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.

The dawn of the tropical Atlantic invasion into the Mediterranean Sea.

The Mediterranean Sea is a marine biodiversity hotspot already affected by climate-driven biodiversity collapses. Its highly endemic fauna is at further risk if global warming triggers an invasion of tropical Atlantic species. Here, we combine modern species occurrences with a unique paleorecord from the Last Interglacial (135 to 116 ka), a conservative analog of future climate, to model the future distribution of an exemplary subset of tropical West African mollusks, currently separated from the Mediterranean by cold upwelling off north-west Africa. We show that, already under an intermediate climate scenario (RCP 4.5) by 2050, climatic connectivity along north-west Africa may allow tropical species to colonize a by then largely environmentally suitable Mediterranean. The worst-case scenario RCP 8.5 leads to a fully tropicalized Mediterranean by 2100. The tropical Atlantic invasion will add to the ongoing Indo-Pacific invasion through the Suez Canal, irreversibly transforming the entire Mediterranean into a novel ecosystem unprecedented in human history.

The molluscan assemblage of a pristine Posidonia oceanica meadow in the eastern Mediterranean.

The seagrass Posidonia oceanica forms extensive meadows in the Mediterranean Sea. Studies on their associated highly diverse invertebrate assemblages are limited to the western Mediterranean. The eastern Mediterranean, however, is a basin undergoing rapid change due to the synergistic effects of climate warming, biological invasions and other human stressors that are driving native biodiversity to regional-scale collapses. We here surveyed the shelled molluscan assemblage of a Posidonia oceanica meadow in Plakias, south-western Crete, the first such study in the eastern Mediterranean Sea. This area has increased its yearly mean temperature by 1 °C in the last 20 years and is under heavy pressure by Lessepsian species. We sampled across a 5- to 20-m depth gradient, in two seasons to capture intra-annual variation and the leaf and rhizome strata separately. Against our expectations, the molluscan assemblage proved to be highly diverse, with species richness, dominant species and trophic guilds comparable to healthy western Mediterranean ones, and with a negligible non-indigenous component. The diversity of the native community (following the biotic resistance hypothesis) and oxygen supersaturation in the meadow may cause greater resistance to biological invasions and warming, respectively, suggesting that Posidonia oceanica meadows may act as a precious refugium for native biodiversity in the fast changing eastern Mediterranean Sea. Supplementary Information: The online version contains supplementary material available at 10.1007/s12526-022-01292-2.

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.

Nomenclator, geographic and stratigraphic distribution of the family Triphoridae (Mollusca: Gastropoda).

The microgastropod family Triphoridae is one of the five most diverse marine molluscan families. It likely hosts a few thousand species worldwide, but its taxonomy has long been considered challenging due to the high diversity and subtle morphological characters needed for species delimitation. Consequently, only a small portion of the species appears to be formally described to date. However, further taxonomic work should be based on robust knowledge on the numerous names introduced so far. In this perspective, we have here compiled a list of all published names that can be attributed to the fossil and extant Triphoridae. We list 958 species and 75 genus names, of which 771 are known as extant species and 146 as fossil species, 41 are known from both fossil and extant records. We provide information on type locality and horizon, type material, synonymy and homonymy. Importantly, based on the review of hundreds of publications, we provide a preliminary overview of the geographic and stratigraphic distribution.

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.

Numerous new records of tropical non-indigenous species in the Eastern Mediterranean highlight the challenges of their recognition and identification.

