New records of non-indigenous species from the eastern Mediterranean Sea (Crustacea, Mollusca), with a revision of genus Isognomon (Mollusca: Bivalvia).

We report new data on non-indigenous invertebrates from the Mediterranean Sea (four ostracods and 20 molluscs), including five new records for the basin: the ostracods Neomonoceratina iniqua, Neomonoceratina aff. mediterranea, Neomonoceratina cf. entomon, Loxoconcha cf. gisellae (Arthropoda: Crustacea)-the first records of non-indigenous ostracods in the Mediterranean-and the bivalve Striarca aff. symmetrica (Mollusca). Additionally, we report for the first time Electroma vexillum from Israel, and Euthymella colzumensis, Joculator problematicus, Hemiliostraca clandestina, Pyrgulina nana, Pyrgulina microtuber, Turbonilla cangeyrani, Musculus aff. viridulus and Isognomon bicolor from Cyprus. We also report the second record of Fossarus sp. and of Cerithiopsis sp. cf. pulvis in the Mediterranean Sea, the first live collected specimens of Oscilla galilae from Cyprus and the northernmost record of Gari pallida in Israel (and the Mediterranean). Moreover, we report the earliest records of Rugalucina angela, Ervilia scaliola and Alveinus miliaceus in the Mediterranean Sea, backdating their first occurrence in the basin by 3, 5 and 7 years, respectively. We provide new data on the presence of Spondylus nicobaricus and Nudiscintilla aff. glabra in Israel. Finally, yet importantly, we use both morphological and molecular approaches to revise the systematics of the non-indigenous genus Isognomon in the Mediterranean Sea, showing that two species currently co-occur in the basin: the Caribbean I. bicolor, distributed in the central and eastern Mediterranean, and the Indo-Pacific I. aff. legumen, at present reported only from the eastern Mediterranean and whose identity requires a more in-depth taxonomic study. Our work shows the need of taxonomic expertise and investigation, the necessity to avoid the unfounded sense of confidence given by names in closed nomenclature when the NIS belong to taxa that have not enjoyed ample taxonomic work, and the necessity to continue collecting samples-rather than relying on visual censuses and bio-blitzes-to enable accurate detection of non-indigenous species.

Ecological baselines in the Eastern Mediterranean Sea shifted long before the availability of observational time series.

Native biodiversity loss and invasions by nonindigenous species (NIS) have massively altered ecosystems worldwide, but trajectories of taxonomic and functional reorganization remain poorly understood due to the scarcity of long-term data. Where ecological time series are available, their temporal coverage is often shorter than the history of anthropogenic changes, posing the risk of drawing misleading conclusions on systems' current states and future development. Focusing on the Eastern Mediterranean Sea, a region affected by massive biological invasions and the largest climate change-driven collapse of native marine biodiversity ever documented, we followed the taxonomic and functional evolution of an emerging "novel ecosystem", using a unique dataset on shelled mollusks sampled in 2005-2022 on the Israeli shelf. To quantify the alteration of observed assemblages relative to historical times, we also analyzed decades- to centuries-old ecological baselines reconstructed from radiometrically dated death assemblages, time-averaged accumulations of shells on the seafloor that constitute natural archives of past community states. Against expectations, we found no major loss of native biodiversity in the past two decades, suggesting that its collapse had occurred even earlier than 2005. Instead, assemblage taxonomic and functional richness increased, reflecting the diversification of NIS whose trait structure was, and has remained, different from the native one. The comparison with the death assemblage, however, revealed that modern assemblages are taxonomically and functionally much impoverished compared to historical communities. This implies that NIS did not compensate for the functional loss of native taxa, and that even the most complete observational dataset available for the region represents a shifted baseline that does not reflect the actual magnitude of anthropogenic changes. While highlighting the great value of observational time series, our results call for the integration of multiple information sources on past ecosystem states to better understand patterns of biodiversity loss in the Anthropocene.

Three common sampling techniques in Pleistocene coral reefs of the Red Sea: a comparison.

