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We hypothesized that supply from macroalgal propagule banks may influence the relative abundance of annual and perennial algae and that this may alter the effects of grazers and nutrients on species composition. In a factorial field experiment in the Baltic Sea littoral system we tested the effects of manipulating propagule banks, the abundance of crustacean and gastropod grazers, and nutrient supply on recruitment and growth of macroalgae over a year. Moreover, we determined seasonal patterns of macroalgal propagule dispersal at the experimental site and quantified algal abundance and recruitment at 25 locations throughout the Baltic Sea. Experimental manipulations had minor effects on adults of the dominating perennial alga, Fucus vesiculosus. Instead, we found that species composition was determined by processes operating at early life stages. Propagule supply from a propagule bank strongly favored the fast-growing annual alga Enteromorpha spp. which then blocked settlement and recruitment of Fucus. Grazers reduced the abundance of annual algae and indirectly favored Fucus recruitment. There was an apparent trade-off between gains from the propagule bank and losses to herbivory in five of seven colonizing species. Nutrient enrichment overrode grazer control of annual algae and accelerated the decline of Fucus only when annual algae had already achieved high densities through the propagule bank. Corroborating the experimental findings, field surveys across the Baltic showed that Fucus recruit densities can be predicted from the cover of annual algae during the period of Fucus reproduction and settlement. Recruitment inhibition by annual algae, which is driven by the abundance of annuals in the propagule bank, increasing nutrient levels, and declining consumer control, is suggested as a mechanistic explanation of the current decline of perennial algae in the Baltic Sea.

In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment ‘‘bottleneck’’ at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (,4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size.

Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis. Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella. Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella. Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha, thereby negatively affecting Pilayella. Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.

Nutrient enrichment of marine sediments or the water column has been used to study plant nutrient limitation and its cascading effects on community structure, Here we develop methodological recommendations for in situ enrichment. We review 18 published enrichment methods. Nutrient concentrations varied through time and among sites, with sediment depth, distance from the source, fertilizer type and load. Combining available data, we could predict an increase in sediment porewater phosphate (r 2 = 0.48) but not ammonium (r 2 = 0.07) concentrations in a multiple regression model. In three comparative field experiments we applied a coated slow-release fertilizer in the sediment and the water column and followed nutrient concentrations over time. We recommend coated fertilizer pellets, because they provide gradual nutrient release, allow for realistic nutrient gradients, and even application but we emphasize that nutrient concentrations need to be monitored through time.

By factorial field experiments we analyzed the relative effects of increased nutrient (N1P) loading and natural grazing pressure on species composition, carbon storage, and nitrogen retention in the Baltic Sea littoral food web, composed of macroalgae, grazers (snails, isopods, amphipods), and predators (shrimps, crabs, fish). Nitrogen was depleted relative to phosphorus throughout most of the year. Increasing nitrogen (6‐200% over ambient concentrations) enhanced algal productivity and cover of fast-growing annual algae, grazer, and predator densities, suggesting a three-level bottom-up effect. With increasing nitrogen loading, annual algae increasingly blocked perennial algal recruitment (65‐98% decrease) and growth. Grazers counteracted the effects of nutrient enrichment on algal species composition through selective consumption of annual algae. Grazer exclusion had equivalent negative effects on perennial recruitment as a 85% increase in nitrogen loading. Nutrient enrichment increased algal nitrogen content and decreased tissue C : N ratios in spring and summer but not in fall. Carbon storage and nitrogen retention, measured as C and N retained in plant biomass at the end of the growth season, were increased by grazers (C: 39%, N: 24%) but decreased with increasing nitrogen loading (C: 271%, N: 274%). Our results emphasize the important role of grazers in buffering moderate eutrophication effects and illustrate how food web interactions and shifts in species composition are tightly linked to coastal ecosystem function. Currently, humans are more than doubling the rate at which nitrogen and phosphorus enter the global biogeochem

Generalisations on the combined effects of consumers and resources on autotrophs in aquatic food webs largely rely on freshwater studies. In this study, we tested these general concepts with marine benthic microalgae, which are important components of coastal food webs. We manipu- lated nitrogen availability and herbivore presence in a factorial field experiment in the Western Baltic Sea. Moreover, we investigated how herbivore control varied among 3 sites and 2 seasons and tested for trophic cascades by enhancing demersal fish density at 2 sites. Nitrogen availability and herbivore presence had strong and antagonistic effects on microalgal biomass, species composition and diver- sity. Herbivores significantly reduced algal biomass, whereas nutrient enrichment led to an increase in biomass. Herbivore effects on microalgal biomass increased with increasing nitrogen availability, indicating a functional response of herbivores to nutrient enrichment. The response of microalgae at the species level suggested a trade-off between nutrient use and grazing resistance which appeared to be linked to algal growth form. Compared to other growth forms, large erect species were most responsive to both nitrogen loading and herbivory. Grazing reduced microalgal diversity at low nutri- ent supply, but enhanced it at high nutrient supply. Herbivore effects varied considerably among dif- ferent sites and were stronger in spring than in summer. Manipulations of fish density during sum- mer did not have any effects on microalgal community structure. In conclusion, our results demonstrate that herbivores and nutrients have strong and balancing effects on marine microbenthic community structure.

