Effects of capability for dispersal on the evolution of diversity in antarctic benthos

The likelihood of marine invertebrates to maintain large geographic ranges is widely dependent on the ability of their early ontogenetic stages to disperse over long distances. Marine benthic invertebrates inhabiting the cold-stenothermal environment of the Southern Ocean are known for their overall reduced number of pelagic larvae, or drifting stages of any kind, when compared with organisms elsewhere in the sea. The diversity of organisms thriving in Antarctic waters is the result of evolution in situ and of the intrusion of species from surrounding seas. The reasons for a high level of endemism and a stunning diversity of benthic invertebrates found today are frequently discussed in the literature, but the mechanisms whereby diversity has been controlled over time remain largely theoretical. Here, I suggest that, indeed, early life-history patterns play a key role in defining the radiation and the speciation potential of Antarctic benthic invertebrates. In arguing this case, I synthesize the growing body of molecular studies on population connectivity in Antarctic benthic invertebrates, and compare this information with knowledge of their life histories and biogeography. I conclude that differences in early life-history patterns are key to the resilience potential of species in response to late Cenozoic glacial periods and propose that there is a direct relationship between rate of speciation and the ability of taxa to disperse.     

Invasion of the Atlantic rock crab (Cancer irroratus) at high latitudes

With the increase in global oceanic trade the establishment of non-indigenous marine organisms has become a major environmental and economic problem worldwide. Recently, the Atlantic rock crab (Cancer irroratus) was reported in Icelandic waters, Eastern North Atlantic. This is the first record of this relatively large crab species outside its natural range, i.e. the east coast of North America. The crab was most likely transferred to Iceland as larvae in ballast water and has successfully established a reproducing population in Icelandic waters. The species is distributed along the southwestern- and western-coast of Iceland. Adult specimens are now common in Faxaflói Bay, Southwest Iceland, but with sporadic occurrences in western and northwestern Icelandic waters. The green crab (Carcinus maenas) and the spider crab (Hyas araneus) are the only native brachyuran decapod species commonly found in its new habitat, but despite its recent colonization the rock crab was the most abundant brachyuran in the areas studied in southwest Iceland. Egg bearing rock crab and green crab females were found from June to October, while egg bearing spider crab females were seen from July to December. In Southwest Iceland both rock crab and green crab larvae were abundant in mid-summer but rare in both spring and autumn, which is opposite of what was observed for the spider crab. The size and abundance of adult crabs, their reproductive conditions, and occurrence of all larval stages, indicate that the Atlantic rock crab has successfully colonized Iceland.     

Patterns of benthic mega-invertebrate habitat associations in the Pacific Northwest continental shelf waters

As human impacts and demands for ocean space increase (fisheries, aquaculture, marine reserves, renewable energy), identification of marine habitats hosting sensitive biological assemblages has become a priority. Epifaunal invertebrates, especially the structure-forming species, are an increasing conservation concern as many traditional (bottom-contact fishing) and novel (marine renewable energy) ocean uses have the potential to displace or otherwise impact these slow-growing organisms. The differences in mega-invertebrate species assemblages between high-relief rocks and low-relief sediments are well documented and likely hold for most marine environments. In anticipation of potential development of marine renewable energy faculties off Oregon and Washington (USA), a survey of the benthic invertebrate assemblages and habitats was conducted on the continental shelf of the Pacific Northwest, using video footage collected by ROV, to more finely characterize these assemblage–habitat associations. Four main associations were found: pure mud/sand dominated by sea whips and burrowing brittle stars; mixed mud–rock (which may be further divided based on size of mixed-in rocks) characterized by various taxa at small densities; consolidated rocks characterized by high diversity and density of sessile or motile mega-invertebrates; and rubble rocks showing less diversity and density than the consolidated rocks, possibly due to the disturbance generated by movement of the unconsolidated rocks. The results of this study will help classify and map the seafloor in a way that represents benthic habitats reflective of biological species assemblage distributions, rather than solely geological features, and support conservation and management planning.     