Line Intercept Transects (LIT), Point Intercept Transects (PIT), and Photoquadrats (PQ) are the most common quantitative sampling techniques in modern and fossil coral reefs. Data from coral reefs obtained by the different methods are generally compared between various reef ages and localities. Quaternary reefs from warmer interglacial periods, which represent climate scenarios projected for the future, are particularly interesting for comparisons with modern reefs. Importantly, fossil reefs differ from modern reefs because they are diagenetically altered and time averaged. While several studies have compared different quantitative methods in modern reefs, very few have dealt with the comparability among fossil and between fossil and modern reefs. Here, we compare LIT, PIT at 10, 20 and 50 cm intervals, and PQ in two Pleistocene reef localities in Egypt. We find that alpha diversity, reef cover and community composition are dependent on the method. Results gained with plotless methods (LIT, PIT) differ strongly from results gained with plot methods (PQ). However, coral cover results are similar between LIT and PIT, and community composition is indistinguishable between the two, but alpha diversity depends on the interval used for PIT. We discuss the implications of our findings for comparing coral reefs of various ages and localities. We recommend surveying Pleistocene reefs with PIT at 20 cm intervals. This is because A) alpha diversity is well captured, B) the amount of time averaging recorded by PIT is reduced compared to PQ, C) the PIT results can be directly compared to reefs analyzed by LIT, and D) the method is less time consuming than LIT and PQ.

Palaeontological evidence for community-level decrease in mesopelagic fish size during Pleistocene climate warming in the eastern Mediterranean.

Mesopelagic fishes are an important element of marine food webs, a huge, still mostly untapped food resource and great contributors to the biological carbon pump, whose future under climate change scenarios is unknown. The shrinking of commercial fishes within decades has been an alarming observation, but its causes remain contended. Here, we investigate the effect of warming climate on mesopelagic fish size in the eastern Mediterranean Sea during a glacial-interglacial-glacial transition of the Middle Pleistocene (marine isotope stages 20-18; 814-712 kyr B.P.), which included a 4°C increase in global seawater temperature. Our results based on fossil otoliths show that the median size of lanternfishes, one of the most abundant groups of mesopelagic fishes in fossil and modern assemblages, declined by approximately 35% with climate warming at the community level. However, individual mesopelagic species showed different and often opposing trends in size across the studied time interval, suggesting that climate warming in the interglacial resulted in an ecological shift toward increased relative abundance of smaller sized mesopelagic fishes due to geographical and/or bathymetric distribution range shifts, and the size-dependent effects of warming.

Potential and limitations of applying the mean temperature approach to fossil otolith assemblages.

Evaluation of the impact of climatic changes on the composition of fish assemblages requires quantitative measures that can be compared across space and time. In this respect, the mean temperature of the catch (MTC) approach has been proven to be a very useful tool for monitoring the effect of climate change on fisheries catch. Lack of baseline data and deep-time analogues, however, prevent a more comprehensive evaluation. In this study, we explore the applicability of the mean temperature approach to fossil fish faunas by using otolith assemblage data from the eastern Mediterranean and the northern Adriatic coastal environments corresponding to the last 8000 years (Holocene) and the interval 2.58-1.80 Ma B. P. (Early Pleistocene). The calculated mean temperatures of the otolith assemblage (MTO) range from 13.5 to 17.3 °C. This case study shows that the MTO can successfully capture compositional shifts in marine fish faunas based on variations in their climatic affinity driven by regional climate differences. However, the index is sensitive to methodological choices and thus requires standardized sampling. Even though theoretical and methodological issues prevent direct comparisons between MTO and MTC values, the MTO offers a useful quantitative proxy for reconstructing spatial and temporal trends in the biogeographic affinity of fossil otolith assemblages.

Bad neighbors? Niche overlap and asymmetric competition between native and Lessepsian limpets in the Eastern Mediterranean rocky intertidal.

The Eastern Mediterranean Sea hosts more non-indigenous species than any other marine region, yet their impacts on the native biota remain poorly understood. Focusing on mollusks from the Israeli rocky intertidal, we explored the hypothesis that this abiotically harsh habitat supports a limited trait diversity, and thus may promote niche overlap and competition between native and non-indigenous species. Indeed, native and non-indigenous assemblage components often had a highly similar trait composition, caused by functionally similar native (Patella caerulea) and non-indigenous (Cellana rota) limpets. Body size of P. caerulea decreased with increasing C. rota prevalence, but not vice versa, indicating potential asymmetric competition. Although both species have coexisted in Israel for >15 years, a rapid 'replacement' of native limpets by C. rota has been reported for a thermally polluted site, suggesting that competition and regionally rapid climate-related seawater warming might interact to progressively erode native limpet performance along the Israeli coast.