Increased nutrient loading favors macroalgal blooms in eutrophied coastal ecosystems. The main counteracting factor on this bottom-up support is top-down control by consumers. We asked (1) whether herbivore control on 2 bloom-forming macroalgae in the Baltic Sea varies between different algal life stages, (2) whether herbivores selectively feed on Enteromorpha spp. (Chlorophyceae) thereby supporting dominance of Pilayella littoralis (Phaeophyceae), and (3) whether various herbivore species differ in their effects. In comparative field and laboratory experiments, we analyzed herbivore pressure and selectivity on germling density and adult thalli of Enteromorpha spp. and P. littoralis. In the field, herbivores reduced macroalgal recruitment by 80% within 14 d indicating strong herbivore control at early life stages. Recruits of Enteromorpha spp. were significantly preferred over P. littoralis Adult thalli of both algae showed similar growth rates, but grazing rates were significantly higher on Enteromorpha spp. In laboratory experiments, Idotea chelipes (Isopoda), Littorina saxatilis (Gastropoda) and Gammarus locusta (Amphipoda) strongly reduced germling density, whereas effects of L. littorea were weak. I. chelipes and L. saxatilis significantly preferred germlings of Enteromorpha spp. over P. littoralis. I. chelipes had strong effects on adult Enteromorpha spp. but not on P. littoralis. The effects of G. locusta and L. littorea on both adult algae were smaller and not selective, and L. saxatilis did not feed on adults at all. Different herbivore feeding modes are discussed. We conclude that strong and selective herbivory on juvenile and adult Enteromorpha spp. favors dominance of P. littoralis in the Baltic. An effective control of macroalgal blooms in eutrophied areas can be supported by sustaining high herbivore density and species richness considering the variable and complementary effects of different herbivore species on different algal life stages.

Although blooms of opportunistic fast-growing macroalgae now occur frequently in coastal ecosystems affected by eutrophication, their initiation and control is little understood. Most previous studies have focused on the ecophysiology of adult algae only. We show that spores and/or germlings may represent critical stages in the life cycles and mass-developments of co-occurring opportunistic macroalgae in the Baltic (Pilayella littoralis and Enteromorpha spp.). We investigated the overwintering of spores, timing of germination, subsequent growth, and grazing on spores and germlings, in order to explain the initiation of mass blooms and species dominance patterns. In the field, Enteromorpha spp. showed 10- to 50-fold higher abundances of overwintering microscopic forms (up to 330 individuals cm-2) than P. littoralis. Moreover, we found continuous production of spores (up to 1.2 million settling spores m-2 h-1) from April to October in Enteromorpha spp., while there was evidence of only a short reproductive period in Pilayella. However, in spring, germlings and adults of P. littoralis appeared earlier in the field and reached a 10-fold higher biomass than Enteromorpha spp. In factorial laboratory experiments including temperature and light, there were clear differences in timing of germination. P. littoralis germinated at 5°C whereas Enteromorpha spp. required temperatures of 10-15°C for germination. In contrast, we detected only minor differences in growth response among adults of P. littoralis and Enteromorpha spp. Germination, not growth of adults, appeared to be the ecophysiological bottleneck for initiating mass spring development. Following the spring Pilayella bloom, Enteromorpha germlings occurred massively in the field (April-September), but rarely developed into adults. In laboratory feeding experiments we tested whether crustacean mesograzers common in summer may control development of Enteromorpha germlings. Both germination of settled spores and growth of germlings were reduced by 93-99% in the presence of grazers (Idotea chelipes and Gammarus locusta). Thus in addition to ecophysiological constraints, grazers, if present, may play a decisive role in the early life stages of macroalgal mass developments. These results mirror patterns of overwintering of seeds, germination control, seed and seedling predation in terrestrial plant communities.

