Quantifying impacts of human pressures on ecosystem services: Effects of widespread non-indigenous species in the Baltic Sea.

Ecosystem services (ES) are the benefits natural ecosystems provide to society, such as food provisioning, water supply, climate regulation and recreational benefits. Biological invasions are a major driver of global change, and several non-indigenous species (NIS) may alter key ecological feedbacks with ultimate consequences to ES, livelihoods and human wellbeing. Nonetheless, the effects of NIS on ES supply remain largely unquantified. Here we present the first quantitative case study assessing the impacts of widespread NIS on ES in the Baltic Sea, by developing and employing a robust and repeatable data-driven approach. All NIS with a sufficient knowledge base pose large and highly significant effects on ES, resulting on average 55 % change in the intensity of ES. Most impacts affected regulation services, concerning both abiotic and biotic realms, with little evidence on cultural and provisioning services. The methodology can be easily employed beyond the current study realm e.g. to better understand the roles of human pressures on ES in any ecosystem. Importantly, the study also identified major biases not only in the availability of taxonomic and sub-regional evidence, but also in the different study types employed to create the evidence base.

Perceived multiple stressor effects depend on sample size and stressor gradient length.

Multiple stressors are continuously deteriorating surface waters worldwide, posing many challenges for their conservation and restoration. Combined effect types of multiple stressors range from single-stressor dominance to complex interactions. Identifying prevalent combined effect types is critical for environmental management, as it helps to prioritise key stressors for mitigation. However, it remains unclear whether observed single and combined stressor effects reflect true ecological processes unbiased by sample size and length of stressor gradients. Therefore, we examined the role of sample size and stressor gradient lengths in 158 paired-stressor response cases with over 120,000 samples from rivers, lakes, transitional and marine ecosystems around the world. For each case, we split the overall stressor gradient into two partial gradients (lower and upper) and investigated associated changes in single and combined stressor effects. Sample size influenced the identified combined effect types, and stressor interactions were less likely for cases with fewer samples. After splitting gradients, 40 % of cases showed a change in combined effect type, 30 % no change, and 31 % showed a loss in stressor effects. These findings suggest that identified combined effect types may often be statistical artefacts rather than representing ecological processes. In 58 % of cases, we observed changes in stressor effect directions after the gradient split, suggesting unimodal stressor effects. In general, such non-linear responses were more pronounced for organisms at higher trophic levels. We conclude that observed multiple stressor effects are not solely determined by ecological processes, but also strongly depend on sampling design. Observed effects are likely to change when sample size and/or gradient length are modified. Our study highlights the need for improved monitoring programmes with sufficient sample size and stressor gradient coverage. Our findings emphasize the importance of adaptive management, as stress reduction measures or further ecosystem degradation may change multiple stressor-effect relationships, which will then require associated changes in management strategies.

Assessing cumulative impacts of human-induced pressures on reef and sandbank habitats and associated biotopes in the northeastern Baltic Sea.

Marine ecosystems are impacted by multiple individual and combined anthropogenic pressures. We used meta-analysis and data-driven PlanWise4Blue decision support tool to predict individual and combined impacts of wind park development, nutrient loading, and invasive species on vulnerable reef and sandbank habitats and associated species-specific biotopes in the northeastern Baltic Sea. Many impacts were not statistically significant due to large between-study variance in effect sizes. Wind park development is predicted to have less impact than nutrient loading and invasive species. Predicted impacts varied greatly among larger-scale habitats versus smaller-scale biotopes with impacts being generally stronger at small scale. Excessive nutrient loading damages algae-based biotopes, the presence of nonnative species has substantial negative impacts on larger-scale reef and sandbank habitats. The results showed that a 25 % reduction of nutrient loading improves all examined benthic habitats, whereas nonnative species, which cannot be removed from ecosystems, pose a significant threat to these habitats.

Persistence in a tropical transition zone? Sargassum forests alternate seasonal growth forms to maintain productivity in warming waters at the expense of annual biomass production.

