M, S] Stephanocyclus meneghiniana (Kützing) Kulikovskiy, Genkal & Kociolek [M] Stephanodiscus hantzschii Grunow [S] Stoermeria americana (Bailey) M.P.Ashworth & P.A.Sims [M] Striatella unipunctata (Lyngbye) C.Agardh* Surirella baldjicki Norman [N] Surirella brebissonii Krammer & Lange-Bertalot [M] Surirella ceylanensis var. oblongistriata F.Hustedt [M] Surirella comis A. Schmidt [N] Surirella crumena Brébisson ex Kützing [M] Surirella gemma var. reniformis Proshkina-Lavrenko [N] Surirella fluminensis Grunow [N] Surirella hybrida Grunow [N, M] Surirella intercedens Grunow [N] Surirella lata W.Smith [M] Surirella minuta Brébisson ex Kützing [N] Surirella ovalis Brébisson [N] Surirella striatula Turpin [M] Surirella venusta Østrup [S] Synedra brockmannii Hustedt [N] Synedra commutata Grunow [M] Synedra distinguenda Hustedt [N] Synedra gaillonii (Bory) Ehrenberg [M, S] Synedra laevis Kützing [S] Synedra pulcherrima Hantzsch [N] Synedrosphenia baculus (W.Gregory) Lobban & Ashworth [N] Synedrosphenia crystallina (C.Agardh) Lobban & Ashworth* Synedrosphenia fulgens (Greville) Lobban & Ashworth* Tabularia fasciculata (C.Agardh) D.M.Williams & Round [M, S] Tabularia gaillonii (Bory) Bukhtiyarova [N] Tabularia investiens (W.Smith) D.M.Williams & Round* Tabularia ktenoeides M.Kuylenstierna [M, S] Tabularia tabulata (C.Agardh) Snoeijs [N] Talaroneis furcigera (Grunow) Sterrenburg [N, M] Tetramphora decussata (Grunow) Stepanek & Kociolek* Tetramphora ostrearia (Brébisson) Mereschkowsky [N, S] Tetramphora rhombica (Kitton) Stepanek & Kociolek [N, M] Tetramphora securicula (Peragallo & Peragallo) Stepanek & Kociolek [N, M] Tetramphora sulcata (Brébisson) Stepanek & Kociolek* Thalassionema frauenfeldii (Grunow) Tempère & Peragallo* Thalassionema nitzschioides (Grunow) Mereschkowsky* Thalassiophysa hyalina (Greville) Paddock & P.A.Sims [M] Thalassiosira angulata (Gregory) Hasle* Thalassiosira eccentrica (Ehrenberg) Cleve* Thalassiosira gravida Cleve [N, S] Thalassiosira hyalina (Grunow) Gran [M] Thalassiosira leptopus (Grunow) Hasle et Fryxell* Thalassiosira mediterranea (Schröder) Hasle [N] Thalassiosira nordenskioeldii Cleve [S] Thalassiosira subtilis (Ostenfeld) Gran [N] Toxarium hennedyanum (Gregory) Pelletan* Toxarium undulatum J.W.Bailey* Toxonidea balearica Grunow [N, S] Toxonidea insignis Donkin [N] Trachyneis aspera (Ehrenberg) Cleve* Trachysphenia australis Petit [N] Triceratium broeckii Lauduger – Fortmorel [S] Triceratium favus Ehrenberg [N, M] Triceratium finnmarchicum Grunow [S] Triceratium pellucidum (Castracane) Y.C.Guo, J.Ye & H.Zhou [N] Triceratium pelagicum (J.L.B.Schröder) Sournia [N, M] Triceratium pentacrinus Wallich [M, S] Triceratium reticulum Ehrenberg [M, S] Trieres mobiliensis (J.W.Bailey) Ashworth & Theriot [M, S] Trigonium arcticum (Brightwell) Cleve [S] Trigonium formosum (Brightwell) Cleve [M, S] Tropidoneis maxima (W.Gregory) Cleve [N] Tropidoneis longa (Cleve) Cleve [N] Tryblionella acuminata W.Smith [N, M] Tryblionella acuta (Cleve) D.G.Mann [N] Tryblionella apiculata W.Gregory [N, M] Tryblionella cf. littoralis (Grunow) D.G.Mann [N] Tryblionella coarctata (Grunow) D.G.Mann [N, S] Tryblionella compressa (Bailey) Poulin [M, S] Tryblionella debilis Arnott ex O'Meara [N] Tryblionella didyma (Hustedt) D.G.Mann [M, S] Tryblionella hantzschiana Grunow* Tryblionella hungarica (Grunow) Frenguelli [M, S] Tryblionella littoralis (Grunow) D.G.Mann [M] Tryblionella marginulata (Grunow) D.G.Mann [N, M] Tryblionella navicularis (Brébisson) Ralfs* Tryblionella pararostrata (Lange-Bertalot) Clavero & Hernández- Mariné [M] Tryblionella persuadens (Cholnoki) K.P.Cavalcante. P.I.Tremarin & T.A.V.Ludwig [M, S] Tryblionella plana (W.Smith) Pelletan [N] Tryblionella punctata W.Smith [M, S] Tryblionella punctata var. elongata Grunow [N] Tryblionella scalaris (Ehrenberg) Siver & P.B.Hamilton [M] Ulnaria longissima (W.Smith) Van de Vijver & D.M.Williams [S] Ulnaria ulna (Nitzsch) Compère [N, S] Undatella lineata (Greville) Paddock & P.A.Sims [M] Undatella quadrata (Brébisson ex Kützing) Paddock & P.A.Sims [M] Vikingea promunturi (Giffen) Witkowski, Lange-Bertalot & Metzeltin [S] Zygoceros rhombus Ehrenberg [N] Tab. 1. Continued BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 133 Taxonomic classification was done according to Round et al. (1990). Previously, some species were given names that differed from those of the latest taxonomic order. An at- tempt was made to classify the species with updated current names according to Guiry and Guiry (2024), Kociolek et al. (2024) and the website Algaebase (http://www. algaebase. org/), which is constantly updated and therefore has a dif- ferent taxonomic hierarchy than the one previously used. The current status of marine benthic diatoms in Adriatic The present work represents the first detailed list of ma- rine benthic diatoms in the Adriatic Sea. An overview of all 52 publications included in this review shows that diatoms from the Northern Adriatic were analysed in most studies (N=32), followed by Middle Adriatic (N=15) and Southern Adriatic (N=12). Only two studies analysed diatoms on a larger spatial scale covering all regions of the Adriatic (On- line Suppl. Tab. 1). The literature was unevenly distributed, with 30 papers for the Italian coast and 18 for the Croatian coast. The coast of Bosnia and Herzegovina, on the other hand, has been the subject of 2 studies, while Slovenia and Albania are represented by 1 study each (On-line Suppl. Tab. 1). The number of publications on diatom research fluctu- ated, with the most productive period being that of 2000- 2024, during which 47 papers were published (On-line Sup- pl. Tab. 1). In certain periods, research activity has increased considerably in particular areas, such as the recent intensive studies of diatoms in the Southern Adriatic. Since the be- ginning of diatom studies mostly in Northern Adriatic, sev- eral authors have made significant contributions to the com- position of diatoms over the years. In this paper, we merge the data with the most recent nomenclature, focusing on the current names of the cited taxa (On-line Suppl Tab. 2). A list of updated species nomen- Fig. 1. A) Major seas of the Mediterranean, B) Locations of the studies of diatom taxa from the Adriatic Sea presented in Suppl. Tab. 1. CAR A., KALELI A. 134 ACTA BOT. CROAT. 