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.

Bioconcentration and lethal effects of gas-condensate and crude oil on nearshore copepod assemblages.

The progressive establishment of gas platforms and increasing petroleum accidents pose a threat to zooplankton communities and thus to pelagic ecosystems. This study is the first to compare the impacts of gas-condensate and crude oil on copepod assemblages. We conducted microcosm experiments simulating slick scenarios at five different concentrations of gas-condensate and crude oil to determine and compare their lethal effects and the bioconcentration of low molecular weight polycyclic aromatic hydrocarbons (LMW-PAHs) in eastern Mediterranean coastal copepod assemblages. We found that gas-condensate had a two-times higher toxic effect than crude oil, significantly reducing copepod survival with increased exposure levels. The LMW-PAHs bioconcentration factor was 1-2 orders of magnitude higher in copepods exposed to gas-condensate than in those exposed to crude oil. The median lethal concentration (LC50) was significantly lower in calanoids vs. cyclopoid copepods, suggesting that calanoids are more susceptible to gas-condensate and crude oil pollution, with potential trophic implications.

Exceptionally high levels of total mercury in deep-sea sharks of the Southeastern Mediterranean sea over the last ∼ 40 years.

Deep-sea habitats are currently recognized as a hot spot for mercury (Hg) accumulation from anthropogenic sources, resulting in elevated concentrations of total mercury (THg) in deep-sea megafauna. Among them, deep-sea sharks (Class Chondrichthyes) are characterized by high trophic position and extended longevity and are, therefore, at high risk for mercury contamination. Despite this, sharks are overexploited by fishing activity in increasingly deeper water, worldwide, imposing health risks to human consumption. While it is imperative to better understand long-term mercury contamination in deep-sea megafauna, few historical data sets exist to capture this process. Here we explore four decades (1985-2022) of THg accumulation in five species of deep-sea sharks (G. melastomus, E. spinax, S. rostratus, C. granulosus, and D. licha) of the ultra-oligotrophic Southeastern Mediterranean Sea (SEMS) sampled during 19 research cruises. We exhibited exceptionally high THg levels (per length/weight), the highest as 16.6 µg g-1 (wet wt.), almost entirely (98.9 %; n = 298 specimens) exceeding the limit for safe consumption (0.3-0.5 µg THg g-1 wet wt.). The maximal THg levels of the long-lived species D. licha and C. granulosus in the SEMS were enriched by a factor of âˆ¼ 7 and >10 compared to counterpart species from other oceanic areas, respectively. We attribute this to the ultra-oligotrophic conditions of the SEMS, which cause slower growth rates and dwarfism in deep-sea sharks, resulting in an extended exposure time to mercury contamination. In the long-lived species, C. granulosus and D. licha, a temporal increase of average THg levels of âˆ¼ 80 % was recorded between 1987-1999 and 2021-2022. This likely reflects the long-term accumulation of historical anthropogenic Hg in deep-sea environments, which is further amplified in marginal seas such as the Mediterranean, impacted by global air pollution crossroads and surrounded by land-based pollution sources. Future consumption of products from deep-sea sharks is potentially high risk to human health.

Accumulation of total mercury in deep-sea sediments and biota across a bathymetric gradient in the Southeastern Mediterranean Sea.

This study explores the accumulation of total mercury (THg) in deep-sea sediments and demersal megafauna of the ultra-oligotrophic Southeastern Mediterranean Sea (SEMS) across bathymetric gradients in the range 35-1900 m, sampled in seven cruises during 2013, 2017-2021, and 2023. Measurements of THg were conducted in surficial (0.0-0.5 cm) and subsurface (9.0-10 cm) sediments, demersal sharks, demersal teleost fish, and benthic crustaceans. Sedimentary organic carbon and biota δ13C and δ15N values were determined to explore possible foraging habitats and dietary sources of THg. The results exhibit an increasing trend of THg in surficial sediments with increasing bottom depth, while in the subsurface, pre-industrial sediments, THg remains lower, slightly increasing with depth. Having no major terrestrial point sources in this area, this increasing trend of THg in surficial sediments across bathymetric gradients is controlled by atmospheric mercury deposition, scavenged by the biological pump, and by lateral transport of particulate Hg in winnowed fine particles from the shelf. Similarly, the THg in benthic crustaceans and demersal fish ranged between 0.02 and 2.71 µg g-1 wet weight (0.06 and 10.8 µg g-1 dry weight) and increased with muscle δ13C as a function of distance offshore, while presenting a low THg-δ15N bio-magnification power. Our results suggest that foraging habitats, longevity, and species-specific depth distribution control their muscle THg bioaccumulation. Despite this complexity, the pooling of THg in megafauna into specific deep zones reflected the trend of increasing anthropogenic THg across bathymetric gradients. Furthermore, many of the biota measurements exceeded safe consumption thresholds for Hg and therefore, should be considered carefully in the development and regulation of deep-sea trawling in this region.