Fisheries exploitation has caused widespread declines in marine predators. Theory predicts that predator depletion will destabilise lower trophic levels, making natural communities more vulnerable to environmental perturbations. However, empirical evidence has been limited. Using a community matrix model, we empirically assessed trends in the stability of a multispecies coastal fish community over the course of predator depletion. Three indices of community stability (resistance, resilience and reactivity) revealed significantly decreasing stability concurrent with declining predator abundance. The trophically downgraded community exhibited weaker top-down control, leading to predator-release processes in lower trophic levels and increased susceptibility to perturbation. At the community level, our results suggest that high predator abundance acts as a stabilising force to the naturally stochastic and highly autocorrelated dynamics in low trophic species. These findings have important implications for the conservation and management of predators in marine ecosystems and provide empirical support for the theory of predatory control.

Sharks and rays’ abundance can decline considerably with fishing. Community changes, however, are more complex because of species interactions, and variable vulnerability and exposure to fishing. We evaluated long-term changes in the elasmobranch community of the Adriatic Sea, a heavily exploited Mediterranean basin where top-predators have been strongly depleted historically, and fishing developed unevenly between the western and eastern side. Combining and standardizing catch data from five trawl surveys from 1948– 2005, we estimated abundance trends and explained community changes using life histories, fish-market and effort data, and historical information.Weidentified a highly depleted elasmobranch community. Since 1948, catch rates have declined by .94% and 11 species ceased to be detected. The exploitation history and spatial gradients in fishing pressure explained most patterns in abundance and diversity, including the absence of strong compensatory increases. Ecological corridors and large-scale protected areas emerged as potential management options for elasmobranch recovery., JRC.G.4-Maritime affairs

Marine mammals have greatly benefitted from a shift from resource exploitation towards conservation. Often lauded as symbols of conservation success, some marine mammal populations have shown remarkable recoveries after severe depletions. Others have remained at low abundance levels, continued to decline, or become extinct or extirpated. Here we provide a quantitative assessment of (1) publicly available population-level abundance data for marine mammals worldwide, (2) abundance trends and recovery status, and (3) historic population decline and recent recovery. We compiled 182 population abundance time series for 47 species and identified major data gaps. In order to compare across the largest possible set of time series with varying data quality, quantity and frequency, we considered an increase in population abundance as evidence of recovery. Using robust log-linear regression over three generations, we were able to classify abundance trends for 92 spatially non-overlapping populations as Significantly Increasing (42%), Significantly Decreasing (10%), Non-Significant Change (28%) and Unknown (20%). Our results were comparable to IUCN classifications for equivalent species. Among different groupings, pinnipeds and other marine mammals (sirenians, polar bears and otters) showed the highest proportion of recovering populations, likely benefiting from relatively fast life histories and nearshore habitats that provided visibility and protective management measures. Recovery was less frequent among cetaceans, but more common in coastal than offshore populations. For marine mammals with available historical abundance estimates (n = 47), larger historical population declines were associated with low or variable recent recoveries so far. Overall, our results show that many formerly depleted marine mammal populations are recovering. However, data-deficient populations and those with decreasing and non-significant trends require attention. In particular, increased study of populations with major data gaps, including offshore small cetaceans, cryptic species, and marine mammals in low latitudes and developing nations, is needed to better understand the status of marine mammal populations worldwide.

In the coming century, life in the ocean will be confronted with a suite of environmental conditions that have no analog in human history. Thus, there is an urgent need to determine which marine species will adapt and which will go extinct. Here, we review the growing literature on marine extinctions and extinction risk in the fossil, historical, and modern records to compare the patterns, drivers, and biological correlates of marine extinctions at different times in the past. Characterized by markedly different environmental states, some past periods share common features with predicted future scenarios. We highlight how the different records can be integrated to better understand and predict the impact of current and projected future environmental changes on extinction risk in the ocean.

Many elasmobranchs have experienced strong population declines, which have been largely attributed to the direct and indirect effects of exploitation. Recently, however, live elasmobranchs are being increasingly valued for their role in marine ecosystems, dive tourism and intrinsic worth. Thus, management plans have been implemented to slow and ultimately reverse negative trends, including shark-specific (e.g. anti-finning laws) to ecosystem-based (e.g. no-take marine reserves) strategies. Yet it is unclear how successful these measures are, or will be, given the degree of depletion and slow recovery potential of most elasmobranchs. Here, current understanding of elasmobranch population recoveries is reviewed. The potential and realized extent of population increases, including rates of increase, timelines and drivers are evaluated. Across 40 increasing populations, only 25% were attributed to decreased anthropogenic mortality, while the majority was attributed to predation release. It is also shown that even low exploitation rates (2–6% per year) can halt or reverse positive population trends in six populations currently managed under recovery plans. Management measures that help restore elasmobranch populations include enforcement or near-zero fishing mortality, protection of critical habitats, monitoring and education. These measures are highlighted in a case study from the south-eastern U.S.A., where some evidence of recovery is seen in Pristis pectinata, Galeocerdo cuvier and Sphyrna lewini populations. It is concluded that recovery of elasmobranchs is certainly possible but requires time and a combination of strong and dedicated management actions to be successful.