84 (2), 2025 clature (On-line Suppl Tab. 2) cited in the previous studies (Cited Name) was given (Tab. 1) and the species/genus names were checked for validity using AlgaeBase (Guiry and Guiry, 2024). Marine diatoms were represented by 822 species (163 genera and 70 families) (Tab. 1). The number of pennate species represented was 719. Families with the highest number of genera were: Naviculaceae (8), Bacillariaceae (7), Surirellaceae (6), Fragilariaceae (6). Additionally, the high- est number of species and infraspecific taxa belonged to the Naviculaceae (95) and followed by Bacillariaceae (88). The taxonomic composition of benthic diatoms in the Adriatic may be defined as the community characterized by genera such as; Mastogloia (65 taxa), Navicula (62), Nitzschia (53), Amphora (52), Diploneis (43), Cocconeis (37), Halamphora (24), Achnanthes (22), Licmophora (22), Tryblionella (19), Fallacia (15), Surirella (14), Grammatophora (13), Lyrella (12). Altogether, 381 taxa were found in the Northern, 524 taxa in the Middle and 373 taxa in the Southern Adriatic Sea. Among the total 822 diatom taxa, 177 taxa were found exclusively in the Northern Adriatic, 195 were found exclu- sively in the Middle and 94 occurred only in Southern Adri- atic (Tab. 1). The studies used in this checklist have shown that many species were documented as first records in the Adriatic. The variation in findings of benthic diatoms across dif- ferent geographical areas may also be attributed to differing research methodologies. In the Adriatic Sea, benthic diatom analyses are often conducted within the framework of oth- er research studies, such as phytoplankton investigations. Additionally, studies of benthic diatoms make use of vari- ous substrates, both natural and artificial. However, the number of diatom species on the Adriatic coasts is similar to the checklist of marine diatoms from Turkish coastal waters (Kaleli and Akçaalan 2021), and the reason for the low number of marine diatoms (767 taxa be- longing to 183 genera, of which 567 taxa belonged to Bacil- lariophyceae in Türkiye) may be attributed to the restricted scope of benthic studies along these coastlines, as highlight- ed by Kaleli and Akçaalan (2021). However, recent findings (Kaleli et al. 2020) suggest a potential increase in the num- ber of taxa in forthcoming studies for both the Turkish coastal waters and the Adriatic Sea, particularly with those specifically geared towards the detection of benthic species. This paper revealed the benthic diatom composition in the Adriatic coastal waters with the latest nomenclature and could be used as a fundamental list for later studies. Conclusion The study of marine diatoms on the coasts of the Adri- atic Sea began more than 100 years ago, and in the last 35 years, extensive studies have been carried out. However, ma- ny geographical areas have not yet been studied and their diatom composition remains unknown. Therefore, this checklist aims to give an overview of the benthic diatoms that have been studied in the last 35 years and to create a database with updated nomenclature to fill the gap regard- ing marine benthic diatoms along the Adriatic coasts. So far, however, there has been very little information on benthic diatoms in the Adriatic Sea. The knowledge of benthic diatom communities in the Adriatic Sea is poor compared to longer-term studies on phytoplankton. In this work, studies on marine plankton and benthos were com- bined to create a dataset that could be comparable for future studies in the Adriatic and other regions. Although check- lists existed for marine plankton, this study provided a list of diatoms that included both forms from the Adriatic coasts. The inclusion of data on benthic taxa improves the knowledge of overall microalgal diversity and of coastal and transitional ecosystem ecology. The list included many ben- thic diatoms as well as the planktonic forms mentioned above but showed that not all the taxa on the Adriatic coasts are fully covered. Future studies should focus on the com- position of benthic diatoms using LM and EM to provide additional data for further implications. Morphological de- tails of benthic diatoms would not only increase the knowl- edge of taxonomy but also of species diversity and geo- graphical distribution. However, as many of the species were not illustrated, future studies may increase the knowledge of accurate identification based on illustrated studies in the region. It should be noted that there have been no changes in the taxonomy of genera and species. The nomenclature has been updated for several taxa that have been transferred or are currently no longer in use. Therefore, further studies should be carried out to clarify the problems of diatom sys- tematics in the Adriatic coastal waters. This is the first exhaustive checklist of benthic diatoms of the Adriatic Sea, including the latest papers, updated with very recent books and websites, and can be used as a valu- able tool for further studies. However, it should be noted that the checklist should be updated over time. Further studies on the structure of the diatom flora would provide additional information that would lead to a better ecologi- cal understanding and an approach to coastal ecosystem monitoring to prevent damage to marine biodiversity, espe- cially for the disturbed Adriatic coasts. Nevertheless, knowl- edge of the composition and spatial distribution of marine benthic diatoms along the coast of the Adriatic coast re- mains limited. Acknowledgements This paper was published on the occasion of the 100th anniversary of Acta Botanica Croatica. We would like to dedicate this review to the late Profes- sor Andrzej Witkowski, an esteemed and passionate diatom taxonomist. We will never forget Andrzej’s enthusiasm, en- ergy, kindness and immense diatom knowledge, which combined historic and modern perspectives and left an in- delible mark worldwide. We will remember him as a noble BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 135 and kind person who was extremely active internationally and whose network spanned numerous countries on most continents. This study is supported by the Croatian Science Foun- dation under project numbers IP-2014-09-2945 (AdMed- Plan), IP-2019-04-9043 (DiVMAd), and UIP 2020-02-7868 (Adri LIFE), and by the Scientific Research Projects Coor- dination Unit of Istanbul University (Grant number: MAB- 2021-37652). The authors thank the researchers who pro- vided literature for this study. This checklist was made possible by the cooperation of many colleagues who pro- vided less accessible literature. We thank the reviewers for their comments, which greatly improved this study. References Accoroni, S., Romagnoli, T., Pichierri, S., Totti, C., 2016: Effects of the bloom of harmful benthic dinoflagellate Ostreopsis cf. ovata on the microphytobenthos community in the northern Adriatic Sea. Harmful Algae 55, 179–190. https://doi. org/10.1016/j.hal.2016.03.003 Akbulut, A., Yıldız, K., 2002: The planktonic diatoms of Lake