New data on 52 non-indigenous mollusks in the Eastern Mediterranean Sea is reported. Fossarus sp. (aff. aptus sensu Blatterer 2019), Coriophora lessepsiana Albano, Bakker & Sabelli, sp. nov., Cerithiopsis sp. aff. pulvis, Joculator problematicus Albano & Steger, sp. nov., Cerithiopsis sp., Elachisina sp., Iravadia aff. elongata, Vitrinella aff. Vitrinella sp. 1 (sensu Blatterer 2019), Melanella orientalis, Parvioris aff. dilecta, Odostomia cf. dalli, Oscilla virginiae, Parthenina cossmanni, Parthenina typica, Pyrgulina craticulata, Turbonilla funiculata, Cylichna collyra, Musculus coenobitus, Musculus aff. viridulus, Chavania erythraea, Scintilla cf. violescens, Iacra seychellarum and Corbula erythraeensis are new records for the Mediterranean. An unidentified gastropod, Skeneidae indet., Triphora sp., Hypermastus sp., Sticteulima sp., Vitreolina cf. philippi, Odostomia (s.l.) sp. 1, Henrya (?) sp., and Semelidae sp. are further potential new non-indigenous species although their status should be confirmed upon final taxonomic assessment. Additionally, the status of Dikoleps micalii, Hemiliostraca clandestina comb. nov. and H. athenamariae comb. nov. is changed to non-indigenous, range extensions for nine species and the occurrence of living individuals for species previously recorded from empty shells only are reported. Opimaphora blattereri Albano, Bakker & Sabelli, sp. nov. is described from the Red Sea for comparison with the morphologically similar C. lessepsiana Albano, Bakker & Sabelli, sp. nov. The taxonomic part is followed by a discussion on how intensive fieldwork and cooperation among institutions and individuals enabled such a massive report, and how the poor taxonomic knowledge of the Indo-Pacific fauna hampers non-indigenous species detection and identification. Finally, the hypothesis that the simultaneous analysis of quantitative benthic death assemblages can support the assignment of non-indigenous status to taxonomically undetermined species is discussed.

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.

Low diversity or poorly explored? Mesophotic molluscs highlight undersampling in the Eastern Mediterranean.

Mesophotic assemblages are the next frontier of marine exploration in the Mediterranean Sea. Located below recreational scuba diving depths, they are difficult to access but host a diverse array of habitats structured by large invertebrate species. The Eastern Mediterranean has been much less explored than the western part of the basin and its mesophotic habitats are virtually unknown. We here describe two mesophotic (77-92 m depth) molluscan assemblages at a rocky reef and on a soft substrate off northern Israel. We record 172 species, of which 43 (25%) are first records for Israel and increase its overall marine molluscan diversity by 7%. Only five of these species have been reported in recent surveys of the nearby Lebanon, suggesting that our results are robust at a broader scale than our study area and that the reported west-to-east declining diversity gradient in the Mediterranean needs a reappraisal based on proper sampling of the eastern basin. We found only four (2%) non-indigenous species, represented by seven (0.5%) specimens. These results suggest that pristine native assemblages still thrive at this depth in Israel, in contrast to the shallow subtidal heavily affected by global warming and biological invasions, calling for strong conservation actions for these valuable but vulnerable habitats.

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.

Management priorities for marine invasive species.

Managing invasive alien species is particularly challenging in the ocean mainly because marine ecosystems are highly connected across broad spatial scales. Eradication of marine invasive species has only been achieved when species were detected early, and management responded rapidly. Generalized approaches, transferable across marine regions, for prioritizing actions to control invasive populations are currently lacking. Here, expert knowledge was elicited to prioritize 11 management actions for controlling 12 model species, distinguished by differences in dispersion capacity, distribution in the area to be managed, and taxonomic identity. Each action was assessed using five criteria (effectiveness, feasibility, acceptability, impacts on native communities, and cost), which were combined in an 'applicability' metric. Raising public awareness and encouraging the commercial use of invasive species were highly prioritized, whereas biological control actions were considered the least applicable. Our findings can guide rapid decision-making on prioritizing management options for the control of invasive species especially at early stages of invasion, when reducing managers' response time is critical.

The potential of large rafting objects to spread Lessepsian invaders: the case of a detached buoy.