A broad-taxa approach as an important concept in ecotoxicological studies and pollution monitoring.

Aquatic invertebrates play a pivotal role in (eco)toxicological assessments because they offer ethical, cost-effective and repeatable testing options. Additionally, their significance in the food chain and their ability to represent diverse aquatic ecosystems make them valuable subjects for (eco)toxicological studies. To ensure consistency and comparability across studies, international (eco)toxicology guidelines have been used to establish standardised methods and protocols for data collection, analysis and interpretation. However, the current standardised protocols primarily focus on a limited number of aquatic invertebrate species, mainly from Arthropoda, Mollusca and Annelida. These protocols are suitable for basic toxicity screening, effectively assessing the immediate and severe effects of toxic substances on organisms. For more comprehensive and ecologically relevant assessments, particularly those addressing long-term effects and ecosystem-wide impacts, we recommended the use of a broader diversity of species, since the present choice of taxa exacerbates the limited scope of basic ecotoxicological studies. This review provides a comprehensive overview of (eco)toxicological studies, focusing on major aquatic invertebrate taxa and how they are used to assess the impact of chemicals in diverse aquatic environments. The present work supports the use of a broad-taxa approach in basic environmental assessments, as it better represents the natural populations inhabiting various ecosystems. Advances in omics and other biochemical and computational techniques make the broad-taxa approach more feasible, enabling mechanistic studies on non-model organisms. By combining these approaches with in vitro techniques together with the broad-taxa approach, researchers can gain insights into less-explored impacts of pollution, such as changes in population diversity, the development of tolerance and transgenerational inheritance of pollution responses, the impact on organism phenotypic plasticity, biological invasion outcomes, social behaviour changes, metabolome changes, regeneration phenomena, disease susceptibility and tissue pathologies. This review also emphasises the need for harmonised data-reporting standards and minimum annotation checklists to ensure that research results are findable, accessible, interoperable and reusable (FAIR), maximising the use and reusability of data. The ultimate goal is to encourage integrated and holistic problem-focused collaboration between diverse scientific disciplines, international standardisation organisations and decision-making bodies, with a focus on transdisciplinary knowledge co-production for the One-Health approach.

Shedding light on the Ophel biome: the trans-Tethyan phylogeography of the sulfide shrimp Tethysbaena (Peracarida: Thermosbaenacea) in the Levant.

Background: Tethysbaena are small peracarid crustaceans inhabiting extreme environments such as subterranean lakes and thermal springs, represented by endemic species found around the ancient Tethys, including the Mediterranean, Arabian Sea, Mid-East Atlantic, and the Caribbean Sea. Two Tethysbaena species are known from the Levant: T. relicta, found along the Dead Sea-Jordan Rift Valley, and T. ophelicola, found in the Ayyalon cave complex in the Israeli coastal plain, both belonging to the same species-group based on morphological cladistics. Along the biospeleological research of the Levantine subterranean fauna, three biogeographic hypotheses determining their origins were proposed: (1) Pliocenic transgression, (2) Mid-late Miocenic transgression, and (3) The Ophel Paradigm, according to which these are inhabitants of a chemosynthetic biome as old as the Cambrian. Methods: Tethysbaena specimens of the two Levantine species were collected from subterranean groundwaters. We used the mitochondrial cytochrome c oxidase subunit I (COI) gene and the nuclear ribosomal 28S (28S rRNA) gene to establish the phylogeny of the Levantine Tethysbaena species, and applied a molecular clock approach for inferring their divergence times. Results: Contrary to the morphological cladistic-based classification, we found that T. relicta shares an ancestor with Tethysbaena species from Oman and the Dominican Republic, whereas the circum-Mediterranean species (including T. ophelicola) share another ancestor. The mean age of the node linking T. relicta from the Dead Sea-Jordan Rift Valley and Tethysbaena from Oman was 20.13 MYA. The mean estimate for the divergence of T. ophelicola from the Mediterranean Tethysbaena clade dated to 9.46 MYA. Conclusions: Our results indicate a two-stage colonization of Tethysbaena in the Levant: a late Oligocene transgression, through a marine gulf extending from the Arabian Sea, leading to the colonization of T. relicta in the Dead Sea-Jordan Rift Valley, whereas T. ophelicola, originating from the Mesogean ancestor, inhabited anchialine caves in the coastal plain of Israel during the Mid-Miocene.