Parallel Evolution,not Always so Parallel: Comparison of Small Benthic Charr, <Emphasis Type="Italic">Salvelinus alpinus</Emphasis>, from Grímsnes and Thingvallavatn,Iceland

Synopsis Iceland is unique in terms of geologically young freshwater systems and rapid adaptations of fresh water fishes to diverse habitats, e.g. lava with ground water flow. Iceland has six species of freshwater fishes, including Arctic charr, Salvelinus alpinus. Previous research has shown great diversity within this species. Four different morphs of Arctic charr are found in one lake, Thingvallavatn, including a small benthivorous charr. Similar populations of small benthic charr are known from several other Icelandic freshwater locations, including Nautavakir in Grímsnes. Our comparison of the small benthic charr morphs in Thingvallavatn and in Grímsnes showed that they are similar in morphology but distinguishable in several characteristics. Small benthic charr in Grímsnes and Thingvallavatn demonstrate similar adaptations and are an example of parallel evolution. However, subtle morphological differences between them indicate further specialized adaptations at each location.     

Global reductions in seafloor biomass in response to climate change

Seafloor organisms are vital for healthy marine ecosystems, contributing to elemental cycling, benthic remineralization, and ultimately sequestration of carbon. Deep‐sea life is primarily reliant on the export flux of particulate organic carbon from the surface ocean for food, but most ocean biogeochemistry models predict global decreases in export flux resulting from 21st century anthropogenically induced warming. Here we show that decadal‐to‐century scale changes in carbon export associated with climate change lead to an estimated 5.2% decrease in future (2091–2100) global open ocean benthic biomass under RCP8.5 (reduction of 5.2 Mt C) compared with contemporary conditions (2006–2015). Our projections use multi‐model mean export flux estimates from eight fully coupled earth system models, which contributed to the Coupled Model Intercomparison Project Phase 5, that have been forced by high and low representative concentration pathways (RCP8.5 and 4.5, respectively). These export flux estimates are used in conjunction with published empirical relationships to predict changes in benthic biomass. The polar oceans and some upwelling areas may experience increases in benthic biomass, but most other regions show decreases, with up to 38% reductions in parts of the northeast Atlantic. Our analysis projects a future ocean with smaller sized infaunal benthos, potentially reducing energy transfer rates though benthic multicellular food webs. More than 80% of potential deep‐water biodiversity hotspots known around the world, including canyons, seamounts, and cold‐water coral reefs, are projected to experience negative changes in biomass. These major reductions in biomass may lead to widespread change in benthic ecosystems and the functions and services they provide.     

Invertebrate biodiversity in cold groundwater fissures in Iceland

Iceland has an abundance of fissures that are parallel to the Mid‐Atlantic Ridge where bedrock cracks as a result of continental rifting. Some fissures penetrate the aquifer and expose the groundwater within the bedrock, becoming springs. As such, groundwater fissures have uniform and constant physical and chemical environment but they can differ greatly in morphology. In addition, there is often great variation in depth within fissures and substrate types contrast between vertical rock wall and more heterogenous horizontal bottom. The variation in morphological environment may create dissimilar habitats with unique characteristics and/or influence distribution of resources. Our objective was to study macrozoobenthos communities in cold groundwater fissures in Iceland in relation to physical habitat by comparing invertebrate diversity and density both between fissures with different morphological characteristics as well as between substrate types and depths within fissures. Samples were collected in two fissures in SW Iceland, Silfra and Flosagjá. Assemblages were similar between fissures except for higher densities of cladocerans in Flosagjá fissure. Within fissures, there was significant difference in Shannon diversity between substrate types in Flosagjá, and ostracods were found in significantly higher densities on the bottom. The distribution of all other taxa groups was homogenous in both fissures regardless of depth gradient and substrate. Invertebrates were found to be living within and around a biofilm that covered the entire substrate. These biofilm mats are made from Cyanobacteria and benthic diatoms, which are successful under low light conditions and may minimize any effect of the heterogeneous habitat creating a uniform and suitable microhabitat for invertebrates regardless of depth and substrate type.     

Effects of dredging on critical ecological processes for marine invertebrates,seagrasses and macroalgae,and the potential for management with environmental windows using Western Australia as a case study