A diverse and abundant fouling community dominated by Lessepsian non-indigenous species was identified on a 13.5-m-long steel buoy stranded on the Israeli coast but originating from Port Said, at the Mediterranean entrance of the Suez Canal, Egypt. The molluscan community was sampled quantitatively by scraping. Three quarters of the individuals and more than half of the species were non-indigenous. Among the latter, a mytilid bivalve, Gregariella cf. ehrenbergi, was first recorded in the Mediterranean Sea on the basis of these samples, suggesting that the full consideration of all potential vectors can contribute to non-indigenous species detection. Large floating objects in coastal waters, such as buoys, are particularly suitable for colonization by Lessepsian species because hard substrates, and artificial ones in particular, are highly susceptible to the establishment of non-indigenous species. Moreover, their size and persistence enable the development of abundant and mature fouling communities that can disseminate propagules as eggs and larvae over long distances and for extended periods if detached. This report highlights the potential for large rafting debris as a vector of the spread of non-indigenous biota within the Mediterranean Sea.

New records of non-indigenous molluscs from the eastern Mediterranean Sea.

We report new findings of non-indigenous Indo-Pacific molluscs from shallow water habitats off Israel, Greece and Egypt, eastern Mediterranean Sea. The bivalves Pillucina vietnamica Zorina, 1978 and Alveinus miliaceus (Issel, 1869) were collected from sandy bottoms off Israel, whereas Gregariella cf. ehrenbergi (Issel, 1869) was recovered from a buoy originating from Port Said, Egypt, and stranded on the Israeli coast. The three species are first records for the Mediterranean Sea. Additionally, we report range extensions for several gastropods: Varicopeza pauxilla (A. Adams, 1855) is recorded from Israel, Phidiana militaris (Alder and Hancock, 1864) from southern Israel (Ashqelon), and Viriola cf. bayani Jousseaume, 1884 from Israel and Crete. Shells and valves of an unidentified lucinid bivalve morphologically distinct from any known Mediterranean species were found along the Israeli Mediterranean shore.

An illustrated catalogue of Rudolf Sturany's type specimens in the Naturhistorisches Museum Wien, Austria (NHMW): deep-sea Eastern Mediterranean molluscs.

The "Pola" expeditions were the first to explore the deep Eastern Mediterranean Sea in the 1890s. They remained the most intense surveys in that area for a century and constitute today a fundamental baseline to assess change in the basin, whose fauna is still inadequately described. Solid taxonomic foundations for the study of deep-sea organisms are needed and we here contribute by revising the name-bearing types of mollusc species introduced by Rudolf Sturany on the basis of the "Pola" material from the Eastern Mediterranean Sea stored in the Natural History Museum in Vienna. Sturany introduced 15 names (Marginella occulta var. minor Sturany, 1896 shall not be considered as the introduction of a new name). He described and established two manuscript names by Monterosato: Jujubinus igneus and Pseudomurex ruderatus. The genus Isorropodon was also introduced together with its type species I. perplexum. For each name, we list the available type material, provide the original description and a translation into English and illustrate the specimens in colour and with SEM imaging.

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.

A risk-based approach to cumulative effect assessments for marine management.

Marine ecosystems are increasingly threatened by the cumulative effects of multiple human pressures. Cumulative effect assessments (CEAs) are needed to inform environmental policy and guide ecosystem-based management. Yet, CEAs are inherently complex and seldom linked to real-world management processes. Therefore we propose entrenching CEAs in a risk management process, comprising the steps of risk identification, risk analysis and risk evaluation. We provide guidance to operationalize a risk-based approach to CEAs by describing for each step guiding principles and desired outcomes, scientific challenges and practical solutions. We reviewed the treatment of uncertainty in CEAs and the contribution of different tools and data sources to the implementation of a risk based approach to CEAs. We show that a risk-based approach to CEAs decreases complexity, allows for the transparent treatment of uncertainty and streamlines the uptake of scientific outcomes into the science-policy interface. Hence, its adoption can help bridging the gap between science and decision-making in ecosystem-based management.

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.