Tar patties are hotspots of hydrocarbon turnover and nitrogen fixation during a nearshore pollution event in the oligotrophic southeastern Mediterranean Sea.

Weathered oil, that is, tar, forms hotspots of hydrocarbon degradation by complex biota in marine environment. Here, we used marker gene sequencing and metagenomics to characterize the communities of bacteria, archaea and eukaryotes that colonized tar patties and control samples (wood, plastic), collected in the littoral following an offshore spill in the warm, oligotrophic southeastern Mediterranean Sea (SEMS). We show potential aerobic and anaerobic hydrocarbon catabolism niches on tar interior and exterior, linking carbon, sulfur and nitrogen cycles. Alongside aromatics and larger alkanes, short-chain alkanes appear to fuel dominant populations, both the aerobic clade UBA5335 (Macondimonas), anaerobic Syntropharchaeales, and facultative Mycobacteriales. Most key organisms, including the hydrocarbon degraders and cyanobacteria, have the potential to fix dinitrogen, potentially alleviating the nitrogen limitation of hydrocarbon degradation in the SEMS. We highlight the complexity of these tar-associated communities, where bacteria, archaea and eukaryotes co-exist, likely exchanging metabolites and competing for resources and space.

Superior growth traits of invaded (Caribbean) versus native (Red sea) populations of the seagrass Halophila stipulacea.

The seagrass Halophila stipulacea is native to the Red Sea. It invaded the Mediterranean over the past century and most of the Caribbean over the last two decades. Understanding the main drivers behind the successful invasiveness of H. stipulacea has become crucial. We performed a comprehensive study including field measurements, a mesocosm experiment, and a literature review to identify 'superior growth traits' that can potentially explain the success story of H. stipulacea. We assessed meadow characteristics and plant traits of three invasive H. stipulacea populations growing off the Island of Sint Eustatius (eastern Caribbean). We compared similar parameters between native (Eilat, northern Red Sea) and invasive (Caribbean) H. stipulacea plants in a common-garden mesocosm. Lastly, we compared our field measurements with published data. The newly arrived H. stipulacea plants from St. Eustatius were characterized by higher percent cover, higher below- and above-ground biomasses, more apical shoots, and faster leaf turnover rates than those measured in both native and older invaded habitats. These results were further confirmed by the mesocosm experiment where the invasive H. stipulacea plants grew faster and developed more apical shoots than the native plants. Results suggest that increased growth vigour is one of the main invasive traits that characterize successful invasive H. stipulacea populations in the Caribbean and potentially in other invaded areas. We encourage long-term monitoring of H. stipulacea in both native and invaded habitats to better understand the future spread of this species and its impacts on communities and their ecosystem functions and services. Supplementary Information: The online version contains supplementary material available at 10.1007/s10530-023-03045-z.

Parasitism enhances gastropod feeding on invasive and native algae while altering essential energy reserves for organismal homeostasis upon warming.