Native biodiversity collapse in the eastern Mediterranean.

Global warming causes the poleward shift of the trailing edges of marine ectotherm species distributions. In the semi-enclosed Mediterranean Sea, continental masses and oceanographic barriers do not allow natural connectivity with thermophilic species pools: as trailing edges retreat, a net diversity loss occurs. We quantify this loss on the Israeli shelf, among the warmest areas in the Mediterranean, by comparing current native molluscan richness with the historical one obtained from surficial death assemblages. We recorded only 12% and 5% of historically present native species on shallow subtidal soft and hard substrates, respectively. This is the largest climate-driven regional-scale diversity loss in the oceans documented to date. By contrast, assemblages in the intertidal, more tolerant to climatic extremes, and in the cooler mesophotic zone show approximately 50% of the historical native richness. Importantly, approximately 60% of the recorded shallow subtidal native species do not reach reproductive size, making the shallow shelf a demographic sink. We predict that, as climate warms, this native biodiversity collapse will intensify and expand geographically, counteracted only by Indo-Pacific species entering from the Suez Canal. These assemblages, shaped by climate warming and biological invasions, give rise to a 'novel ecosystem' whose restoration to historical baselines is not achievable.

Ecological regime shift preserved in the Anthropocene stratigraphic record.

Palaeoecological data are unique historical archives that extend back far beyond the last several decades of ecological observations. However, the fossil record of continental shelves has been perceived as too coarse (with centennial-millennial resolution) and incomplete to detect processes occurring at yearly or decadal scales relevant to ecology and conservation. Here, we show that the youngest (Anthropocene) fossil record on the northern Adriatic continental shelf provides decadal-scale resolution that accurately documents an abrupt ecological change affecting benthic communities during the twentieth century. The magnitude and the duration of the twentieth century shift in body size of the bivalve Corbula gibba is unprecedented given that regional populations of this species were dominated by small-size classes throughout the Holocene. The shift coincided with compositional changes in benthic assemblages, driven by an increase from approximately 25% to approximately 70% in median per-assemblage abundance of C. gibba. This regime shift increase occurred preferentially at sites that experienced at least one hypoxic event per decade in the twentieth century. Larger size and higher abundance of C. gibba probably reflect ecological release as it coincides with an increase in the frequency of seasonal hypoxia that triggered mass mortality of competitors and predators. Higher frequency of hypoxic events is coupled with a decline in the depth of intense sediment mixing by burrowing benthic organisms from several decimetres to less than 20 cm, significantly improving the stratigraphic resolution of the Anthropocene fossil record and making it possible to detect sub-centennial ecological changes on continental shelves.

Drivers of beta diversity in modern and ancient reef-associated soft-bottom environments.

Beta diversity, the compositional variation among communities, is often associated with environmental gradients. Other drivers of beta diversity include stochastic processes, priority effects, predation, or competitive exclusion. Temporal turnover may also explain differences in faunal composition between fossil assemblages. To assess the drivers of beta diversity in reef-associated soft-bottom environments, we investigate community patterns in a Middle to Late Triassic reef basin assemblage from the Cassian Formation in the Dolomites, Northern Italy, and compare results with a Recent reef basin assemblage from the Northern Bay of Safaga, Red Sea, Egypt. We evaluate beta diversity with regard to age, water depth, and spatial distance, and compare the results with a null model to evaluate the stochasticity of these differences. Using pairwise proportional dissimilarity, we find very high beta diversity for the Cassian Formation (0.91 ± 0.02) and slightly lower beta diversity for the Bay of Safaga (0.89 ± 0.04). Null models show that stochasticity only plays a minor role in determining faunal differences. Spatial distance is also irrelevant. Contrary to expectations, there is no tendency of beta diversity to decrease with water depth. Although water depth has frequently been found to be a key factor in determining beta diversity, we find that it is not the major driver in these reef-associated soft-bottom environments. We postulate that priority effects and the biotic structuring of the sediment may be key determinants of beta diversity.