Dredging can have significant impacts on benthic marine organisms through mechanisms such as sedimentation and reduction in light availability as a result of increased suspension of sediments. Phototrophic marine organisms and those with limited mobility are particularly at risk from the effects of dredging. The potential impacts of dredging on benthic species depend on biological processes including feeding mechanism, mobility, life history characteristics (LHCs), stage of development and environmental conditions. Environmental windows (EWs) are a management technique in which dredging activities are permitted during specific periods throughout the year; avoiding periods of increased vulnerability for particular organisms in specific locations. In this review we identify these critical ecological processes for temperate and tropical marine benthic organisms; and examine if EWs could be used to mitigate dredging impacts using Western Australia (WA) as a case study. We examined LHCs for a range of marine taxa and identified, where possible, their vulnerability to dredging. Large gaps in knowledge exist for the timing of LHCs for major species of marine invertebrates, seagrasses and macroalgae, increasing uncertainty around their vulnerability to an increase in suspended sediments or light attenuation. We conclude that there is currently insufficient scientific basis to justify the adoption of generic EWs for dredging operations in WA for any group of organisms other than corals and possibly for temperate seagrasses. This is due to; 1) the temporal and spatial variation in the timing of known critical life history stages of different species; and 2) our current level of knowledge and understanding of the critical life history stages and characteristics for most taxa and for most areas being largely inadequate to justify any meaningful EW selection. As such, we suggest that EWs are only considered on a case-by-case basis to protect ecologically or economically important species for which sufficient location-specific information is available, with consideration of probable exposures associated with a given mode of dredging.     

Feeding ecology of mackerel and dietary overlap with herring in Icelandic waters

The Northeast Atlantic mackerel (Scomber scombrus) has been extending its summer feeding distribution north and west, including around Iceland, since around 2006. The objective of this study is to quantify the weight gain and total food consumption of mackerel and to evaluate the food competition between mackerel and herring (Clupea harengus) feeding in the marine ecosystem around Iceland during the summers 2009–2011. Mackerel feeding in Icelandic waters gained around 43% on average in weight during these summers. Based on swept-area abundance estimates of mackerel from an international survey in 2011 and available estimates of food conversion efficiency in mackerel, the weight gain in Icelandic waters in 2011 corresponded to a total consumption of around 3.4 (2.4?4.5) million tonnes. Overall, 98% of 2314 mackerel, 91% of 398 Icelandic summer-spawning herring and 96% of 424 Norwegian spring-spawning herring stomachs, collected over the summers 2009?2011, contained food. The mean weight of the stomach content of mackerel was higher than for herring in all the years. While the stomach content weight of mackerel was generally highest in southeastern and southwestern areas, it was highest for herring in western, eastern and northern areas. Analysis of stomach contents showed that Copepoda were the most important food of mackerel in most areas, while Copepoda and Euphausiacea were the most important food items for herring. Fish prey contributed a higher proportion in stomachs of mackerel than herring, although relatively low for both species.     

Diversity and genomics of Antarctic marine micro-organisms

Marine bacterioplanktons are thought to play a vital role in Southern Ocean ecology and ecosystem function, as they do in other ocean systems. However, our understanding of phylogenetic diversity, genome-enabled capabilities and specific adaptations to this persistently cold environment is limited. Bacterioplankton community composition shifts significantly over the annual cycle as sea ice melts and phytoplankton bloom. Microbial diversity in sea ice is better known than that of the plankton, where culture collections do not appear to represent organisms detected with molecular surveys. Broad phylogenetic groupings of Antarctic bacterioplankton such as the marine group I Crenarchaeota, alpha-Proteobacteria (Roseobacter-related and SAR-11 clusters), gamma-Proteobacteria (both cultivated and uncultivated groups) and Bacteriodetes-affiliated organisms in Southern Ocean waters are in common with other ocean systems. Antarctic SSU rRNA gene phylotypes are typically affiliated with other polar sequences. Some species such as Polaribacter irgensii and currently uncultivated gamma-Proteobacteria (Ant4D3 and Ant10A4) may flourish in Antarctic waters, though further studies are needed to address diversity on a larger scale. Insights from initial genomics studies on both cultivated organisms and genomes accessed through shotgun cloning of environmental samples suggest that there are many unique features of these organisms that facilitate survival in high-latitude, persistently cold environments.     

Gradient in seaweed vegetation patterns along the North Icelandic coast,related to hydrographic conditions

The benthic algal vegetation of the North Icelandic coast exhibits particular features, different from those found in southern and western Iceland on the one hand and in eastern Iceland on the other. It is characterised by low-eulittoral belts of Devaleraea ramentacea, Petalonia species and Chordaria flagelliformis as well as by meadows of diverse Acrosiphonia species. In the tide pools Atlantic and typical North Icelandic associations occur side by side. This area thus represents an intermediate region between Atlantic and subarctic growth conditions. A certain gradient in the west-east direction was observed within the North Icelandic vegetation type and is possibly related to the gradual cooling and dilution of the water masses which pass the North Icelandic shelf. The hydrographic discontinuity in the extreme NW creates a sharp floristic and vegetational boundary for benthic algae, whereas in the NE a gradual transition towards the subarctic East Icelandic vegetation was observed. The Atlantic character of the vegetation was more pronounced in the western than in the eastern part of the north coast. An enclave of warm water vegetation was interposed along the western banks of the Tjörnes peninsula.This paper is based on a presentation at the XIII^th International Seaweed Symposium in Vancouver, Canada.     