Background Around the world, researchers are using the observations and experiences of citizens to describe patterns in animal populations. This data is often collected via ongoing sampling or by synthesizing past experiences. Since elasmobranchs are relatively rare, obtaining data for broad-scale trend analysis requires high sampling effort. Elasmobranchs are also relatively large and conspicuous and therefore it may be possible to enlist recreational divers to collect data on their occurrence and relative abundance from daily dive activities. For this, however, a good understanding of the value of data collected by recreational divers is essential. Methodology/Principal Findings Here, we explore the value of recreational divers for censusing elasmobranchs using a diverse set of data sources. First, we use a simulation experiment to explore detection rates of the roving diver technique, used by recreational divers, across a range of fish densities and speeds. Next, using a field survey, we show that inexperienced recreational divers detect and count elasmobranchs as well as experienced recreational divers. Finally, we use semi-structured interviews of recreational dive instructors to demonstrate the value of their recollections in terms of effort and their descriptions of spatial and temporal distributions of sharks in Thailand. Conclusions/Significance Overall, this study provides initial ground-work for using recreational divers for monitoring elasmobranch populations. If used appropriately, citizen-collected data may provide additional information that can be used to complement more standardized surveys and to describe population trends across a range of spatial and temporal scales. Due to the non-extractive nature of this data, recreational divers may also provide important insight into the success of conservation initiatives, such as shark sanctuaries and no-take zones.

The ocean has a history. Over the past hundreds, thousands, and millions of years the ocean and life within it have evolved and changed. This history determines the ocean’s present state and shapes its path into the future. We cannot understand how the ocean functions today without knowing its ecological history. Likewise, we cannot predict future changes without knowing the origin, cause, and trajectory of change in the past. Finally, we cannot effectively restore degraded marine populations, communities, or ecosystems without historical baselines to use as reference points.

25 pages, 8 figures, 6 tables electronic supplementary material in https://doi.org/10.1007/s10021-010-9404-8, The Mediterranean Sea has been strongly influenced by human activities for millennia. Although the environmental history of its surrounding terrestrial ecosystems has received considerable study, historical changes in its marine realm are less known. We used a multidisciplinary approach combining paleontological, archeological, historical, fisheries, and ecological data to reconstruct past changes in marine populations, habitats, and water quality in the Adriatic Sea. Then, we constructed binary food webs for different historical periods to analyze possible changes in food-web structure and functioning over time. Our results indicate that human activities have influenced marine resource abundance since at least Roman times and accelerated in the nineteenth and twentieth centuries. Today, 98% of traditional marine resources are depleted to less than 50% of former abundance, with large (>1 m) predators and consumers being most affected. With 37% of investigated species rare and 11% extirpated, diversity has shifted towards smaller, lower trophic-level species, further aggravated by more than 40 species invasions. Species providing habitat and filter functions have been reduced by 75%, contributing to the degradation of water quality and increased eutrophication. Increased exploitation and functional extinctions have altered and simplified food-web structure over time, especially by changing the proportions of top predators, intermediate consumers, and basal species. Moreover, simulations of species losses indicate that today's ecosystems may be less robust to species extinctions than in the past. Our results illustrate the long-term and far-reaching consequences human activities can have on marine food webs and ecosystems, Financial support was provided by the National Center for Ecological Analysis and Synthesis’ Long-term Marine Records working group, the Census of Marine Life’s FMAP project, and post-doctoral fellowships to M.C. from the Spanish Ministry of Science and Technology and the Marie-Curie International Outgoing Fellowships (IOF; Call: FP7-PEOPLE-2007-4-1-IOF) to ECOFUN

11 pages, 3 figures, 2 tables, Many marine populations and ecosystems have experienced strong historical depletions, yet reports of recoveries are increasing. Here, we review the growing research on marine recoveries to reveal how common recovery is, its magnitude, timescale and major drivers. Overall, 10–50% of depleted populations and ecosystems show some recovery, but rarely to former levels of abundance. In addition, recovery can take many decades for long-lived species and complex ecosystems. Major drivers of recovery include the reduction of human impacts, especially exploitation, habitat loss and pollution, combined with favorable life-history and environmental conditions. Awareness, legal protection and enforcement of management plans are also crucial. Learning from historical recovery successes and failures is essential for implementing realistic conservation goals and promising management strategies, Financial support was provided by the Sloan Foundation’s Census of Marine Life FMAP Program, the Natural Sciences and Engineering Research Council of Canada and Dalhousie University. MC was funded by the European Community Marie-Curie Post-doctoral Fellowship through the International Outgoing Fellowships (IOF; Call: FP7-PEOPLE-2007-4-1-IOF) to ECOFUN and LA by the European Commission through FP7.2009-1, Contract 244104–THESEUS