Marine bioinvasions are of increasing attention due to their potential of causing ecological and economic loss. The seaweed Gracilaria vermiculophylla has recently invaded the Baltic Sea, where, under certain conditions, it was found to outcompete the native alga Fucus vesiculosus. Parasites of grazers and temperature are among the potential factors which might indirectly modulate the interactions between these co-occurring algae through their single and combined effects on grazing rates. We tested the temperature and parasitism effects on the feeding of the gastropod Littorina littorea on F. vesiculosus vs. G. vermiculophylla. Uninfected and trematode-infected gastropods were exposed to 10, 16, 22, and 28 °C for 4 days while fed with either algae. Faeces production was determined as a proxy for grazing rate, and HSP70 expression, glycogen and lipid concentrations were used to assess the gastropod's biochemical condition. Gracilaria vermiculophylla was grazed more than F. vesiculosus. Trematode infection significantly enhanced faeces production, decreased glycogen concentrations, and increased lipid concentrations in the gastropod. Warming significantly affected glycogen and lipid concentrations, with glycogen peaking at 16 °C and lipids at 22 °C. Although not significant, warming and trematode infection increased HSP70 levels. Increased faeces production in infected snails and higher faeces production by L. littorea fed with G. vermiculophylla compared to those which fed on F. vesiculosus, suggest parasitism as an important indirect modulator of the interaction between these algae. The changes in the gastropod's biochemical condition indicate that thermal stress induced the mobilization of energy reserves, suggesting a possible onset of compensatory metabolism. Finally, glycogen decrease in infected snails compared to uninfected ones might make them more susceptible to thermal stress.

Machine learning techniques to characterize functional traits of plankton from image data.

Plankton imaging systems supported by automated classification and analysis have improved ecologists' ability to observe aquatic ecosystems. Today, we are on the cusp of reliably tracking plankton populations with a suite of lab-based and in situ tools, collecting imaging data at unprecedentedly fine spatial and temporal scales. But these data have potential well beyond examining the abundances of different taxa; the individual images themselves contain a wealth of information on functional traits. Here, we outline traits that could be measured from image data, suggest machine learning and computer vision approaches to extract functional trait information from the images, and discuss promising avenues for novel studies. The approaches we discuss are data agnostic and are broadly applicable to imagery of other aquatic or terrestrial organisms.

Jellyfish swarm impair the pretreatment efficiency and membrane performance of seawater reverse osmosis desalination.

Circumstantial evidence has suggested that jellyfish swarms impair the operation of seawater reverse osmosis desalination facilities. However, only limited information is currently available on the pretreatment efficiency of jellyfish and their effects on reverse osmosis (RO) membrane performance. Here, we have comprehensively tested the pretreatment efficiency of a dual-media gravity filter and cartridge micro-filtration following the addition of jellyfish into the feedwater. Concurrently, the fouling propensity and performance of the RO membranes were examined. We show that jellyfish demise resulted in seawater eutrophication that triggered a significant increase in bacterial biomass (∼50-fold), activity (∼7-fold), and release of transparent exopolymer particles (∼5-fold), peaking three days after the addition of jellyfish into the feedwater. In parallel, a significant reduction in permeate water flux was recorded (∼10%) while trans-membrane pressure sharply increased (15%), reaching the operation pressure limit of our system (75 bar) after five days. At the conclusion of the experiments, the membrane surface was heavily covered by large chunks of organic-rich material and multilayered biofilms. Our results provide a holistic view on the operational challenges of seawater reverse osmosis (SWRO) desalination triggered by jellyfish swarms in coastal areas. Following the above, it can be inferred that freshwater production will likely be halted three days after drawing the jellyfish into the pretreatment system. Outcomes from these results may lead to the development of science-based operational protocols to cope with growing occurrence of jellyfish swarms around the intake of SWRO desalination facilities worldwide.

Sea level rise can severely reduce biodiversity and community net production on rocky shores.

Sea level rise (SLR), driven by anthropogenic climate change, can be a major threat to coastal ecosystems. Among the most biologically diverse but SLR-threatened coastal ecosystems are rocky shores, especially in regions with a small tidal range. Nonetheless, the impacts of SLR on rocky shore biodiversity, community structure and ecosystem functions have rarely been studied. Here, we use the biogenic intertidal ecosystem, Mediterranean vermetid reefs on the Israeli coast, as case study for testing the potential impact of SLR on reef communities, with surveys, 3D topographic mapping plus SLR simulations, and a manipulative community translocation experiment. We show that: (1) biodiversity is much lower on very shallow, permanently submerged, horizontal rocky surfaces compared to that on intertidal reef platforms, (2) the extensive intertidal platforms will permanently drown under even modest SLR scenarios, (3) the rich intertidal community will transform, when permanently submerged, either to a very different but still rich community when protected from grazing by highly abundant invasive fish (rabbitfish), or to a much poorer turf community when exposed to such fish grazing, and (4) the reef community net production will drastically drop under permanent submersion. Because the main ecosystem engineer of the vermetid reefs, Dendropoma anguliferum (Monterosato, 1878), is nearly extinct in the southeast Levant, it is unlikely that new reefs will be formed higher on the shore in the future, presumably resulting in extensive coastal ecological shifts. Considerable coastal community shifts are forecasted for many regions globally due to SLR, as many shorelines are predicted to suffer from "coastal squeeze". Hence, similar manipulative experiments are encouraged in other regions to test for generality vs. context dependency in SLR ecological impacts. We suggest that in cases where essential/unique intertidal habitats like vermetid reefs are expected to vanish by SLR, constructing carefully-planned, ecologically friendly, artificial alternatives should be considered.