Macroalgal forests provide productivity and biomass that underpins the function of many coastal ecosystems globally. The phenology of forests is seasonally driven by environmental conditions, with the environment-productivity relationship understood for most coastlines of the world. Climatic transition zones, however, have characteristics of temperate and tropical regions, creating large fluctuations in environmental conditions, and potentially limiting productivity and the persistence of macroalgal forests. The response of a forest-forming, dimorphic seaweed (Sargassum hemiphyllum) to seasonal temperature and light conditions in a rapidly warming tropical-temperate transitional zone (Hong Kong) was quantified by measuring in situ growth, net primary productivity (NPP), respiration, and photosynthetic potential. These physiological responses of S. hemiphyllum were then experimentally tested in response to changing temperatures (16.5-27 °C) and irradiances (20, 110, and 300 µmol m-2 s-1) in laboratory mesocosms. In contrast to predictions, S. hemiphyllum demonstrated asynchronous NPP and growth patterns, with growth maximized in cooler conditions but, counter-intuitively, highest photosynthetic rates in summer after annual senescence and dormancy were established. This discrepancy between peak photosynthetic rates and growth may provide regional populations of S. hemiphyllum the ability to survive higher temperatures in the near future, resisting the predicted range shifts under ocean warming. In contrast, warming is likely to drive a shorter growth season, longer dormancy, and reduced annual biomass production in bi-phasic seaweeds inhabiting climatic transition zones, potentially reducing system-wide productivity of these algal forests.

Assessing the potential for sea-based macroalgae cultivation and its application for nutrient removal in the Baltic Sea.

Marine eutrophication is a pervasive and growing threat to global sustainability. Macroalgal cultivation is a promising circular economy solution to achieve nutrient reduction and food security. However, the location of production hotspots is not well known. In this paper the production potential of macroalgae of high commercial value was predicted across the Baltic Sea region. In addition, the nutrient limitation within and adjacent to macroalgal farms was investigated to suggest optimal site-specific configuration of farms. The production potential of Saccharina latissima was largely driven by salinity and the highest production yields are expected in the westernmost Baltic Sea areas where salinity is >23. The direct and interactive effects of light availability, temperature, salinity and nutrient concentrations regulated the predicted changes in the production of Ulva intestinalis and Fucus vesiculosus. The western and southern Baltic Sea exhibited the highest farming potential for these species, with promising areas also in the eastern Baltic Sea. Macroalgal farming did not induce significant nutrient limitation. The expected spatial propagation of nutrient limitation caused by macroalgal farming was less than 100-250 m. Higher propagation distances were found in areas of low nutrient and low water exchange (e.g. offshore areas in the Baltic Proper) and smaller distances in areas of high nutrient and high water exchange (e.g. western Baltic Sea and Gulf of Riga). The generated maps provide the most sought-after input to support blue growth initiatives that foster the sustainable development of macroalgal cultivation and reduction of in situ nutrient loads in the Baltic Sea.

Blueprint for the ideal microplastic effect study: Critical issues of current experimental approaches and envisioning a path forward.

This article presents a novel conceptual blueprint for an 'ideal', i.e., ecologically relevant, microplastic effect study. The blueprint considers how microplastics should be characterized and applied in laboratory experiments, and how biological responses should be measured to assure unbiased data that reliably reflect the effects of microplastics on aquatic biota. This 'ideal' experiment, although practically unachievable, serves as a backdrop to improve specific aspects of experimental research on microplastic effects. In addition, a systematic and quantitative literature review identified and quantified departures of published experiments from the proposed 'ideal' design. These departures are related mainly to the experimental design of microplastic effect studies failing to mimic natural environments, and experiments with limited potential to be scaled-up to ecosystem level. To produce a valid and generalizable assessment of the effect of microplastics on biota, a quantitative meta-analysis was performed that incorporated the departure of studies from the 'ideal' experiment (a measure of experimental quality) and inverse variance (a measure of the study precision) as weighting coefficients. Greater weights were assigned to experiments with higher quality and/or with lower variance in the response variables. This double-weighting captures jointly the technical quality, ecological relevance and precision of estimates provided in each study. The blueprint and associated meta-analysis provide an improved baseline for the design of ecologically relevant and technically sound experiments to understand the effects of microplastics on single species, populations and, ultimately, entire ecosystems.

Meta-analysis on the ecological impacts of widely spread non-indigenous species in the Baltic Sea.