BACKGROUND: In recent decades, large pelagic and coastal shark populations have declined dramatically with increased fishing; however, the status of sharks in other systems such as coral reefs remains largely unassessed despite a long history of exploitation. Here we explore the contemporary distribution and sighting frequency of sharks on reefs in the greater-Caribbean and assess the possible role of human pressures on observed patterns. METHODOLOGY/PRINCIPAL FINDINGS: We analyzed 76,340 underwater surveys carried out by trained volunteer divers between 1993 and 2008. Surveys were grouped within one km2 cells, which allowed us to determine the contemporary geographical distribution and sighting frequency of sharks. Sighting frequency was calculated as the ratio of surveys with sharks to the total number of surveys in each cell. We compared sighting frequency to the number of people in the cell vicinity and used population viability analyses to assess the effects of exploitation on population trends. Sharks, with the exception of nurse sharks occurred mainly in areas with very low human population or strong fishing regulations and marine conservation. Population viability analysis suggests that exploitation alone could explain the large-scale absence; however, this pattern is likely to be exacerbated by additional anthropogenic stressors, such as pollution and habitat degradation, that also correlate with human population. CONCLUSIONS/SIGNIFICANCE: Human pressures in coastal zones have lead to the broad-scale absence of sharks on reefs in the greater-Caribbean. Preventing further loss of sharks requires urgent management measures to curb fishing mortality and to mitigate other anthropogenic stressors to protect sites where sharks still exist. The fact that sharks still occur in some densely populated areas where strong fishing regulations are in place indicates the possibility of success and encourages the implementation of conservation measures.

Whereas many land predators disappeared before their ecological roles were studied, the decline of marine apex predators is still unfolding. Large sharks in particular have experienced rapid declines over the last decades. In this study, we review the documented changes in exploited elasmobranch communities in coastal, demersal, and pelagic habitats, and synthesize the effects of sharks on their prey and wider communities. We show that the high natural diversity and abundance of sharks is vulnerable to even light fishing pressure. The decline of large predatory sharks reduces natural mortality in a range of prey, contributing to changes in abundance, distribution, and behaviour of small elasmobranchs, marine mammals, and sea turtles that have few other predators. Through direct predation and behavioural modifications, top-down effects of sharks have led to cascading changes in some coastal ecosystems. In demersal and pelagic communities, there is increasing evidence of mesopredator release, but cascading effects are more hypothetical. Here, fishing pressure on mesopredators may mask or even reverse some ecosystem effects. In conclusion, large sharks can exert strong top-down forces with the potential to shape marine communities over large spatial and temporal scales. Yet more empirical evidence is needed to test the generality of these effects throughout the ocean.

Estuaries are vital ecosystems that have sustained human and marine life since earliest times. Yet, no other part of the ocean has been so fundamentally shaped by human activities. Understanding the magnitude, drivers and consequences of past changes is essential to determine current trends and realistic management goals. This review provides a detailed account of human-induced changes in Massachusetts, Delaware, Chesapeake, Galveston and San Francisco Bays and Pamlico Sound. Native Americans have lived off these estuaries for millennia, yet left few signs of local resource depletion. European colonisation, commercialisation and industrialisation dramatically depleted and degraded valuable species, habitats and water quality. Exploitation and habitat loss were the main factors depleting 95% of valued species, with 35% being rare and 3% extirpated. Twentieth century conservation efforts enabled 10% of species to recover. Such profound changes in species diversity have altered the structure and functions of estuarine ecosystems as well as their services for human well-being. Thus, undesirable health risks and societal costs have increased over past decades. Protecting and restoring the diversity and vitality of estuaries will enhance their resilience towards current and future disturbances, yet require better governance of these often-neglected ecosystems. Their documented historical richness and essential role for marine life and people may increase the necessary awareness and appreciation.

Whereas many land predators disappeared before their ecological roles were studied, the decline of marine apex predators is still unfolding. Large sharks in particular have experienced rapid declines over the last decades. In this study, we review the documented changes in exploited elasmobranch communities in coastal, demersal, and pelagic habitats, and synthesize the effects of sharks on their prey and wider communities. We show that the high natural diversity and abundance of sharks is vulnerable to even light fishing pressure. The decline of large predatory sharks reduces natural mortality in a range of prey, contributing to changes in abundance, distribution, and behaviour of small elasmobranchs, marine mammals, and sea turtles that have few other predators. Through direct predation and behavioural modifications, top-down effects of sharks have led to cascading changes in some coastal ecosystems. In demersal and pelagic communities, there is increasing evidence of mesopredator release, but cascading effects are more hypothetical. Here, fishing pressure on mesopredators may mask or even reverse some ecosystem effects. In conclusion, large sharks can exert strong top-down forces with the potential to shape marine communities over large spatial and temporal scales. Yet more empirical evidence is needed to test the generality of these effects throughout the ocean.