Çıldır (Ardahan-Turkey). Turkish Journal of Botany 26(2), 55–75. https://journals.tubitak.gov.tr/botany/vol26/iss2/1 Aleem, A. A., 1950: Some new records of marine algae from the Mediterranean Sea. With reference to their geographical dis- tribution. Acta Horti Gothoburgensis 18, 276–288. Altuğ, G., Aktan, Y., Oral, M., Topaloğlu, B., Dede, A., Keskin, Ç., İşinibilir, M., Çardak, M., Çiftçi, P. S., 2011: Biodiversity of the northern Aegean Sea and southern part of the Sea of Marmara, Turkey. Marine Biodiversity Records 4, E65. https://doi.org/10.1017/S1755267211000662 Alverson, A. J., 2008: Molecular systematics and the diatom ape- cies. Protist 159, 339–353. https://doi.org/10.1016/j.pro- tis.2008.04.001 Aragão, L., Mentaschi, L., Pinardi, N., Verri, G., Senatore, A., Di Sabatino, S., 2024: The freshwater discharge into the Adri- atic Sea revisited. Frontiers in Climate 6: 1368456. https://doi. org/10.3389/fclim.2024.1368456 Aslan, H., Gönülal, O., Can-Yılmaz, E., Çamur Elipek, B., Baytut, Ö., Tosunoğlu, M., Karabacak, E. Kurt, Y., 2018: Species di- versity in lentic, lotic, marine and terrestrial biotopes of Gok- ceada Salt Lake wetland (Canakkale, Turkiye). Fresenius En- vironmental Bulletin 27(5), 2853–2866. Baldassarre, L., Natali, V., De Pascale, F., Vezzi, A., Banchi, E., Bazzaro, M., Relitti, F., Tagliapietra, D., Cibic, T., 2023: The Impact of MOSE (Experimental Electromechanical Module) flood barriers on microphytobenthic community of the Ven- ice Lagoon. Microorganisms 11(4), 936. https://doi. org/10.3390/microorganisms11040936 Barbiero, R. P., 2000: A multi–lake comparison of epilithic diatom communities on natural and artificial substrates. Hydrobiolo- gia 438, 157–170. https://doi.org/10.1023/A:1004182231973 Barinova, S., Bondarenko, A., Ryabushko, L., Kapranov, S., 2019: Microphytobenthos as an indicator of water quality and or- ganic pollution in the western coastal zone of the Sea of Azov. Oceanological and Hydrobiological Studies 48(2), 125–139. https://doi.org/10.1515/ohs-2019-0013 Barranguet, C., 1997: The role of microphytobenthic primary production in a Mediterranean mussel culture area. Estua- rine, Coastal and Shelf Science 44(6), 753–765. Barranguet, C., Plante-Cuny, M. R., Alivon, E., 1996: Microphy- tobenthos production in the Gulf of Fos, French Mediterra- nean coast. Hydrobiologia 333, 181–193. https://doi. org/10.1007/BF00013432 Bartole, L., Welker, C., Dellavalle, G., Bressan, G., 1991–1994: Primi stadi di colonizzazione “a diatomee” di un substrato duro artificiale in due stazioni fisse del Golfo di Trieste (nord Adriatico). Nova Thalassia 12, 163–189. Batistić, M., Garić, R., Molinero, J. C., 2014: Interannual varia- tions in Adriatic Sea zooplankton mirror shifts in circulation regimes in the Ionian Sea. Climate Research 61(3), 231–240. https://doi.org/10.3354/cr01248 B-Béres, V., Stenger-Kovács, C., Buczkó, K., Padisák, J., Selmeczy, G. B., Lengyel, E., Tapolczai, K. 2023: Ecosystem services pro- vided by freshwater and marine diatoms. Hydrobiologia 850, 2707–2733. https://doi.org/10.1007/s10750-022-04984-9 Belegratis, M. R., 2002: Studies on periphytic algae in marine sites of Evoikos Gulf. PhD Thesis. University of Athens, Greece. Belegratis, M. R., Economou-Amilli, A., 2002: Seasonal and ver- tical distribution of marine epiphytic algae in the Evoikos Gulf (Aegean Sea, Greece). Algological Studies 107, 131–151. https://doi.org/10.1127/algol_stud/107/2002/131 Blanco, I. A., Blanco, S., 2014: Benthic diatoms from Mediterra- nean coasts. Bibliotheca Diatomologica 60, 1–409. Box, A., Sureda, A., Tauler, P., Terrados, J., Marbà, N., Pons, A., Deudero, S., 2010: Seasonality of caulerpenyne content in na- tive Caulerpa prolifera and invasive C. taxifolia and C. race- mosa var. cylindracea in the western Mediterranean Sea. Bo- tanica Marina 53(4), 367–375. https://doi.org/10.1515/ BOT.2010.034 Box, A., Sureda, A., Terrados, J., Pons, A., Deudero, S., 2008: An- tioxidant response and caulerpenyne production of the alien Caulerpa taxifolia (Vahl) epiphytized by the invasive algae Lophocladia lallemandii (Montagne). Journal of Experimen- tal Marine Biology and Ecology 364, 24–28. https://doi. org/10.1016/j.jembe.2008.06.029 Burić, Z., Caput, K., Viličić, D., 2004: Distribution of the diatom Cocconeis scutellum in the karstic estuary (Zrmanja, eastern Adriatic Sea). Biologia, Bratislava 59(1), 1–7. Caput Mihalić, K., Viličić, D., Ahel, M., Burić, Z., Carić, M., 2008: Periphytic algae development in the upper reach of the Zr- manja estuary (eastern Adriatic coast). Vie et Milieu 58(3-4), 203–213. Car, A., Hafner, D., Dupčić Radić, I., Kaleli, A., Ljubimir, S., So- lak, C. N., 2021: Comparison of benthic diatom community structures on natural and artificial substrates in marine lake (Adriatic Sea). Acta Adriatica: international journal of Ma- rine Sciences 62(1), 21–44. https://doi.org/10.32582/aa.62.1.1 Car, A., Hafner, D., Ljubimir, S., Dupčić Radić, I., Bobanović- Ćolić, S., Jasprica, N., 2020: Colonization of bacteria and dia- toms on an artificial substrate in a marine lake (eastern Adri- atic Sea, NE Mediterranean). Acta Botanica Croatica 79(2), 212–227. https://doi.org/10.37427/botcro-2020-028 Car, A., Witkowski, A., Dobosz, S., Burfeind, D.D., Meinesz, A., Jasprica, N., Ruppel, M., Kurzydłowski, K. J., Płociński, T., 2012: Description of a new marine diatom, Cocconeis cauler- pacola sp. nov. (Bacillariophyceae), epiphytic on invasive Caulerpa species. European Journal of Phycology 47(4), 433– 448. https://doi.org/10.1080/09670262.2012.735255 Car, A., Witkowski, A., Dobosz, S., Jasprica, N., Ljubimir, S., 2019a: Epiphytic diatom assemblages on invasive algae Caul- erpa taxifolia and autochthonous Halimeda tuna and Padina sp. from the Adriatic Sea – summer/autumn aspect. Oceano- logical and Hydrobiological Studies 48(3), 209–226. https:// doi.org/10.2478/ohs-2019-0019 Car, A., Witkowski, A., Jasprica, N., Ljubimir, S., Čalić, M., Do- bosz, S., Dupčić Radić, I., Hrustić, E., 2019b: Epilithic diatom communities from areas of invasive Caulerpa species (Caul- https://doi.org/10.1127/algol_stud/107/2002/131 https://doi.org/10.2478/ohs-2019-0019 https://doi.org/10.2478/ohs-2019-0019 CAR A., KALELI A. 136 ACTA BOT. CROAT. 