The introduction of non-indigenous species (NIS) is a major driver for global change in species biogeography, often associated with significant consequences for recipient ecosystems and services they provide for humans. Despite mandated by several high-level international legislative instruments, comprehensive quantitative evaluation on ecosystem impacts of marine NIS is scarce and lack a robust and data-driven assessment framework. The current study is aiming at fulfilling this gap, through quantitative assessment on the effects of the widespread NIS of the Baltic Sea on multiple ecosystem features and components including direct food-web effects. The outcomes of this study allowed identifying the most impacting widespread NIS, together with defining the processes underlying the most significant changes and outlined major sources of uncertainty. Lack and/or bias in the availability of evidence of impacts was recorded for several (both recent and early) introductions. Realizing a sophisticated, data and information-hungry framework for the evaluation of ecosystem impacts of NIS is not pragmatic for management purposes in the foreseeable future. Instead, simple approaches, such as application of common statistical parameters like absolute effect size, are more likely to result in tangible outcomes. As bearing no unit, effect sizes can be later easily aggregated across taxa, affected ecosystem features or spatial scales. The proposed approach enables performing systematic comparisons on the severity of impacts of different NIS along different study disciplines and ecosystems.

Seagrass beds reveal high abundance of microplastic in sediments: A case study in the Baltic Sea.

Microplastic (MPL) contamination in the marine environment is extensively studied yet little is known about the extent of MPL abundance in seagrass beds. The aim of this study was to evaluate MPL accumulation in coastal seagrass (Zostera marina) beds in the Baltic Sea, Estonia. Surface water was sampled by pumping using 40 µm plankton net, and sediments by trowel. MPL was extracted with NaCl, identified by microscopy and ATR-FTIR on selected samples. Surface water in the seagrass beds had 0.04-1.2 (median 0.14) MPL/L, similar to other areas of the Baltic Sea. Sediments had 0-1817 (median 208) MPL/kg (dwt), much higher than previously recorded from adjacent unvegetated and offshore sediments, thereby suggesting a strong ability of the sediments in seagrass beds to retain MPL. Of identified MPL, blue fibres were dominant in both the sampled media. Sediment characterization showed a correlation between MPL counts with poorly sorted sediments.

From ecosystems to socio-economic benefits: A systematic review of coastal ecosystem services in the Baltic Sea.

Seagrass meadows, algal forests and mussel beds are widely regarded as foundation species that support communities providing valuable ecosystem services in many coastal regions; however, quantitative evidence of the relationship is scarce. Using the Baltic Sea as a case study, a region of significant socio-economic importance in the northern hemisphere, we systematically synthesized the primary literature and summarized the current knowledge on ecosystem services derived from seagrass, macroalgae, and mussels (see animated video summary of the manuscript: Video abstract). We found 1740 individual ecosystem service records (ESR), 61% of which were related to macroalgae, 26% to mussel beds and 13% to seagrass meadows. The most frequently reported ecosystem services were raw material (533 ESR), habitat provision (262 ESR) and regulation of pollutants (215 ESR). Toxins (356 ESR) and nutrients (302 ESR) were the most well-documented pressures to services provided by coastal ecosystems. Next, we assessed the current state of knowledge as well as knowledge transfer of ecosystem services to policies through natural, social, human and economic dimensions, using a systematic scoring tool, the Eco-GAME matrix. We found good quantitative information about how ecosystems generated the service but almost no knowledge of how they translate into socio-economic benefits (8 out of 657 papers, 1.2%). While we are aware that research on Baltic Sea socio-economic benefits does exist, the link with ecosystems providing the service is mostly missing. To close this knowledge gap, we need a better analytical framework that is capable of directly linking existing quantitative information about ecosystem service generation with human benefit.

Predicting lake dissolved organic carbon at a global scale.