Global patterns of species richness and their structuring forces have fascinated biologists since Darwin and provide critical context for contemporary studies in ecology, evolution and conservation. Anthropogenic impacts and the need for systematic conservation planning have further motivated the analysis of diversity patterns and processes at regional to global scales. Whereas land diversity patterns and their predictors are known for numerous taxa, our understanding of global marine diversity has been more limited, with recent findings revealing some striking contrasts to widely held terrestrial paradigms. Here we examine global patterns and predictors of species richness across 13 major species groups ranging from zooplankton to marine mammals. Two major patterns emerged: coastal species showed maximum diversity in the Western Pacific, whereas oceanic groups consistently peaked across broad mid-latitudinal bands in all oceans. Spatial regression analyses revealed sea surface temperature as the only environmental predictor highly related to diversity across all 13 taxa. Habitat availability and historical factors were also important for coastal species, whereas other predictors had less significance. Areas of high species richness were disproportionately concentrated in regions with medium or higher human impacts. Our findings indicate a fundamental role of temperature or kinetic energy in structuring cross-taxon marine biodiversity, and indicate that changes in ocean temperature, in conjunction with other human impacts, may ultimately rearrange the global distribution of life in the ocean.

Background Increasingly, underwater visual censuses (UVC) are used to assess fish populations. Several studies have demonstrated the effectiveness of protected areas for increasing fish abundance or provided insight into the natural abundance and structure of reef fish communities in remote areas. Recently, high apex predator densities (>100,000 individuals·km−2) and biomasses (>4 tonnes·ha−1) have been reported for some remote islands suggesting the occurrence of inverted trophic biomass pyramids. However, few studies have critically evaluated the methods used for sampling conspicuous and highly mobile fish such as sharks. Ideally, UVC are done instantaneously, however, researchers often count animals that enter the survey area after the survey has started, thus performing non-instantaneous UVC. Methodology/Principal Findings We developed a simulation model to evaluate counts obtained by divers deploying non-instantaneous belt-transect and stationary-point-count techniques. We assessed how fish speed and survey procedure (visibility, diver speed, survey time and dimensions) affect observed fish counts. Results indicate that the bias caused by fish speed alone is huge, while survey procedures had varying effects. Because the fastest fishes tend to be the largest, the bias would have significant implications on their biomass contribution. Therefore, caution is needed when describing abundance, biomass, and community structure based on non-instantaneous UVC, especially for highly mobile species such as sharks. Conclusions/Significance Based on our results, we urge that published literature state explicitly whether instantaneous counts were made and that survey procedures be accounted for when non-instantaneous counts are used. Using published density and biomass values of communities that include sharks we explore the effect of this bias and suggest that further investigation may be needed to determine pristine shark abundances and the existence of inverted biomass pyramids. Because such studies are used to make important management and conservation decisions, incorrect estimates of animal abundance and biomass have serious and significant implications.

Large animals are severely depleted in many ecosystems, yet we are only beginning to understand the ecological implications of their loss. To empirically measure the short-term effects of removing large animals from an ocean ecosystem, we used exclosures to remove large fish from a near-pristine coral reef at Palmyra Atoll, Central Pacific Ocean. We identified a range of effects that followed from the removal of these large fish. These effects were revealed within weeks of their removal. Removing large fish (1) altered the behavior of prey fish; (2) reduced rates of herbivory on certain species of reef algae; (3) had both direct positive (reduced mortality of coral recruits) and indirect negative (through reduced grazing pressure on competitive algae) impacts on recruiting corals; and (4) tended to decrease abundances of small mobile benthic invertebrates. Results of this kind help advance our understanding of the ecological importance of large animals in ecosystems. © 2010 The Author(s).