A decline in molluscan carbonate production driven by the loss of vegetated habitats encoded in the Holocene sedimentary record of the Gulf of Trieste.

Carbonate sediments in non-vegetated habitats on the north-east Adriatic shelf are dominated by shells of molluscs. However, the rate of carbonate molluscan production prior to the 20th century eutrophication and overfishing on this and other shelves remains unknown because: (i) monitoring of ecosystems prior to the 20th century was scarce; and (ii) ecosystem history inferred from cores is masked by condensation and mixing. Here, based on geochronological dating of four bivalve species, carbonate production during the Holocene is assessed in the Gulf of Trieste, where algal and seagrass habitats underwent a major decline during the 20th century. Assemblages of sand-dwelling Gouldia minima and opportunistic Corbula gibba are time-averaged to >1000 years and Corbula gibba shells are older by >2000 years than shells of co-occurring Gouldia minima. This age difference is driven by temporally disjunct production of two species coupled with decimetre-scale mixing. Stratigraphic unmixing shows that Corbula gibba declined in abundance during the highstand phase and increased again during the 20th century. In contrast, one of the major contributors to carbonate sands - Gouldia minima - increased in abundance during the highstand phase, but declined to almost zero abundance over the past two centuries. Gouldia minima and herbivorous gastropods associated with macroalgae or seagrasses are abundant in the top-core increments but are rarely alive. Although Gouldia minima is not limited to vegetated habitats, it is abundant in such habitats elsewhere in the Mediterranean Sea. This live-dead mismatch reflects the difference between highstand baseline communities (with soft-bottom vegetated zones and hard-bottom Arca beds) and present-day oligophotic communities with organic-loving species. Therefore, the decline in light penetration and the loss of vegetated habitats with high molluscan production traces back to the 19th century. More than 50% of the shells on the sea floor in the Gulf of Trieste reflect inactive production that was sourced by heterozoan carbonate factory in algal or seagrass habitats.

Molluscan benthic communities at Brijuni Islands (northern Adriatic Sea) shaped by Holocene sea-level rise and recent human eutrophication and pollution.

The effects of and the interplay between natural and anthropogenic influences on the composition of benthic communities over long time spans are poorly understood. Based on a 160-cm-long sediment core collected at 44 m water depth in the NE Adriatic Sea (Brijuni Islands, Croatia), we document changes in molluscan communities since the Holocene transgression ~11,000 years ago and assess how they were shaped by environmental changes. We find that (1) a transgressive lag deposit with a mixture of terrestrial and marine species contains abundant seagrass-associated gastropods and epifaunal suspension-feeding bivalves, (2) the maximum-flooding phase captures the establishment of epifaunal bivalve-dominated biostromes in the photic zone, and (3) the highstand phase is characterized by increasing infaunal suspension feeders and declining seagrass-dwellers in bryozoan-molluscan muddy sands. Changes in the community composition between the transgressive and the highstand phase can be explained by rising sea level, reduced light penetration, and increase in turbidity, as documented by the gradual up-core shift from coarse molluscan skeletal gravel with seagrass-associated molluscs to bryozoan sandy muds. In the uppermost 20 cm (median age <200 years), however, epifaunal and grazing species decline and deposit-feeding and chemosymbiotic species increase in abundance. These changes concur with rising concentrations of nitrogen and organic pollutants due to the impact of eutrophication, pollution, and trawling in the 20th century. The late highstand benthic assemblages with abundant bryozoans, high molluscan diversity, and abundance of soft-bottom epi- and infaunal filter feeders and herbivores represent the circalittoral baseline community largely unaffected by anthropogenic impacts.

20th century increase in body size of a hypoxia-tolerant bivalve documented by sediment cores from the northern Adriatic Sea (Gulf of Trieste).