84 (2), 2025 erpa taxifolia and Caulerpa cylindracea) in the Adriatic Sea, NE Mediterranean. Mediterranean Marine Science 20(1), 151–173. https://doi.org/10.12681/mms.14330 Caraus, I., 2017: Algae of Romania. A distributional checklist of actual algae. Version 2.4. Studii si Cercetari Biologie 7, 1–1002. Carreira-Flores, D., Neto, R., Ferreira, H., Cabecinha, E., Díaz- Agras, G., Gomes, P. T., 2020: Artificial substrates as sam- pling devices for marine epibenthic fauna: A quest for stan- dardization. Regional Studies in Marine Science 37, 1–11. https://doi.org/10.1016/j.rsma.2020.101331 Cibic, T., Baldassarre, L., Cerino, F., Comici, C., Fornasaro, D., Kralj, M., Giani, M., 2022: Benthic and pelagic contributions to primary production: experimental insights from the Gulf of Trieste (Northern Adriatic Sea). Frontiers in Marine Sci- ence 9, 877935. https://doi.org/10.3389/fmars.2022.877935 Cibic, T., Blasutto, O., 2011: Living marine benthic diatoms as in- dicators of nutrient enrichment: a case study in the Gulf of Trieste. In: Compton, J. C. (ed.), Diatoms: ecology and life cycle, 169–184. Nova Science Publishers, Hauppauge, New York. Cibic, T., Blasutto, O., Falconi, C., Fonda Umani, S., 2007a: Mi- crophytobenthic biomass, species composition and nutrient availability in sublittoral sediments of the Gulf of Trieste (northern Adriatic Sea). Estuarine, Coastal and Shelf Science 75(1-2), 50–62. https://doi.org/10.1016/j.ecss.2007.01.020 Cibic, T., Blasutto, O., Fonda Umani, S., 2007b: Biodiversity of settled material in a sediment trap in the Gulf of Trieste (northern Adriatic Sea). Hydrobiologia 580, 57–75. https:// doi.org/10.1007/s10750-006-0465-9 Cibic, T., Comici, C., Bussani, A., Del Negro, P., 2012: Benthic diatom response to changing environmental conditions. Es- tuarine, Coastal and Shelf Science 115, 158–169. https://doi. org/10.1016/j.ecss.2012.03.033 Cibic, T., Facca, C., 2010: Microphytobenthos. In: Relini, G. (ed.), Checklist della flora e della fauna dei mari italiani. Parte II. Biologia Marina Mediterranea 17 (suppl. 1), 754–800. Cibic, T., Fazi, S., Nasi, F., Pin, L., Alvisi, F., Berto, D., Viganò, L., Zoppini, A., Del Negro, P., 2019: Natural and anthropogenic disturbances shape benthic phototrophic and heterotrophic microbial communities in the Po River Delta system. Estua- rine, Coastal and Shelf Science 222, 168–182. https://doi. org/10.1016/j.ecss.2019.04.009 Civitarese, G., Gačić, M., Batistić, M., Bensi, M., Cardin, V., Dulčić, J., Garić, R., Menna, M., 2023: The BiOS mechanism: History, theory, implications. Progress in Oceanography 216, 103056. https://doi.org/10.1016/j.pocean.2023.103056 Civitarese, G., Gačić, M., Lipizer, M., Eusebi Borzelli, G. L., 2010: On the impact of the Bimodal Oscillating System (BiOS) on the biogeochemistry and biology of the Adriatic and Ionian Seas (Eastern Mediterranean). Biogeosciences 7, 3987–3997. https://doi.org/10.5194/bg-7-3987-2010 Cleve, P.T., Grunow, A. 1880: Beiträge zur kenntniss der arctisch- en diatomeen. Kongliga Svenska vetenskaps-Akademiens Handlingar 17(2), 1–121. Çelekli, A., Külköylüoğlu, O., 2006: Net planktonic piatom (Bac- illariophyceae) composition of Lake Abant (Bolu). Turkish Journal of Botany 30(5), 331–347. https://journals.tubitak.gov. tr/botany/vol30/iss5/2 Çolak Sabancı, F., 2010: Contributions to the knowledge of algal flora of Homa Lagoon (Aegean Sea, Turkey). Journal of Black Sea / Mediterranean Environment 16(3), 311–327. Retrieved October 16, 2023 from https://dergipark.org.tr/en/pub/jbme/ issue/9843/121871 Çolak Sabancı, F., 2011: Relationship of epilithic diatom commu- nities to environmental variables in Homa lagoon (Izmir, Turkey). Aquatic Biology 13(3), 233–241. https://doi. org/10.3354/ab00367 Çolak Sabancı, F., 2012a: An illustrated survey on the morpho- logical characters in three species of the diatom genus Mas- togloia (Bacillariophyceae). Turkish Journal of Botany 36(6), 727–737. https://doi.org/10.3906/bot-1111-18 Çolak Sabancı, F., 2012b: Taxonomic survey of benthic diatoms on natural substrata from coastal lagoon (Aegean Sea, Tur- key). Turkish Journal of Fisheries and Aquatic Sciences 12(4), 841–849. https://doi.org/10.4194/1303-2712-v12_4_12 Çolak Sabancı, F., 2013: Species of Mastogloia (Bacillariophyceae) - new for the Aegean coast of Turkey. Mediterranean Marine Science 14(1), 129–140. https://doi.org/10.12681/mms.331 Çolak Sabancı, F., Koray, T., 2010: Four new records for the ben- thic diatoms (genera Cocconeis, Seminavis, Synedra, and Tra- chysphenia) from the Aegean Sea. Turkish Journal of Botany 34(6), 531–540. https://doi.org/10.3906/bot-0912-285 Cox, E. J., 2014: Diatom identification in the face of changing spe- cies concepts and evidence of phenotypic plasticity. Journal of Micropalaeontology 33, 111–120, https://doi.org/10.1144/ jmpaleo2014-014 Cozzi, S., Giani, M., 2011: River water and nutrient discharges in the Northern Adriatic Sea: Current importance and long term changes. Continental Shelf Research 31(18), 1881–1893. https://doi.org/10.1016/j.csr.2011.08.010 Delgado, M., 1989: Abundance and distribution of microphyto- benthos in the bays of Ebro Delta (Spain). Estuarine, Coastal and Shelf Science 29(2), 183–194. https://doi.org/10.1016/0272- 7714(89)90007-3 Delgado, M., De Jonge, V. N., Peletier, H., 1991: Effect of sand movement on the growth of benthic diatoms, Journal of Ex- perimental Marine Biology and Ecology 145(2), 221–231. https://doi.org/10.1016/0022-0981(91)90177-X De Stefano, M., Marino. D., 2003: Morphology and taxonomy of Amphicocconeis gen. nov. (Achnanthales, Bacillariophyceae, Bacillariophyta) with considerations on its relationship to other monoraphid diatom genera. European Journal of Phy- cology 38(4), 361–370. https://doi.org/10.1080/096702603100 01612646 De Stefano, M., Marino, D., Mazzella, L., 2000: Marine taxa of Cocconeis on leaves of Posidonia oceanica, including a new species and two new varieties. European Journal of Phycol- ogy 35(3), 225–242. https://doi.org/10.1080/09670260010001 735831 De Stefano, M., Romero, O. E., Totti, C., 2008: Ultrastructure and taxonomy of Cocconeis scutellum and its varieties. Botanica Marina 51(6), 506–536. https://doi.org/10.1515/BOT.2008.058 De Stefano, M., Sacchi, U., Totti, C., Romero, O. E., 2006: Cocco- neis distans Gregory and Amphicocconeis debesi (Hustedt) De Stefano comb. nov. (Bacillariophyta), an intricate taxonomi- cal history. Botanica Marina 49(5-6), 438–449. https://doi. org/10.1515/BOT.2006.055 Economou-Amilli, A., 1980: Marine diatoms from Greece. I. Di- atoms from the Saronikos Gulf. Nova Hedwigia 32(1), 63– 104. Elmacı, A., Obalı, O., 1998: Aksehir gölü kıyı bölgesi alg florası (The algal flora of the littoral region in Lake Akşehir). Turk- ish Journal of Biology 22(1), 81–98. Facca, C., Sfriso, A., 2007: Epipelic diatom spatial and temporal distribution and relationship with the main environmental parameters in coastal waters. Estuarine, Coastal and Shelf Science 75(1-2), 35–49. https://doi.org/10.1016/j. ecss.2007.03.033 Facca, C., Sfriso, A., Ghetti, P. F., 2004: Abbondanza e diversità del fitoplancton e delle diatomee bentoniche