The pool of dissolved organic carbon (DOC), is one of the main regulators of the ecology and biogeochemistry of inland water ecosystems, and an important loss term in the carbon budgets of land ecosystems. We used a novel machine learning technique and global databases to test if and how different environmental factors contribute to the variability of in situ DOC concentrations in lakes. In order to estimate DOC in lakes globally we predicted DOC in each lake with a surface area larger than 0.1 km2. Catchment properties and meteorological and hydrological features explained most of the variability of the lake DOC concentration, whereas lake morphometry played only a marginal role. The predicted average of the global DOC concentration in lake water was 3.88 mg L-1. The global predicted pool of DOC in lake water was 729 Tg from which 421 Tg was the share of the Caspian Sea. The results provide global-scale evidence for ecological, climate and carbon cycle models of lake ecosystems and related future prognoses.

Trans-Atlantic Distribution and Introgression as Inferred from Single Nucleotide Polymorphism: Mussels Mytilus and Environmental Factors.

Large-scale climate changes influence the geographic distribution of biodiversity. Many taxa have been reported to extend or reduce their geographic range, move poleward or displace other species. However, for closely related species that can hybridize in the natural environment, displacement is not the only effect of changes of environmental variables. Another option is subtler, hidden expansion, which can be found using genetic methods only. The marine blue mussels Mytilus are known to change their geographic distribution despite being sessile animals. In addition to natural dissemination at larval phase-enhanced by intentional or accidental introductions and rafting-they can spread through hybridization and introgression with local congeners, which can create mixed populations sustaining in environmental conditions that are marginal for pure taxa. The Mytilus species have a wide distribution in coastal regions of the Northern and Southern Hemisphere. In this study, we investigated the inter-regional genetic differentiation of the Mytilus species complex at 53 locations in the North Atlantic and adjacent Arctic waters and linked this genetic variability to key local environmental drivers. Of seventy-nine candidate single nucleotide polymorphisms (SNPs), all samples were successfully genotyped with a subset of 54 SNPs. There was a clear interregional separation of Mytilus species. However, all three Mytilus species hybridized in the contact area and created hybrid zones with mixed populations. Boosted regression trees (BRT) models showed that inter-regional variability was important in many allele models but did not prevail over variability in local environmental factors. Local environmental variables described over 40% of variability in about 30% of the allele frequencies of Mytilus spp. For the 30% of alleles, variability in their frequencies was only weakly coupled with local environmental conditions. For most studied alleles the linkages between environmental drivers and the genetic variability of Mytilus spp. were random in respect to "coding" and "non-coding" regions. An analysis of the subset of data involving functional genes only showed that two SNPs at Hsp70 and ATPase genes correlated with environmental variables. Total predictive ability of the highest performing models (r2 between 0.550 and 0.801) were for alleles that discriminated most effectively M.trossulus from M.edulis and M.galloprovincialis, whereas the best performing allele model (BM101A) did the best at discriminating M.galloprovincialis from M. edulis and M.trossulus. Among the local environmental variables, salinity, water temperature, ice cover and chlorophyll a concentration were by far the greatest predictors, but their predictive performance varied among different allele models. In most cases changes in the allele frequencies along these environmental gradients were abrupt and occurred at a very narrow range of environmental variables. In general, regions of change in allele frequencies for M.trossulus occurred at 8-11 psu, 0-10 C, 60%-70% of ice cover and 0-2 mg m-3 of chlorophyll a, M. edulis at 8-11 and 30-35 psu, 10-14 C and 60%-70% of ice cover and for M.galloprovincialis at 30-35 psu, 14-20 C.

Cleaning up seas using blue growth initiatives: Mussel farming for eutrophication control in the Baltic Sea.

Eutrophication is a serious threat to aquatic ecosystems globally with pronounced negative effects in the Baltic and other semi-enclosed estuaries and regional seas, where algal growth associated with excess nutrients causes widespread oxygen free "dead zones" and other threats to sustainability. Decades of policy initiatives to reduce external (land-based and atmospheric) nutrient loads have so far failed to control Baltic Sea eutrophication, which is compounded by significant internal release of legacy phosphorus (P) and biological nitrogen (N) fixation. Farming and harvesting of the native mussel species (Mytilus edulis/trossulus) is a promising internal measure for eutrophication control in the brackish Baltic Sea. Mussels from the more saline outer Baltic had higher N and P content than those from either the inner or central Baltic. Despite their relatively low nutrient content, harvesting farmed mussels from the central Baltic can be a cost-effective complement to land-based measures needed to reach eutrophication status targets and is an important contributor to circularity. Cost effectiveness of nutrient removal is more dependent on farm type than mussel nutrient content, suggesting the need for additional development of farm technology. Furthermore, current regulations are not sufficiently conducive to implementation of internal measures, and may constitute a bottleneck for reaching eutrophication status targets in the Baltic Sea and elsewhere.