An increase in the frequency of hypoxia, mucilages, and sediment pollution occurred in the 20th century in the Adriatic Sea. To assess the effects of these impacts on bivalves, we evaluate temporal changes in size structure of the opportunistic bivalve Corbula gibba in four sediment cores that cover the past ~500 years in the northern, eutrophic part and ~10,000 years in the southern, mesotrophic part of the Gulf of Trieste. Assemblages exhibit a stable size structure during the highstand phase but shift to bimodal distributions and show a significant increase in the 95th percentile size during the 20th century. This increase in size by 2-3 mm is larger than the northward size increase associated with the transition from mesotrophic to eutrophic habitats. It coincides with increasing concentrations of total organic carbon and nitrogen, and can be related to enhanced food supply and by the tolerance of C. gibba to hypoxia.

Historical ecology of a biological invasion: the interplay of eutrophication and pollution determines time lags in establishment and detection.

Human disturbance modifies selection regimes, depressing native species fitness and enabling the establishment of non-indigenous species with suitable traits. A major impediment to test the effect of disturbance on invasion success is the lack of long-term data on the history of invasions. Here, we overcome this problem and reconstruct the effect of disturbance on the invasion of the bivalve Anadara transversa from sediment cores in the Adriatic Sea. We show that (1) the onset of major eutrophication in the 1970s shifted communities towards species tolerating hypoxia, and (2) A. transversa was introduced in the 1970s but failed to reach reproductive size until the late 1990s because of metal contamination, resulting in an establishment and detection lag of ~25 years. Subfossil assemblages enabled us to (1) disentangle the distinct stages of invasion, (2) quantify time-lags and (3) finely reconstruct the interaction between environmental factors and the invasion process, showing that while disturbance does promote invasions, a synergism of multiple disturbances can shift selection regimes beyond tolerance limits and induce significant time lags in establishment. The quantification of these time lags enabled us to reject the hypothesis that aquaculture was an initial vector of introduction, making shipping the most probable source.

Reliable estimates of beta diversity with incomplete sampling.

Beta diversity, the compositional variation among communities or assemblages, is crucial to understanding the principles of diversity assembly. The mean pairwise proportional dissimilarity expresses overall heterogeneity of samples in a data set and is among the most widely used and most robust measures of beta diversity. Obtaining a complete list of taxa and their abundances requires substantial taxonomic expertise and is time consuming. In addition, the information is generally incomplete due to sampling biases. Based on the concept of the ecological significance of dominant taxa, we explore whether determining proportional dissimilarity can be simplified based on dominant species. Using simulations and six case studies, we assess the correlation between complete community compositional data and reduced subsets of a varying number of dominant species. We find that gross beta diversity is usually depicted accurately when only the 80th percentile or five of the most abundant species of each site is considered. In data sets with very high evenness, at least the 10 most abundant species should be included. Focusing on dominant species also maintains the rank-order of beta diversity among sites. Our new approach will allow ecologists and paleobiologists to produce a far greater amount of data on diversity patterns with less time and effort, supporting conservation studies and basic science.

Holocene ecosystem shifts and human-induced loss of Arca and Ostrea shell beds in the north-eastern Adriatic Sea.

The molluscan assemblages in a sediment core from the north-eastern Adriatic show significant compositional changes over the past 10,000yrs related to (1) natural deepening driven by the post-glacial sea-level rise, (2) increasing abundance of skeletal sand and gravel, and (3) anthropogenic impacts. The transgressive phase (10,000-6000 BP) is characterized by strongly time-averaged communities dominated by infaunal bivalves. During the early highstand (6000-4000 BP), the abundance of epifaunal filter feeders and grazers increases, and gastropods become more important. Epifaunal dominance culminates during the late highstand (4000-2000 BP) with the development of extensive shell beds formed by large-sized Arca noae and Ostrea sp. bivalves. This community persists until the early 20th century, when it falls victim to multiple anthropogenic impacts, mainly bottom trawling, and is substituted by an infauna-dominated community indicative of instability, disturbance and organic enrichment. The re-establishment of this unique shell-bed ecosystem can be a goal for restoration efforts.

Climate change and body size shift in Mediterranean bivalve assemblages: unexpected role of biological invasions.