in laguna di Venezia. Biologia Ambientale 18(2), 19–24. Retrieved October BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 137 16, 2023 from http://www.cisba.eu/images/rivista/biologia_ ambientale/Ba2004-2/Ba_2004-2_Facca_Diatomee_VE.pdf Facca, C., Sfriso, A., Socal, G., 2002a: Temporal and spatial dis- tribution of diatoms in the surface sediments of the Venice Lagoon. Botanica Marina 45(2), 170–183. https://doi. org/10.1515/BOT.2002.016 Facca, C., Sfriso, A., Socal, G., 2002b: Changes in abundance and composition of phytoplankton and microphytobenthos due to increased sediment fluxes in the Venice Lagoon, Italy. Es- tuarine, Coastal and Shelf Science 54(5), 773–792. https://doi. org/10.1006/ecss.2001.0848 Facca, C., Sfriso, A., Socal, G., 2003: Diatomee planctoniche e bentoniche: abbondanza e diversità nella laguna di Venezia. Biologia Marina Mediterranea 10(2), 1006–1009. Fain, A. M. V., Ogston, A. S., Sternberg, R. W., 2007: Sediment transport event analysis on the western Adriatic continental shelf. Continental Shelf Research 27(3-4), 431–451. Falkowski, P. G., Schofield, O., Katz, M. E., Van De Schootbrugge, B., Knoll, A. H., 2004: Why is the land green and the ocean red? In: Thierstein, H. R. and Young, J. R. (eds.), Coccolitho- phores: from molecular processes to global impact, 429–453. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3- 662-06278-4_16 Foged, N., 1985a: Diatoms in Samos, a Greek island in the Ae- gean. Bibliotheca Diatomologica 10, 1-119. Foged, N., 1985b: Diatoms in Kos and Kalymnos, two Greek is- lands in the Aegean. Bibliotheca Diatomologica 10, 1–105. Franzo, A., Cibic, T., Del Negro, P., De Vittor, C., 2015: Spatial distribution of microphytobenthos, meiofauna and macro- fauna in the north-western Adriatic Sea: a synoptic study. Advances in Oceanography and Limnology 6(1-2), 58–75. https://doi.org/10.4081/aiol.2015.5470 Franzo, A., Cibic, T., Del Negro, P., Solidoro, C., 2014: Microphy- tobenthic response to mussel farm biodeposition in coastal sediments of the northern Adriatic Sea. Marine Pollution Bulletin 79(1-2), 379–388. https://doi.org/10.1016/j.marpol- bul.2013.11.002 Gačić, M., Kovačević, V., Manca, B., Papageorgiou, E., Poulain, P.-M., Scarazzato, P., Vetrano, A., 1996: Thermohaline prop- erties and circulation in the Otranto Strait. Bulletin de l’Institut oceanographique 17, 117–145. Gambi, C., Totti, C., Manini, E., 2003: Impact of organic loads and environmental gradients on microphytobenthos and meiofau- nal distribution in a coastal lagoon. Chemistry and Ecology 19(2-3), 207–223. https://doi.org/10.1080/0275754031000119951 Gligora Udovič, M., Ljubešić, Z., 2021: The Croatian National Diatom Collection. Acta Botanica Croatica, 80(1). Retrieved October 7, 2024 from https://www.abc.botanic.hr/index.php/ abc/article/view/3184 Grunow, A., 1860: Ueber neue oder ungenügend gekannte Algen. Erste folge, diatomeen, familie Naviculaceen. Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesell- schaft in Wien 10, 503–582. Guiry, M. D., Guiry, G. M., 2024: AlgaeBase. World-wide elec- tronic publication, National University of Ireland, Galway. Retrieved October 18, 2024 from http://www.algaebase.org Hafner, D., Jasprica, N., Car, A., 2018a: Taxonomic survey of ben- thic diatoms in Neum Bay, southeastern Adriatic. Natura Croatica 27(1), 1–26. https://doi.org/10.20302/NC.2018.27.1 Hafner, D., Car, A., Jasprica, N., Kapetanović, T., Dupčić Radić, I., 2018b: Relationship between marine epilithic diatoms and environmental variables in oligotrophic bay, NE Mediterra- nean. Mediterranean Marine Science 19(2), 223–239. https:// doi.org/10.12681/mms.14151 Hendey, N. I., 1974: A revised check-list of British marine dia- toms. Journal of the Marine Biological Association of the United Kingdom 54, 277–300. Hoagland, K. D., Zlotsky, A., Peterson, C. G., 1986: Chapter 2 The source of algal colonizers on rock substrates in a freshwater impoundment. In: Evans, L. V., Hoagland, K. D. (eds.), Algal biofouling, 21–39. Elsevier, Amsterdam, https://doi. org/10.1016/S0166-1116(08)72168-X Jahn, R., Zetzsche, H., Reinhardt, R., Gemeinholzer, B., 2007: Diatoms and DNA barcoding: a pilot study on an environ- mental sample. In: Kusber W. H., Jahn R. (eds.), Proceedings of the 1st Central European Diatom Meeting 2007. Berlin, 63–68. Botanic Garden and Botanical Museum Berlin- Dahlem, Freie Universität Berlin, Berlin. Kaddeche, H., Noune, F., Dzizi, S., Chaib, N., Boudjellab, Z. E., Blanco, S., 2022: Determinant factors of diatom assemblage’s distribution along the coastal Central Constantine (North- eastern Algeria). Aquatic Ecology 56, 1245–1269. https://doi. org/10.1007/s10452-022-09980-8 Kaleli, A., 2019: Benthic diatom composition of Iztuzu coastal lake, Dalyan (Aegean Sea, Turkey). Aquatic Sciences and En- gineering 34(4), 122–130. https://doi.org/10.26650/ ASE2019575987 Kaleli, A., 2022: Contributions to the diatom flora of the Aegean Sea and the Mediterranean Sea from the turkish coasts with remarks on rare taxa. Mediterranean Marine Science 23(4), 952–967. https://doi.org/10.12681/mms.28075 Kaleli, A., Akçaalan, R., 2021: Checklist of marine diatoms from Turkish coastal waters with updated nomenclature. Aquatic Research 4(1), 88–115. https://doi.org/10.3153/AR21008 Kaleli, A., Car, A., Witkowski, A., Krzywda, M., Riaux-Gobin, C., Solak, C. N., Kaska, Y., Zgłobicka, I., Płociński, T., Wróbel, R., Kurzydłowski, K., 2020: Biodiversity of carapace epibiont diatoms in lggerhead sea turtles (Caretta caretta Linnaeus 1758) in the Aegean Sea Turkish coast. PeerJ 8: e9406. https:// doi.org/10.7717/peerj.9406 Kanjer, L., Mucko, M., Car, A., Bosak, S., 2019: Epiphytic diatoms on Posidonia oceanica (l.) Delile leaves from eastern Adriatic Sea. Natura Croatica 28(1), 1–20. https://doi.org/10.20302/ NC.2019.28.1 Kociolek, J. P., Balasubramanian, K., Blanco, S., Coste, M., Ector, L., Liu, Y., Kulikovskiy, M., Lundholm, N., Ludwig, T., Pota- pova, M., Rimet, F., Sabbe, K., Sala, S., Sar, E., Taylor, J., Van De Vijver, B., Wetzel, C. E., Williams, D. M., Witkowski, A., Witkowski, J., 2024, DiatomBase. Retrieved February 23, 2024 from http://www.diatombase.org Konucu, M., Develi, E. E., 2021: New records of pennate diatom species from the Mediterranean Sea. Mediterranean Fisher- ies and Aquaculture Research 4(2), 43–51. Retrieved October 16, 2023 from https://dergipark.org.tr/en/pub/medfar/is- sue/62622/868097 Koray, T., 2001: Türkiye denizleri fitoplankton türleri kontrol lis- tesi (A check-list for phytoplankton of Turkish seas). Ege Journal of Fisheries and Aquatic Sciences 18(1-2), 1–23. Re- trieved October 16, 2023 from https://dergipark.org.tr/en/ pub/medfar/issue/62622/868097 Kützing, F. T., 1844: Die Kieselschaligen. Bacillarien oder Diato- meen. Nordhausen, Germany. Kützing, F. T. 1845: Phycologia germanica, d.i. Deutschlands al- gen in bündigen beschreibungen. Nebst einer anleitung zum untersuchen und bestimmen dieser gewächse für anfäger. Nordhausen, Germany. Lamberti, G. A., Resh, V. H., 1985: Distribution of benthic algae and macroinvertebrates along a thermal stream gradient. Hy- drobiologia. 