Geographic variation in fitness-related traits of the bladderwrack Fucus vesiculosus along the Baltic Sea-North Sea salinity gradient.

In the course of the ongoing global intensification and diversification of human pressures, the study of variation patterns of biological traits along environmental gradients can provide relevant information on the performance of species under shifting conditions. The pronounced salinity gradient, co-occurrence of multiple stressors, and accelerated rates of change make the Baltic Sea and its transition to North Sea a suitable region for this type of study. Focusing on the bladderwrack Fucus vesiculosus, one of the main foundation species on hard-bottoms of the Baltic Sea, we analyzed the phenotypic variation among populations occurring along 2,000 km of coasts subjected to salinities from 4 to >30 and a variety of other stressors. Morphological and biochemical traits, including palatability for grazers, were recorded at 20 stations along the Baltic Sea and four stations in the North Sea. We evaluated in a common modeling framework the relative contribution of multiple environmental drivers to the observed trait patterns. Salinity was the main and, in some cases, the only environmental driver of the geographic trait variation in F. vesiculosus. The decrease in salinity from North Sea to Baltic Sea stations was accompanied by a decline in thallus size, photosynthetic pigments, and energy storage compounds, and affected the interaction of the alga with herbivores and epibiota. For some traits, drivers that vary locally such as wave exposure, light availability or nutrient enrichment were also important. The strong genetic population structure in this macroalgae might play a role in the generation and maintenance of phenotypic patterns across geographic scales. In light of our results, the desalination process projected for the Baltic Sea could have detrimental impacts on F. vesiculosus in areas close to its tolerance limit, affecting ecosystem functions such as habitat formation, primary production, and food supply.

Integrating experimental and distribution data to predict future species patterns.

Predictive species distribution models are mostly based on statistical dependence between environmental and distributional data and therefore may fail to account for physiological limits and biological interactions that are fundamental when modelling species distributions under future climate conditions. Here, we developed a state-of-the-art method integrating biological theory with survey and experimental data in a way that allows us to explicitly model both physical tolerance limits of species and inherent natural variability in regional conditions and thereby improve the reliability of species distribution predictions under future climate conditions. By using a macroalga-herbivore association (Fucus vesiculosus - Idotea balthica) as a case study, we illustrated how salinity reduction and temperature increase under future climate conditions may significantly reduce the occurrence and biomass of these important coastal species. Moreover, we showed that the reduction of herbivore occurrence is linked to reduction of their host macroalgae. Spatial predictive modelling and experimental biology have been traditionally seen as separate fields but stronger interlinkages between these disciplines can improve species distribution projections under climate change. Experiments enable qualitative prior knowledge to be defined and identify cause-effect relationships, and thereby better foresee alterations in ecosystem structure and functioning under future climate conditions that are not necessarily seen in projections based on non-causal statistical relationships alone.

Knowledge to decision in dynamic seas: Methods to incorporate non-indigenous species into cumulative impact assessments for maritime spatial planning.

Incorporating ecosystem changes from non-indigenous species (NIS) is an important task of maritime spatial planning. Maritime spatial planning requires a framework that emphasises ecological functioning in a state of dynamic change, including changes to ecosystem services from functions introduced by new NIS. Adaptable modelling toolsets should be developed that can readily incorporate knowledge of new NIS. In the Baltic Sea, recent NIS examples are the North American mud crab Rhithropanopeus harrisii and the Ponto-Caspian round goby Neogobius melanostomus. We performed environmental niche modelling that predicted N. melanostomus will spread across large areas of the Baltic Sea coast while R. harrisii will be limited to regions with high temperature and low salinity conditions. We then performed a meta-analysis on literature showing effects in the Baltic Sea from these NIS and calculated the standardised effect-sizes on relevant ecosystem services. Half the impacts identified for N. melanostomus were considered to increase ecosystem service outcomes, while all R. harrisii impacts caused apparent decreases. Effect coefficients were incorporated into an online impact assessment tool developed by the Estonian Marine Institute. Users with or without science training can use the portal to estimate areas impacted and changes to natural assets (km2) caused by these NIS and cumulative effects from other pressure-types. Impact estimates are based on best available knowledge from manipulative and correlative experiments and thus form a link between science and management. Dynamic modelling techniques informed from varied ecological and methodological perspectives will effectively advise spatial planners about rapid maritime changes and mitigation actions to reduce NIS impacts especially in the focus areas.