Body size is a synthetic functional trait determining many key ecosystem properties. Reduction in average body size has been suggested as one of the universal responses to global warming in aquatic ecosystems. Climate change, however, coincides with human-enhanced dispersal of alien species and can facilitate their establishment. We address effects of species introductions on the size structure of recipient communities using data on Red Sea bivalves entering the Mediterranean Sea through the Suez Canal. We show that the invasion leads to increase in median body size of the Mediterranean assemblage. Alien species are significantly larger than native Mediterranean bivalves, even though they represent a random subset of the Red Sea species with respect to body size. The observed patterns result primarily from the differences in the taxonomic composition and body-size distributions of the source and recipient species pools. In contrast to the expectations based on the general temperature-size relationships in marine ectotherms, continued warming of the Mediterranean Sea indirectly leads to an increase in the proportion of large-bodied species in bivalve assemblages by accelerating the entry and spread of tropical aliens. These results underscore complex interactions between changing climate and species invasions in driving functional shifts in marine ecosystems.

Responses of molluscan communities to centuries of human impact in the northern Adriatic Sea.

In sediment cores spanning ~500 years of history in the Gulf of Trieste, down-core changes in molluscan community structure are characterized by marked shifts in species and functional composition. Between the 16th and 19th century, a strong heavy metal contamination of the sediments, most notably by Hg, together with the effects of natural climatic oscillations (increased sedimentation and organic enrichment) drive community changes. Since the early 20th century up to 2013, the combined impacts of cultural eutrophication, frequent hypoxic events and intensifying bottom trawling replace heavy metal contamination and climatic factors as the main drivers. The pollution-tolerant and opportunistic bivalve Corbula gibba and the scavenging gastropod Nassarius pygmaeus significantly increase in abundance during the 20th century, while species more sensitive to disturbances and hypoxia such as Turritella communis and Kurtiella bidentata become rare or absent. An infaunal life habit and scavenging emerge as the dominant life strategies during the late 20th century. Down-core shifts in the proportional abundances of molluscan species and functional groups represent a sensitive proxy for past ecological changes and reveal a century-long anthropogenic impact as the main driver behind these processes in the northern Adriatic Sea, offering also a unique perspective for other shallow marine ecosystems worldwide.

Lenticellaria and Hillerella, new kraussinoid genera (Kraussinoidea, Brachiopoda) from Indo-Pacific and Red Sea waters: evolution in the subfamily Megerliinae.

Two new kraussinid brachiopod genera, namely Lenticellaria gen. nov. and Hillerella gen. nov. are described from Pacific waters in the sub-equatorial zone in the Indonesian Archipelago, from Indian Ocean waters in Madagascar and from Red Sea waters in Egypt (Gulf of Aqaba) and Sudan. This fills the equatorial gap in the distribution of the superfamily Kraussinoidea, known from higher latitudes in both hemispheres. The micromorphic new material described is an excellent example of homeomorphy in brachiopods. It also provides new information on the distribution of the genus Megerlia sensu stricto and illustrates subtle variations in the evolutionary process of the reduced brachidium in Kraussinoidea.

An innovative piston corer for large-volume sediment samples.

Coring is one of several standard procedures to extract sediments and their faunas from open marine, estuarine, and limnic environments. Achieving sufficiently deep penetration, obtaining large sediment volumes in single deployments, and avoiding sediment loss upon retrieval remain problematic. We developed a piston corer with a diameter of 16 cm that enables penetration down to 1.5 m in a broad range of soft bottom types, yields sufficient material for multiple analyses, and prevents sediment loss due to a specially designed hydraulic core catcher. A novel extrusion system enables very precise slicing and preserves the original sediment stratification by keeping the liners upright. The corer has moderate purchase costs and a robust and simple design that allows for a deployment from relatively small vessels as available at most marine science institutions. It can easily be operated by two to three researchers rather than by specially trained technicians. In the northern Adriatic Sea, the corer successfully extracted more than 50 cores from a range of fine mud to coarse sand, at water depths from three to 45 m. The initial evaluation of the cores demonstrated their usefulness for fauna sequences along with heavy metal, nutrient and pollutant analyses. Their length is particularly suited for historical ecological work requiring sedimentary and faunal sequences to reconstruct benthic communities over the last millennia.