128, 13–21. https://doi.org/10.1007/BF00008935 https://www.sciencedirect.com/journal/estuarine-coastal-and-shelf-science https://www.sciencedirect.com/journal/estuarine-coastal-and-shelf-science https://doi.org/10.1006/ecss.2001.0848 https://doi.org/10.1006/ecss.2001.0848 CAR A., KALELI A. 138 ACTA BOT. CROAT. 84 (2), 2025 Lane, C. M., Taffs, K. H., Corfield, J. L., 2003: A comparison of diatom community structure on natural and artificial sub- strates. Hydrobiologia 493, 65–79. https://doi. org/10.1023/A:1025498732371 Levkov, Z., Caput Mihalić, K., Ector, L., 2010: A taxonomical study of Rhoicosphenia Grunow (Bacillariophyceae) with a key for identification of selected taxa. Fottea 10(2), 145–200. https://doi.org/10.5507/fot.2010.010 Li, C., Gastineau, R., Turmel, M., Witkowski, A., Otis, C., Car, A., Lemieux, C., 2019: Complete chloroplast genome of the tiny marine diatom Nanofrustulum shiloi (Bacillariophyta) from the Adriatic Sea. Mitochondrial DNA Part B: Resources 4(2), 3374–3376. https://doi.org/10.1080/23802359.2019.1673245 Li, C. L., Witkowski, A., Ashworth, M. P., Dąbek, P., Sato, S., Zgłobicka, I., Witak, M., Khim, J. S., Kwon, C. J., 2018: The morphology and molecular phylogenetics of some marine di- atom taxa within the Fragilariaceae, including twenty unde- scribed species and their relationship to Nanofrustulum, Opephora and Pseudostaurosira. Phytotaxa 355(1), 1–104. https://doi.org/10.11646/phytotaxa.355.1.1 Lobarzewski [Lobarzewsky], H. R. von, 1840: Einige neue dia- tomaceen Agardh et auct. der ostküste des Adriatischen meeres. Linnaea 14: 265–277. Lobban, C. S., Ashworth, M. P., Car, A., Herwig, W., Ulanova, A., 2015: Licmosphenia revisited: transfer to Licmophora, rede- scription of L. clevei Mereschkowsky and descriptions of three new species. Diatom research 30(3), 227–236. https:// doi.org/10.1080/0269249X.2015.1045037 Loir, M., 2010–2014. Marine benthic diatoms. Retrieved August 16, 2023 from http://www.diatomloir.eu/Site Diatom/Com- munities.html. Louvrou, I., 2007: Periphyton and its colonization in marine hy- drothermal regions of island Milos (Greece). PhD, University of Athens, Greece. Louvrou, I, Danielidis, D. B., Economou-Amilli, A., 2006. Taxo- nomic status of Detonia Frenguelli and the establishment of Detonia dobrinae sp. nov. (Bacillariophyceae). Ιn: Ognjanova Rumenova, N., Manoylov, K. (eds.), Advances in Phycologi- cal Studies: Festschrift in honour of Prof. Dobrina Temnis- kova-Topalova. 123–131. Pensoft Publisher, St. Kliment Ohridski University Press, Sofia, Bulgaria. Louvrou, I., Danielidis, D. B., Economou-Amilli, A., 2012: Melo- neis gen. nov., a new epipsammic genus of Rhaphoneidaceae (Bacillariophyceae). PLoS ONE 7(3), e32198. https://doi. org/10.1371/journal.pone.0032198 Ljubimir, S., Jasprica, N., Čalić, M., Hrustić, E., Dupčić Radić, I., Car, A., Batistić, M., 2017: Interannual (2009–2013) variabil- ity of winter-spring phytoplankton in the open South Adri- atic Sea: effects of deep convection and lateral advection. Continental Shelf Research 143, 311–321. https://doi. org/10.1016/j.csr.2017.05.007 MacIntyre, H. L., Geider, R. J., Miller, D. C., 1996: Microphytob- enthos: The ecological role of the “secret garden” of unvege- tated, shallow-water marine habitats. I. Distribution, abun- dance and primary production. Estuaries 19, 186–201. https:// doi.org/10.2307/1352224 Majewska, R., D’alelio, D., De Stefano, M., 2014: Cocconeis Eh- renberg (Bacillariophyta), a genus dominating diatom com- munities associated with Posidonia oceanica Delile (mono- cotyledons) in the Mediterranean Sea. Aquatic Botany 112, 48–56. https://doi.org/10.1016/j.aquabot.2013.07.008 Matek, A., Ljubešić, Z., 2024: Primary production research in the Adriatic Sea - a review. Acta Adriatica 65(2). https://doi. org/10.32582/aa.65.2.1 Mazzella, L., Buia, M. C., Spinoccia, L., 1994: Biodiversity of ep- iphytic diatom community on leaves of Posidonia oceanica. In: Marino, D., Montresor, M. (eds.), Proceedings of the 13th International Diatom Symposium, 241–251. Biopress, Bristol. Medlin, L. K., 2018: Mini review: diatom species as seen through a molecular window. Brazilian Journal of Botany 41, 457–469. https://doi.org/10.1007/s40415-018-0444-1 Medlin, L. K., Kaczmarska, I., 2004: Evolution of the diatoms: V. Morphological and cytological support for the major clades and a taxonomic revision. Phycologia 43(3), 245–270. https:// doi.org/10.2216/i0031-8884-43-3-245.1 Mejdandžić, M., Ivanković, T., Pfannkuchen, M., Godrijan, J., Marić Pfannkuchen, D., Hrenović, J., Ljubešić, Z., 2015: Col- onization of diatoms and bacteria on artificial substrates in the northeastern coastal Adriatic Sea. Acta Botanica Croati- ca 74(2), 407–422. https://doi.org/10.1515/botcro-2015-0030 Menna, M., Gačić, M., Martellucci, R., Notarstefano, G., Fedele, G., Mauri, E., Gerin, R., Poulain, P.-M., 2022: Climatic, decadal, and interannual variability in the upper layer of the Mediterranean Sea using remotely sensed and in-situ data. Remote Sensing. 14, 1322. https://doi.org/10.3390/rs14061322 Miho, A., Witkowski, A., 2005: Diatom (Bacillariophyta) flora of Albanian coastal wetlands taxonomy and ecology: A review. Proceedings of the California Academy of Sciences 56(12), 129–145. Morales, E. A., Wetzel, C. E., Novais, M. H., Buczkó, K., Morais, M. M., Ector, L., 2019: Morphological reconsideration of the araphid genus Pseudostaurosira (Bacillariophyceae), a revi- sion of Gedaniella, Popovskayella and Serratifera, and a de- scription of a newNanofrustulum species. Plant Ecology and Evolution 152(2), 262–284. https://doi.org/10.5091/plece- vo.2019.1604 Munda, I. M., 2005: Seasonal fouling by diatoms on artificial sub- strata at different depths near Piran (Gulf of Trieste, North- ern Adriatic). Acta Adriatica 46(2), 137–157. https://hrcak. srce.hr/86 Myers, N., Mittermeier, R. A., Mittermeier, C. G., Da Fonseca, G. A., Kent, J., 2000: Biodiversity hotspots for conservation pri- orit ies. Nature 403(6772), 853–858. https://doi. org/10.1038/35002501 Natali, V., Malfatti, F., Cibic, T., 2023: Ecological effect of different- ly treated wooden materials on microalgal biofilm formation in the Grado Lagoon (Northern Adriatic Sea). Microorganisms 11, 2196. https://doi.org/10.3390/microorganisms11092196 