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.

Community dynamics and ecological shifts on Mediterranean vermetid reefs.

Mediterranean coastal ecosystems experience many local and global stressors and require long-term monitoring to detect and follow trends in community structure. Between 2009 and 2017, we seasonally and annually monitored the spatiotemporal community dynamics at 11 sites on the rocky shores of the southeastern Mediterranean, focusing on the understudied intertidal vermetid reef ecosystem. Marked seasonal trends were found in biodiversity, with the highest diversity in winter and spring. Canopy-forming brown algae, dominating the northwestern Mediterranean intertidal reefs, were generally scarce on the reef platform and almost only found in tidepools. Interannual shifts in community structure were driven mostly by sharp fluctuations in a few dominant native and alien species and the regional mass mortality of an Indo-Pacific mussel in summer 2016. Compared to an older macroalgae dataset, dating back to 1973-1995, we found that some warm-affinity (summer) taxa became more dominant and cold-affinity (winter) species less dominant, while one once conspicuous species, Halimeda tuna, completely disappeared. The observed community shifts are probably driven mostly by stressors related to climate change. We encourage forming a network of long-term, multi-site ecological monitoring programs in the Mediterranean to improve our understanding of ecosystem change and to enable making better predictions at the basin scale.

Epiphytes provide micro-scale refuge from ocean acidification.

Coralline algae, a major calcifying component of coastal shallow water communities, have been shown to be one of the more vulnerable taxonomic groups to ocean acidification (OA). Under OA, the interaction between corallines and epiphytes was previously described as both positive and negative. We hypothesized that the photosynthetic activity and the complex structure of non-calcifying epiphytic algae that grow on corallines ameliorate the chemical microenvironmental conditions around them, providing protection from OA. Using mesocosm and microsensor experiments, we showed that the widespread coralline Ellisolandia elongata is less susceptible to the detrimental effects of OA when covered with non-calcifying epiphytic algae, and its diffusive boundary layer is thicker than when not covered by epiphytes. By modifying the microenvironmental carbonate chemistry, epiphytes, facilitated by OA, create micro-scale shield (and refuge) with more basic conditions that may allow the persistence of corallines associated with them during acidified conditions. Such ecological refugia could also assist corallines under near-future anthropogenic OA conditions.

Multiple transgressions and slow evolution shape the phylogeographic pattern of the blind cave-dwelling shrimp Typhlocaris.