36 p. The Mediterranean Sea is a marine biodiversity hot spot. Here we combined an extensive literature analysis with expert opinions to update publicly available estimates of major taxa in this marine ecosystem and to revise and update several species lists. We also assessed overall spatial and temporal patterns of species diversity and identified major changes and threats. Our results listed approximately 17,000 marine species occurring in the Mediterranean Sea. However, our estimates of marine diversity are still incomplete as yet-undescribed species will be added in the future. Diversity for microbes is substantially underestimated, and the deep-sea areas and portions of the southern and eastern region are still poorly known. In addition, the invasion of alien species is a crucial factor that will continue to change the biodiversity of the Mediterranean, mainly in its eastern basin that can spread rapidly northwards and westwards due to the warming of the Mediterranean Sea. Spatial patterns showed a general decrease in biodiversity from northwestern to southeastern regions following a gradient of production, with some exceptions and caution due to gaps in our knowledge of the biota along the southern and eastern rims. Biodiversity was also generally higher in coastal areas and continental shelves, and decreases with depth. Temporal trends indicated that overexploitation and habitat loss have been the main human drivers of historical changes in biodiversity. At present, habitat loss and degradation, followed by fishing impacts, pollution, climate change, eutrophication, and the establishment of alien species are the most important threats and affect the greatest number of taxonomic groups. All these impacts are expected to grow in importance in the future, especially climate change and habitat degradation. The spatial identification of hot spots highlighted the ecological importance of most of the western Mediterranean shelves (and in particular, the Strait of Gibraltar and the adjacent Alboran Sea), western African coast, the Adriatic, and the Aegean Sea, which show high concentrations of endangered, threatened, or vulnerable species. The Levantine Basin, severely impacted by the invasion of species, is endangered as well. The authors gratefully acknowledge the support given by the European Census of Marine Life, the Total Foundation and the Alfred P. Sloan Foundation. MC was supported financially by the European Commission Marie Curie Postdoctoral Fellowship through the International Outgoing Fellowships (Call FP7-PEOPLE-2007-4-1-IOF) for the ECOFUN project and by Dalhousie University (Halifax, Canada). EB acknowledges financial support from GRACCIE project C5D2007-00067 funded by the Spanish Ministry of Science and Innovation. BG thanks the support of SESAME funded by the European Commission's Sixth Framework Program on Sustainable Development, Global Change and Ecosystems. JISS acknowledges ECOLIFE (CGL2008-05407-C03-03) funds of the Ministry of Science and Innovation of Spain. The present paper is a contribution of JG, DM and XT to the projects 2009SRG665 and 2009SGR484 of the “Generalitat de Catalunya”, and of DM and XT to project CTM2007-66635 funded by the Ministry of Science and Innovation of Spain. Research by C. N. Bianchi on Mediterranean Sea biodiversity is partly supported by the project “The impacts of biological invasions and climate change on the biodiversity of the Mediterranean Sea” (Italy-Israel co-operation) funded by the Italian Ministry of the Environment. The funders had no role in study design, data collection and analysis, decision to published, or preparation of the manuscript.

After a long history of overexploitation, increasing efforts to restore marine ecosystems and rebuild fisheries are under way. Here, we analyze current trends from a fisheries and conservation perspective. In 5 of 10 well-studied ecosystems, the average exploitation rate has recently declined and is now at or below the rate predicted to achieve maximum sustainable yield for seven systems. Yet 63% of assessed fish stocks worldwide still require rebuilding, and even lower exploitation rates are needed to reverse the collapse of vulnerable species. Combined fisheries and conservation objectives can be achieved by merging diverse management actions, including catch restrictions, gear modification, and closed areas, depending on local context. Impacts of international fleets and the lack of alternatives to fishing complicate prospects for rebuilding fisheries in many poorer regions, highlighting the need for a global perspective on rebuilding marine resources. Fil: Worm, Boris. Dalhousie University Halifax; Canadá Fil: Hilborn, Ray. University of Washington; Estados Unidos Fil: Baum, Julia K.. University of California at San Diego. Scripps Institution of Oceanography; Estados Unidos Fil: Branch, Trevor A.. University of Washington; Estados Unidos Fil: Collie, Jeremy S.. University of Rhode Island; Estados Unidos Fil: Costello, Christopher. University of California; Estados Unidos Fil: Fogarty, Michael J.. Noaa National Marine Fisheries Service Northeast Fisheries Science Center; Estados Unidos Fil: Fulton, Elizabeth A.. Commonwealth Scientific and Industrial Research Organisation; Australia Fil: Hutchings, Jeffrey A.. Dalhousie University Halifax; Canadá Fil: Jennings, Simon. University of East Anglia; Reino Unido. Centre for the Environment Fisheries and Aquaculture Science; Reino Unido Fil: Jensen, Olaf P.. University of Washington; Estados Unidos Fil: Lotze, Heike K.. Dalhousie University Halifax; Canadá Fil: Mace, Pamela M.. Ministry of Fisheries; Nueva Zelanda Fil: McClanahan, Tim R.. Wildlife Conservation Society Marine Programs; Kenia Fil: Minto, Cóilín. Dalhousie University Halifax; Canadá Fil: Palumbi, Stephen R.. University of Stanford; Estados Unidos Fil: Parma, Ana María. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Nacional Patagónico; Argentina Fil: Ricard, Daniel. Dalhousie University Halifax; Canadá Fil: Rosenberg, Andrew A.. University of New Hampshire. Institute for the Study of Earth, Oceans, and Space; Estados Unidos Fil: Watson, Reg. University of British Columbia; Canadá Fil: Zeller, Dirk. University of British Columbia; Canadá