Nenadović, T., Šarčević, T., Čižmek, H., Godrijan, J., Marić Pfannkuchen, D., Pfannkuchen, M., Ljubešić, Z., 2015: De- velopment of periphytic diatoms on different artificial sub- strates in the Eastern Adriatic Sea. Acta Botanica Croatica 74(2), 377–392. https://doi.org/10.1515/botcro-2015-0026 Neri, F., Ubaldi, M., Accoroni, S., Ricci, S., Banchi, E., Romag- noli, T., Totti, C., 2024: Comparative analysis of phytoplank- ton diversity using microscopy and metabarcoding: insights from an eLTER station in the Northern Adriatic Sea. Hydro- biologia. https://doi.org/10.1007/s10750-024-05692-2 Park, J. S., Lobban, C. S., Lee, K. W., 2018: Diatoms associated with seaweeds from Moen Island in Chuuk Lagoon, Micro- nesia. Phytotaxa 351(2), 101–140. https://doi.org/10.11646/ phytotaxa.351.2.1 Pennesi, C., Danovaro, R., 2017: Assessing marine environmental status through microphytobenthos assemblages colonizing the Autonomous Reef Monitoring Monitoring Structures (ARMS) and their potential in coastal marine restoration. Marine Pollution Bulletin 125(1-2), 56–65. https://doi. org/10.1016/j.marpolbul.2017.08.001 Pennesi, C., Poulin, M., De Stefano, M., Romagnoli, T., Totti, C., 2011: New insights to the ultrastructure of some marine Mas- togloia species section Sulcatae (Bacillariophyceae), including https://doi.org/10.1371/journal.pone.0032198 https://doi.org/10.1371/journal.pone.0032198 BENTHIC DIATOMS FROM THE ADRIATIC SEA ACTA BOT. CROAT. 84 (2), 2025 139 Mastogloia neoborneensis sp. nov. Phycologia 50(5), 548–562. https://doi.org/10.2216/10-39.1 Pennesi, C., Poulin, M., De Stefano, M., Romagnoli, T., Totti, C., 2012: Morphological studies of some marine Mastogloia (Bacillariophyceae) belonging to the section sulcatae, includ- ing the description of new species. Journal of Phycology 48(5), 1248–1264. https://doi.org/10.1111/j.1529-8817.2012.01215.x Pérez-Burillo, J., Valoti, G., Witkowski, A., Prado, P., Mann, D. G., Trobajo, R., 2022: Assessment of marine benthic diatom communities: insights from a combined morphological–me- tabarcoding approach in Mediterranean shallow coastal wa- ters. Marine Pollution Bulletin 174, 113183. https://doi. org/10.1016/j.marpolbul.2021.113183 Poulain, P. M., 2001: Adriatic Sea surface circulation as derived from drifter data between 1990 and 1999. Journal of Marine Systems 29, 3–32. https://doi.org/10.1016/S0924-7963(01) 00007-0 Ravaioli, M., Alvisi, F., Menegazzo-Vitturi, L., 2003: Dolomite as a tracer for sediment transport and deposition on the north- western Adriatic continental shelf (Adriatic Sea, Italy). Con- tinental Shelf Research 23, 1359–1377. https://doi.org/10.1016/ S0278-4343(03)00121-3 Riaux-Gobin, C., Vétion, G., Neveux, J., Duchêne, J.-C., 1998: Mi- crophytobenthos et phytoplancton en baie de Banyuls (Golfe du Lion): Biomasses pigmentaires et facteurs hydroclima- tiques. Vie et Milieu 48(1), 1–13. Ribeiro, L., Figueira, R., Oiry, S., Buchet, R., Sabbe, K., Barillé, L., 2022: Benthic marine diatoms from French coastal and tran- sitional waters. Version 1.5. Université de Nantes. Retrieved October 10, 2024 from https://doi.org/10.15468/byyhxu ac- cessed via GBIF.org Rogelja, M., Cibic, T., Belmonte, M., Del Negro, P., 2018: Active and resting microbenthos in differently contaminated ma- rine coastal areas: insights from the Gulf of Trieste (northern Adriatic, Mediterranean Sea). Hydrobiologia 806, 283–301. https://doi.org/10.1007/s10750-017-3366-1 Rogelja, M., Cibic, T., Pennesi, C., De Vittor, De, 2016: Microphy- tobenthic community composition and primary production at gas and thermal vents in the Aeolian Islands (Tyrrhenian Sea, Italy), Marine Environmental Research 118, 31–44. https://doi.org/10.1016/j.marenvres.2016.04.009 Romagnoli, T., Bavestrello, G., Cucchiari, E. M., De Stefano, M., Di Camillo, C. G., Pennesi, C., Puce, S., Totti, C., 2007: Mi- croalgal communities epibiontic on the marine hydroid Eu- dendrium racemosum in the Ligurian Sea during an annual cycle. Marine Biology 151, 537–552. https://doi.org/10.1007/ s00227-006-0487-x Romagnoli, T., Totti, C., Accoroni, S., De Stefano, M., Pennesi, C., 2014: SEM analysis of the epibenthic diatoms on Euden- drium racemosum (Hydrozoa) from the Mediterranean Sea. Turkish Journal of Botany 38(3), 566–594. https://doi. org/10.3906/bot-1305-52 Round, F. E., Crawford, R. M., Mann, D. G., 1990: The Diatoms. Biology and morphology of the genera. Cambridge Univer- sity Press, Cambridge. Rubino, F., Cibic, T., Belmonte, M., Rogelja, M., 2016: Microben- thic community structure and trophic status of sediments in the Mar Piccolo of Taranto (Mediterranean, Ionian Sea). En- vironmental Science and Pollution Research 23, 12624– 12644. https://doi.org/10.1007/s11356-015-5526-z Sdrigotti, E., Barbariol, V., Welker, C., 1999: Diatom assemblages in coastal shallow waters at the water-sediment interface (Gulf of Trieste, North Adriatic Sea). Annales. Series historia naturalis 9(17), 191–202. Retrieved October 16, 2023 from https://www.dlib.si/details/URN:NBN:SI:DOC-BW6NZCZP Seveno, J., Car, A., Sirjacobs, D., Fullgrabe, L., Dupčić Radić, I., Lejeune, P., Leignel, V., Mouget, J-L., 2023: Benthic diatom blooms of blue Haslea spp. in the Mediterranean Sea. Marine Drugs 21(11), 583. https://doi.org/10.3390/md21110583 Seveno, J., Witkowski, A., Car, A., Gastineau, R., Sirjacobs, D., Leignel, V., Mouget, J-L., 2024: Impact of blue Haslea spp. blooms on benthic diatom and bacterial communities. Phy- cology 4(3), 465–507. https://doi.org/10.3390/phycolo- gy4030027 Sıvacı, E. R., Kilinc, S., Dere, Ş., 2007: Relationships between epipelic diatom taxa (Bacillariophyceae) and nutrients in two karstic lakes in central Anatolia, Turkey. Journal of Freshwa- ter Ecology 22(3), 515–520. https://doi.org/10.1080/02705060 .2007.9664181 Sorhannus, U., 2007: A nuclear-encoded small-subunit ribosom- al RNA timescale for diatom evolution. Marine Micropale- ontology 65(1-2), 1–12. https://doi.org/10.1016/j.marmi- cro.2007.05.002 Souffreau, C., Verbruggen, H., Wolfe, A. P., Vanormelingen, P., Siver, P. A., Cox, E. J., Mann, D. G., Van de Vijver, B., Sabbe, K., Vyverman, W., 2011: A time-calibrated multi-gene phy- logeny of the diatom genus Pinnularia. Molecular Phyloge- netics and Evolution 61(3), 866–879. https://doi.org/10.1016/j. ympev.2011.08.031 Stevenson, R. J., Pan, Y., 1999: Assessing environmental conditions in rivers and streams with diatoms. In: Stoermer, E. F., Smol, J. P. (eds.), The Diatoms: applications for the environmental and earth sciences. 