BACKGROUND: Aquatic subterranean species often exhibit disjunct distributions, with high level of endemism and small range, shaped by vicariance, limited dispersal, and evolutionary rates. We studied the disjunct biogeographic patterns of an endangered blind cave shrimp, Typhlocaris, and identified the geological and evolutionary processes that have shaped its divergence pattern. METHODS: We collected Typlocaris specimens of three species (T. galilea, T. ayyaloni, and T. salentina), originating from subterranean groundwater caves by the Mediterranean Sea, and used three mitochondrial genes (12S, 16S, cytochrome oxygnese subunit 1 (COI)) and four nuclear genes (18S, 28S, internal transcribed spacer, Histon 3) to infer their phylogenetic relationships. Using the radiometric dating of a geological formation (Bira) as a calibration node, we estimated the divergence times of the Typhlocaris species and the molecular evolution rates. RESULTS: The multi-locus ML/Bayesian trees of the concatenated seven gene sequences showed that T. salentina (Italy) and T. ayyaloni (Israel) are sister species, both sister to T. galilea (Israel). The divergence time of T. ayyaloni and T. salentina from T. galilea was 7.0 Ma based on Bira calibration. The divergence time of T. ayyaloni from T. salentina was 5.7 (4.4-6.9) Ma according to COI, and 5.8 (3.5-7.2) Ma according to 16S. The computed interspecific evolutionary rates were 0.0077 substitutions/Myr for COI, and 0.0046 substitutions/Myr for 16S. DISCUSSION: Two consecutive vicariant events have shaped the phylogeographic patterns of Typhlocaris species. First, T. galilea was tectonically isolated from its siblings in the Mediterranean Sea by the arching uplift of the central mountain range of Israel ca. seven Ma. Secondly, T. ayyaloni and T. salentina were stranded and separated by a marine transgression ca. six Ma, occurring just before the Messinian Salinity Crisis. Our estimated molecular evolution rates were in one order of magnitude lower than the rates of closely related crustaceans, as well as of other stygobiont species. We suggest that this slow evolution reflects the ecological conditions prevailing in the highly isolated subterranean water bodies inhabited by Typhlocaris.

Diverse effects of invasive ecosystem engineers on marine biodiversity and ecosystem functions: A global review and meta-analysis.

Invasive ecosystem engineers (IEE) are potentially one of the most influential types of biological invaders. They are expected to have extensive ecological impacts by altering the physical-chemical structure of ecosystems, thereby changing the rules of existence for a broad range of resident biota. To test the generality of this expectation, we used a global systematic review and meta-analysis to examine IEE effects on the abundance of individual species and communities, biodiversity (using several indices) and ecosystem functions, focusing on marine and estuarine environments. We found that IEE had a significant effect (positive and negative) in most studies testing impacts on individual species, but the overall (cumulative) effect size was small and negative. Many individual studies showed strong IEE effects on community abundance and diversity, but the direction of effects was variable, leading to statistically non-significant overall effects in most categories. In contrast, there was a strong overall effect on most ecosystem functions we examined. IEE negatively affected metabolic functions and primary production, but positively affected nutrient flux, sedimentation and decomposition. We use the results to develop a conceptual model by highlighting pathways whereby IEE impact communities and ecosystem functions, and identify several sources of research bias in the IEE-related invasion literature. Only a few of the studies simultaneously quantified IEE effects on community/diversity and ecosystem functions. Therefore, understanding how IEE may alter biodiversity-ecosystem function relationships should be a primary focus of future studies of invasion biology. Moreover, the clear effects of IEE on ecosystem functions detected in our study suggest that scientists and environmental managers ought to examine how the effects of IEE might be manifested in the services that marine ecosystems provide to humans.

Future warming and acidification effects on anti-fouling and anti-herbivory traits of the brown alga Fucus vesiculosus (Phaeophyceae).

Human-induced ocean warming and acidification have received increasing attention over the past decade and are considered to have substantial consequences for a broad range of marine species and their interactions. Understanding how these interactions shift in response to climate change is particularly important with regard to foundation species, such as the brown alga Fucus vesiculosus. This macroalga represents the dominant habitat former on coastal rocky substrata of the Baltic Sea, fulfilling functions essential for the entire benthic community. Its ability to withstand extensive fouling and herbivory regulates the associated community and ecosystem dynamics. This study tested the interactive effects of future warming, acidification, and seasonality on the interactions of a marine macroalga with potential foulers and consumers. F. vesiculosus rockweeds were exposed to different combinations of conditions predicted regionally for the year 2100 (+∆5°C, +∆700 µatm CO2 ) using multifactorial long-term experiments in novel outdoor benthic mesocosms ("Benthocosms") over 9-12-week periods in four seasons. Possible shifts in the macroalgal susceptibility to fouling and consumption were tested using consecutive bioassays. Algal susceptibility to fouling and grazing varied substantially among seasons and between treatments. In all seasons, warming predominantly affected anti-fouling and anti-herbivory interactions while acidification had a subtle nonsignificant influence. Interestingly, anti-microfouling activity was highest during winter under warming, while anti-macrofouling and anti-herbivory activities were highest in the summer under warming. These contrasting findings indicate that seasonal changes in anti-fouling and anti-herbivory traits may interact with ocean warming in altering F. vesiculosus community composition in the future.