Linking atmospheric, terrestrial and aquatic environments: Regime shifts in the Estonian climate over the past 50 years.

Climate change in recent decades has been identified as a significant threat to natural environments and human wellbeing. This is because some of the contemporary changes to climate are abrupt and result in persistent changes in the state of natural systems; so called regime shifts (RS). This study aimed to detect and analyse the timing and strength of RS in Estonian climate at the half-century scale (1966-2013). We demonstrate that the extensive winter warming of the Northern Hemisphere in the late 1980s was represented in atmospheric, terrestrial, freshwater and marine systems to an extent not observed before or after the event within the studied time series. In 1989, abiotic variables displayed statistically significant regime shifts in atmospheric, river and marine systems, but not in lake and bog systems. This was followed by regime shifts in the biotic time series of bogs and marine ecosystems in 1990. However, many biotic time series lacked regime shifts, or the shifts were uncoupled from large-scale atmospheric circulation. We suggest that the latter is possibly due to complex and temporally variable interactions between abiotic and biotic elements with ecosystem properties buffering biotic responses to climate change signals, as well as being affected by concurrent anthropogenic impacts on natural environments.

Description of a new species of Sabellidae (Polychaeta, Annelida) from fresh and brackish waters in Europe, with some remarks on the branchial crown of Laonome.

In 2009, a hitherto unknown Laonome species was found in the Canal Ghent-Terneuzen in the Netherlands and subsequently in other Dutch rivers, canals and estuaries. A few years later, more unknown Laonome specimens were found in the eastern part of the Baltic Sea and in the Don River estuary, Sea of Azov. Initially, it was assumed that these specimens could represent Laonome calida Capa, 2007, originally described from Australia. In the present study we examine all these unknown European Laonome specimens and compare these specimens with the type material of L. calida from Australia. This lead to two main results: First, all specimens from Europe have the same diagnostic characters and therefore belong to one species. This finding was also supported by the results of a correspondence analysis, and genetic analyses using four different DNA sequences (COI, 16S, 28S). Second, it turned out that the type material of L. calida contains two morphologically distinct groups of specimens. The holotype and 7 paratypes are similar to each other but differ significantly from the other also similar 16 paratypes, and from all European specimens. On the basis of these observations, the Laonome specimens from European waters are described here as L. xeprovala sp. nov. We also provide the characters of the branchial crown of three Laonome species for a prospective revision of this genus.

Predicting the cover and richness of intertidal macroalgae in remote areas: a case study in the Antarctic Peninsula.

Antarctica is an iconic region for scientific explorations as it is remote and a critical component of the global climate system. Recent climate change causes a dramatic retreat of ice in Antarctica with associated impacts to its coastal ecosystem. These anthropogenic impacts have a potential to increase habitat availability for Antarctic intertidal assemblages. Assessing the extent and ecological consequences of these changes requires us to develop accurate biotic baselines and quantitative predictive tools. In this study, we demonstrated that satellite-based remote sensing, when used jointly with in situ ground-truthing and machine learning algorithms, provides a powerful tool to predict the cover and richness of intertidal macroalgae. The salient finding was that the Sentinel-based remote sensing described a significant proportion of variability in the cover and richness of Antarctic macroalgae. The highest performing models were for macroalgal richness and the cover of green algae as opposed to the model of brown and red algal cover. When expanding the geographical range of the ground-truthing, even involving only a few sample points, it becomes possible to potentially map other Antarctic intertidal macroalgal habitats and monitor their dynamics. This is a significant milestone as logistical constraints are an integral part of the Antarctic expeditions. The method has also a potential in other remote coastal areas where extensive in situ mapping is not feasible.

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.