Plant survival in Iceland during periods of glaciation?

Aim The paper addresses the classical question of possible plant survival in Iceland during the last glacial period in the light of a palaeobotanical record from northern Iceland, spanning the period 11,300–9000 BP , including the Younger Dryas stadial. We review the Late Cenozoic fossil plant record, the past debate on glacial plant refugia in Iceland, and the evidence for ice-free areas during the Weichselian. Location The investigated lake sediment record comes from Lake Torfadalsvatn, which is situated in the northwestern part of the Skagi peninsula in northern Iceland. Methods The sediment chronology was constructed from the cccurrence of the Vedde Ash and the Saksunarvatn ash, two well-dated Icelandic tephras, together with the results from five AMS and conventional radiocarbon dates performed on bulk sediment samples. The vegetational reconstruction was based on detailed pollen analysis of the sediment sequence. Results The pollen analysis revealed that many of the taxa present in the area prior to the Younger Dryas stadial continued to produce pollen during that cold event. The more or less immediate reappearance of a few other pollen taxa at the Younger Dryas-Preboreal boundary suggests that these plants also survived, even if they did not produce sufficient pollen to be recorded during the Younger Dryas stadial. Main conclusions We conclude that the relatively high plant diversity found in high Arctic areas and present-day nunataks in Iceland and Greenland, together with the fact that many plant species were able to survive the Younger Dryas stadial on the Skagi peninsula, suggest that species with high tolerance for climate fluctuations also survived the whole Weichselian in Iceland. This conclusion is supported by recent palaeoclimatic data from ice-cores and deep-sea sediments, indicating that Icelandic climate during the last glacial was only occasionally slightly colder than during the Younger Dryas stadial.     

From fronds to fish: the use of indicators for ecological monitoring in marine benthic ecosystems,with case studies from temperate Western Australia

Ecological indicators are used for monitoring in marine habitats the world over. With the advent of Ecosystem Based Fisheries Management (EBFM), the need for cost effective indicators of environmental impacts and ecosystem condition has intensified. Here, we review the development, utilisation and analysis of indicators for monitoring in marine benthic habitats, and outline important advances made in recent years. We use the unique, speciose benthic system of Western Australia (WA) as a detailed case study, as the development of indicators for EBFM in this region is presently ongoing, and major environmental drivers (e.g. climate change) and fishing practices are currently influencing WA marine systems. As such, the work is biased towards, but not restricted to, indicators that may be important tools for EBFM, such as biodiversity surrogates and indicators of fishing pressure. The review aimed to: (1) provide a concise, up-to-date account of the use of ecological indicators in marine systems; (2) discuss the current, and potential, applications of indicators for ecological monitoring in WA; and (3) highlight priority areas for research and pressing knowledge gaps. We examined indicators derived from benthic primary producers, benthic invertebrates and fish to achieve these goals.     

Fine-scale parallel patterns in diversity of small benthic Arctic charr (Salvelinus alpinus) in relation to the ecology of lava/groundwater habitats

It is critical to study factors that are important for origin and maintenance of biological diversity. A comparative approach involving a large number of populations is particularly useful. We use this approach to study the relationship between ecological factors and phenotypic diversity in Icelandic Arctic charr (Salvelinus alpinus). Numerous populations of small benthic charr have evolved in lava springs in Iceland. These charr appear morphologically similar, but differ in important morphological features related to feeding. We found a clear relationship between diversity in morphology, diet, and ecological factors among populations. In particular, there were clear differences in morphology and diet between fish coming from habitats where the lava spring flowed on as a stream compared to habitats where the lava spring flowed into a pond. Our study shows that ecological factors are important for the origin and maintenance of biological diversity. The relationship between phenotype and ecological factors are observed on a fine scale, when comparing numerous populations that are phenotypically similar. This strongly suggests that for understanding, managing, and conserving biological diversity important ecological variables have to be taken into the account.     