11–40. Cambridge University Press, New York. https://doi.org/10.1017/CBO9780511613005.003 Tas, S., 2013: Phytoplankton composition and abundance in the coastal waters of the Datça and Bozburun Peninsulas, south- eastern Aegean Sea (Turkey). Mediterranean Marine Science 15(1), 84–94. https://doi.org/10.12681/mms.418 Theriot, E. C., Ashworth, M., Ruck, E., Nakov, T., Jansen, R. K., 2010: A preliminary multigene phylogeny of the diatoms (Bacillariophyta): challenges for future research. Plant Ecol- ogy and Evolution 143(3), 278–296. https://doi.org/10.5091/ plecevo.2010.418 Tolomio, C., 2004: Approche expérimentale pour l'étude des di- atomées des sédiments de la lagune de Venise, Italie. Diatom Research 19(1), 81–101. https://doi.org/10.1080/026924 9X.2004.9705609 Tolomio, C., Andreoli, C., 1989: Recherches sur le periphyton à diatomées dans un vivier de la Lagune de Venise (mai 1984- mai 1985). Diatom Research 4(1), 151–162. https://doi.org/10 .1080/0269249X.1989.9705060 Tolomio, C., Andreoli, C., Avignone, M. T., 1991: Contribution a la connaissance du periphyton diatomique des eaux saumâtres de la Valle Sparesera (Suite). Diatom Research 6(2), 391–399. https://doi.org/10.1080/0269249X.1991.9705183 Tolomio, C., Moro, I., Moschin, E., Valandro, A., 1999: Résultats préliminaires sur les diatomées benthiques de substrats meu- bles dans la lagune de Venise, Italie (Mars 1994-Janvier 1995). Diatom Research 14(2), 367–379. https://doi.org/10.1080/026 9249X.1999.9705478 Tolomio, C., Moschin, E., Duzzin, B., 2002: Distribution des di- atomées benthiques de substrats meubles dans le bassin sud de la lagune de Venise, Italie. Diatom Research 17(2), 401–414. https://doi.org/10.1080/0269249X.2002.9705557 Totti, C., 2003: Influence of the plume of the river Po on the dis- tribution of subtidal microphytobenthos in the northern Adriatic Sea. Botanica Marina 46(2), 161–178. https://doi. org/10.1515/BOT.2003.017 Totti, C., Cucchiari, E., De Stefano, M., Pennesi, C., Romagnoli, T., Bavestrello, G., 2007: Seasonal variations of epilithic dia- toms on different hard substrates, in the northern Adriatic https://doi.org/10.1007/s00227-006-0487-x https://doi.org/10.1007/s00227-006-0487-x CAR A., KALELI A. 140 ACTA BOT. CROAT. 84 (2), 2025 Sea. Journal of the Marine Biological Association of the Unit- ed Kingdom 87(3), 649–658. https://doi.org/10.1017/ S0025315407054665 Traykovski, P., Wiberg, P. L., Geyer, W. R., 2007: Observations and modelling wave-supported sediment gravity flows on the Po prodelta and comparison to prior observations from the Eel shelf. Continental Shelf Research 27(3-4), 357–399. https:// doi.org/10.1016/j.csr.2005.07.008 Tuchman, M. L., Stevenson, R. J., 1980: Comparison of clay tile, sterilized rock, and natural substrate diatom communities in a small stream in Southeastern Michigan, USA. Hydrobiolo- gia 75, 73–79. https://doi.org/10.1007/BF00006564 Van Heurck, H., 1881: Synopsis des Diatomées de Belgique Atlas. Ducaju et Cie, Anvers, Belgium. Velâsquez, Z. R., Cruzado, A., 1995: Inventory of the diatom flo- ra of the Nw Mediterranean Sea. Vie et Milieu / Life & Envi- ronment 249–263. Verri, G., Furnari, L., Gunduz, M., Senatore, A., Santos da Costa, V., De Lorenzis, A., Fedele, G., Manco, I., Mentaschi, L., Cle- menti, E., Coppini, G., Mercogliano, P., Mendicino, G., Pi- nardi, N., 2024: Climate projections of the Adriatic Sea: role of river release. Frontiers in Climate 6, 1368413. https://doi. org/10.3389/fclim.2024.1368413 Vilibić, I., Pranić, P., Denamiel, C., 2023: North Adriatic Dense Water: lessons learned since the pioneering work of Mira Zore-Armanda 60 years ago, Acta Adriatica 64, 53–78, https://doi.org/10.32582/aa.64.1.11, 2023 Viličić, D., 2014: Ecology and composition of phytoplankton in the Adriatic Sea. Koeltz Scientific Books, Konigstein. Viličić, D., Marasović, I., Mioković, D., 2002: Checklist of phyto- plankton in the eastern Adriatic Sea. Acta Botanica Croatica 61(1), 57–91. Retrieved October 16, 2023 from https://hrcak. srce.hr/file/5822 Voigt, M., 1963: Some new and interesting Mastogloia from the Mediterranean Area and the Far East. Journal of the Royal Microscopical Society 82(2), 111–121. https://doi. org/10.1111/j.1365-2818.1963.tb05308.x Welker, C., Sdrigotti, E., Covelli, S., Faganeli, J., 2002: Microphy- tobenthos in the Gulf of Trieste (Northern Adriatic Sea): Re- lationship with Labile Sedimentary Organic Matter and Nu- trients. Estuarine, Coastal and Shelf Science 55(2), 259–273. https://doi.org/10.1006/ecss.2001.0901 Witkowski, A., Lange-Bertalot, H., Metzelin, D., 2000: Diatom flora of marine coasts I. In: Lange-Bertalot, H. (ed.), Icono- graphia diatomologica, vol. 7, Diversity-taxonomy-identifica- tion, 1–925. A.R.G. Gantner, Ruggell, Germany. Witkowski, A., Li, C., Zgłobicka, I., Yu, S., Ashworth, M., Dabek, P., Qin, S., Tang, C., Krzywda, M., Ruppel, M., Theriot, E. C., Jansen, R. K., Car, A., Płociński, T., Wang, Y., Sabir, J.S.M., Daniszewska-Kowalczyk, G., Kierzek, A., Hajrah, N. H., 2016: Multigene assessment of biodiversity of diatom (Bacillario- phyceae) assemblages from the littoral zone of the Bohai and Yellow Seas in Yantai Region of northeast China with some remarks on ubiquitous taxa. Journal of Coastal Research. 74(sp1), 166–195. https://doi.org/10.2112/SI74-016.1 Zalat, A. A., 2001: New araphid diatom species of the genus Tra- chysphenia Petit from recent coastal sediments of the eastern side of the Gulf of Suez, Egypt. Diatom Research 16(2), 417– 432. https://doi.org/10.1080/0269249X.2001.9705528 Zalat, A. A., 2002: Distribution and origin of diatoms in the bot- tom sediments of the Suez Canal lakes and adjacent areas, Egypt. Diatom Research 17(1), 243–266. https://doi.org/10.1 080/0269249X.2002.9705542 Zalat, A. A., El-Sheekh, M. M., Gaballa, M., 2019: Distribution pattern of diatom flora in the surface sediments of Bardawil Lagoon (North Sinai), Egypt. Thalassas: An International Journal of Marine Sciences volume 35, 531–539. https://doi. org/10.1007/s41208-019-00160-4 Zenetos, A., Cinar, M. E., Pancucci-Papadopoulou, M. A., Harmelin, J. G., Furnari, G., Andaloro, F., Bellou, N., Stref- taris, N., Zibrowius, H, 2005: Annotated list of alien marine species in the Mediterranean with emphasis on worst invasive species. Mediterranean Marine Science 6(2), 63–118. https:// doi.org/10.12681/mms.186 Zidarova, R., Ivanov, P., Hristova, O., Dzhurova, B., Hineva, E., 2022: The unexpected diversity in Amphora sensu lato (Bac- illariophyta) at Sozopol Bay, the western Black Sea. Phytotaxa 544(2), 103–127. https://doi.org/10.11646/phytotaxa.544.2 Zimmermann J., Glöckner G., Jahn R., Enke N., Gemeinholzer B., 2015: Metabarcoding vs. morphological identification to assess diatom diversity in environmental studies. Molecular Ecology Resources 15(3), 526–542. https://onlinelibrary.wi- ley.com/doi/full/10.1111/1755-0998.12336