Gene flow and lack of population differentiation in Atlantic cod, Gadus morhua L., from Iceland, and comparison of cod from Norway and Newfoundland

Restriction analysis of mitochondrial DNA was used to study genetic variation and geographic population structure of Atlantic cod from localities around Iceland. Gene phylogenies were constructed and geographic locations superimposed on these. The variation was not localized. Estimated gene flow was large. Thus, Atlantic cod in Iceland belong to a single genetic population. Analyses of published sequence variation of cod from Norway and Newfoundland showed that the extensive continuity of intraspecific phylogeny of cod in Iceland, which is very similar to other marine organisms with a similar life history, extends from Norway to Newfoundland and possibly to larger areas of the Atlantic.     

Effects of fishing and acidification‐related benthic mortality on the southeast Australian marine ecosystem

Oceanic uptake of anthropogenic carbon dioxide (CO₂) is altering the carbonate chemistry of seawater, with potentially negative consequences for many calcifying marine organisms. At the same time, increasing fisheries exploitation is impacting on marine ecosystems. Here, using increased benthic‐invertebrate mortality as a proxy for effects of ocean acidification, the potential impact of the two stressors of fishing and acidification on the southeast Australian marine ecosystem to year 2050 was explored. The individual and interaction effects of the two stressors on biomass and diversity were examined for the entire ecosystem and for regional assemblages. For 61 functional groups or species, the cumulative effects of moderate ocean acidification and fishing were additive (30%), synergistic (33%), and antagonistic (37%). Strong ocean acidification resulted in additive (22%), synergistic (40%), and antagonistic (38%) effects. The greatest impact was on the demersal food web, with fishing impacting predation and acidification affecting benthic production. Areas that have been subject to intensive fishing were the most susceptible to acidification effect, although fishing also mitigated some of the decline in biodiversity observed with moderate acidification. The model suggested that ocean acidification and long‐term fisheries exploitation could act synergistically with the increasing sensitivity to change from long‐term (decades) fisheries exploitation potentially causing unexpected restructuring of the pelagic and demersal food webs. Major regime shifts occur around year 2040. Greater focus is needed on how differential fisheries exploitation of marine resources may exacerbate or accelerate effects of environmental changes such as ocean acidification.     

Diversity of urea‐degrading microorganisms in open‐ocean and estuarine planktonic communities

Urea is an important and dynamic natural component of marine nitrogen cycling and also a major contributor to anthropogenic eutrophication of coastal ecosystems, yet little is known about the identities or diversity of ureolytic marine microorganisms. Primers targeting the gene encoding urease were used to PCR‐amplify, clone and sequence 709 urease gene fragments from 31 plankton samples collected at both estuarine and open‐ocean locations. Two hundred and eighty‐six amplicons belonged to 22 distinct sequence types that were closely enough related to named organisms to be identified, and included urease sequences both from typical marine planktonic organisms and from bacteria usually associated with terrestrial habitats. The remaining 423 amplicons were not closely enough related to named organisms to be identified, and belonged to 96 distinct sequence types of which 43 types were found in two or more different samples. The distributions of unidentified urease sequence types suggested that some represented truly marine microorganisms while others reflected terrestrial inputs to low‐salinity estuarine areas. The urease primers revealed this great diversity of ureolytic organisms because they were able to amplify many previously unknown, environmentally relevant urease genes, and they will support new approaches for exploring the role of urea in marine ecosystems.     

Beyond corals and fish: the effects of climate change on noncoral benthic invertebrates of tropical reefs

Climate change is threatening tropical reefs across the world, with most scientists agreeing that the current changes in climate conditions are occurring at a much faster rate than in the past and are potentially beyond the capacity of reefs to adapt and recover. Current research in tropical ecosystems focuses largely on corals and fishes, although other benthic marine invertebrates provide crucial services to reef systems, with roles in nutrient cycling, water quality regulation, and herbivory. We review available information on the effects of environmental conditions associated with climate change on noncoral tropical benthic invertebrates, including inferences from modern and fossil records. Increasing sea surface temperatures may decrease survivorship and increase the developmental rate, as well as alter the timing of gonad development, spawning, and food availability. The broad latitudinal distribution and associated temperature ranges of several pantropical taxa suggest that some reef communities may have an in‐built adaptive capacity. Tropical benthic invertebrates will also show species‐specific sublethal and lethal responses to sea‐level rise, ocean acidification, physical disturbance, runoff, turbidity, sedimentation, and changes in ocean circulation. In order to accurately predict a species' response to these stressors, we must consider the magnitude and duration of exposure to each stressor, as well as the physiology, mobility, and habitat requirements of the species. Stressors will not act independently, and many organisms will be exposed to multiple stressors concurrently, including anthropogenic stressors. Environmental changes associated with climate change are linked to larger ecological processes, including changes in larval dispersal and recruitment success, shifts in community structure and range extensions, and the establishment and spread of invasive species. Loss of some species will trigger economic losses and negative effects on ecosystem function. Our review is intended to create a framework with which to predict the vulnerability of benthic invertebrates to the stressors associated with climate change, as well as their adaptive capacity. We anticipate that this review will assist scientists, managers, and policy‐makers to better develop and implement regional research and management strategies, based on observed and predicted changes in environmental conditions.     

Harsh climate selects for small body size among Iceland's Arctic foxes

We studied the effect of the two environmental indices, the sub‐polar gyre (SPG), and winter and summer North Atlantic Oscillation (NAO), together with mean annual winter and summer temperatures and geographic location on mandible size and body mass of Arctic foxes in Iceland (6345 and 2732 specimens, respectively) during the year of their death. We predicted that when favorable conditions prevailed, large specimens would be selected for, and vice versa. Body size and body mass were significantly affected by the environmental parameters (i.e. SPG, NAO, ambient temperature and cloud cover) prevailing during the year of death. The effect of environmental conditions on body size was much stronger in the less productive region of eastern Iceland, apparently because in areas where food availability is meager, even a small difference in climate may tilt the balance from food sufficiency to food shortage. Western Iceland comprises only a quarter of the total surface area of the country, but its productive seashores are twice as long as those of all the rest of the country combined. It is interesting to note that the effect of the SPG, a marine phenomenon in the oceans surrounding Iceland, is reflected in the condition of the foxes more than the other climatic variables we used in this study, which are largely land‐related. Because Arctic foxes in Iceland feed largely on marine birds and invertebrates, the SPG seems to encompass more accurate information regarding the direct ocean forces that affect food availability to the foxes.     

Possible effects of global environmental changes on Antarctic benthos: a synthesis across five major taxa

Because of the unique conditions that exist around the Antarctic continent, Southern Ocean (SO) ecosystems are very susceptible to the growing impact of global climate change and other anthropogenic influences. Consequently, there is an urgent need to understand how SO marine life will cope with expected future changes in the environment. Studies of Antarctic organisms have shown that individual species and higher taxa display different degrees of sensitivity to environmental shifts, making it difficult to predict overall community or ecosystem responses. This emphasizes the need for an improved understanding of the Antarctic benthic ecosystem response to global climate change using a multitaxon approach with consideration of different levels of biological organization. Here, we provide a synthesis of the ability of five important Antarctic benthic taxa (Foraminifera, Nematoda, Amphipoda, Isopoda, and Echinoidea) to cope with changes in the environment (temperature, pH, ice cover, ice scouring, food quantity, and quality) that are linked to climatic changes. Responses from individual to the taxon-specific community level to these drivers will vary with taxon but will include local species extinctions, invasions of warmer-water species, shifts in diversity, dominance, and trophic group composition, all with likely consequences for ecosystem functioning. Limitations in our current knowledge and understanding of climate change effects on the different levels are discussed.     

Biodiversity gradient in the Baltic Sea: a comprehensive inventory of macrozoobenthos data

In the Helsinki Commission Red List project 2009–2012, taxonomic and distributional data of benthic (macro) invertebrates were compiled by the present authors in a comprehensive checklist of the Baltic Sea fauna. Based on the most recent and comprehensive data, this paper presents the diversity patterns observed among benthic invertebrates in the Baltic Sea. As expected, the total number of species per sub-region generally declined along the salinity gradient from the Danish Straits to the northern Baltic Sea. This relationship is well known from the Baltic Sea and has resulted in a general assumption of an exponentially positive relationship between species richness and salinity for marine species, and a negative relationship for freshwater species. In 1934, Remane produced a diagram to describe the hypothetical distribution of benthic invertebrate diversity along a marine–freshwater salinity gradient. Our results clearly indicated the validity of this theory for the macrozoobenthic diversity pattern within the Baltic Sea. Categorisation of sub-regions according to species composition showed both separation and grouping of some sub-regions and a strong alignment of similarity patterns of zoobenthic species composition along the salinity gradient.