Ambient temperature is very likely the most important environmental factor determining the distribution and diversity of life in the oceans. Hence, climate change is expected to alter marine biodiversity on a global scale. Here we review observed and predicted effects of climate change on the diversity of marine species. Overall, an increasing number of studies demonstrate that effects of climate change on marine biodiversity are already apparent from local to global scales. So far, long-term fish and plankton monitoring data have provided the most compelling evidence for climate-driven changes in species distribution and diversity, but studies involving other groups such as corals, seaweeds, and mammals are increasing. As a general pattern, tropical regions often experience a loss of species due to elevated heat stress, whereas temperate regions increase in diversity, as species migrate polewards. Net increases in diversity are also expected in the polar regions, but so far there are few observations to support this. Complex patterns of change can emerge where ocean warming is accompanied by the effects of sea level rise, acidification, habitat change, changes in ocean circulation, stratification, and other aspects of global change. From a management perspective, the conservation of biological diversity will provide insurance and resilience in the face of rapid global change. Cumulative impacts of exploitation, habitat destruction and other threats to biodiversity need to be minimized to maintain the adaptive capacity of marine ecosystems in the present and coming centuries. This might be particularly pressing in tropical regions and developing countries, which will face exceptional socio-economic and climate-related pressures, as well as in the polar regions, which are faced with a multitude of emerging pressures.

Current trends in marine ecosystems need to be interpreted against a solid understanding of the magnitude and drivers of past changes. Over the last decade, marine scientists from different disciplines have engaged in the emerging field of marine historical ecology to reconstruct past changes in the sea. Here we review the diversity of approaches used and resulting patterns of historical changes in large marine mammals, birds, reptiles and fish. Across 256 reviewed records, exploited populations declined 89% from historical abundance levels (range: 11–100%). In many cases, long-term fluctuations are related to climate variation, rapid declines to overexploitation and recent recoveries to conservation measures. These emerging historical patterns offer new insights into past ecosystems, and provide important context for contemporary ocean management.

Evidence for severe declines in large predatory fishes is increasing around the world. Because of its long history of intense fishing, the Mediterranean Sea offers a unique perspective on fish population declines over historical timescales. We used a diverse set of records dating back to the early 19th and mid 20th century to reconstruct long-term population trends of large predatory sharks in the northwestern Mediterranean Sea. We compiled 9 time series of abundance indices from commercial and recreational fishery landings, scientific surveys, and sighting records. Generalized linear models were used to extract instantaneous rates of change from each data set, and a meta-analysis was conducted to compare population trends. Only 5 of the 20 species we considered had sufficient records for analysis. Hammerhead (Sphyrna spp.), blue (Prionace glauca), mackerel (Isurus oxyrinchus and Lamna nasus), and thresher sharks (Alopias vulpinus) declined between 96 and 99.99% relative to their former abundance. According to World Conservation Union (IUCN) criteria, these species would be considered critically endangered. So far, the lack of quantitative population assessments has impeded shark conservation in the Mediterranean Sea. Our study fills this critical information gap, suggesting that current levels of exploitation put large sharks at risk of extinction in the Mediterranean Sea. Possible ecosystem effects of these losses involve a disruption of top-down control and a release of midlevel consumers.

Human-dominated marine ecosystems are experiencing accelerating loss of populations and species, with largely unknown consequences. We analyzed local experiments, long-term regional time series, and global fisheries data to test how biodiversity loss affects marine ecosystem services across temporal and spatial scales. Overall, rates of resource collapse increased and recovery potential, stability, and water quality decreased exponentially with declining diversity. Restoration of biodiversity, in contrast, increased productivity fourfold and decreased variability by 21%, on average. We conclude that marine biodiversity loss is increasingly impairing the ocean's capacity to provide food, maintain water quality, and recover from perturbations. Yet available data suggest that at this point, these trends are still reversible.

Estuarine and coastal transformation is as old as civilization yet has dramatically accelerated over the past 150 to 300 years. Reconstructed time lines, causes, and consequences of change in 12 once diverse and productive estuaries and coastal seas worldwide show similar patterns: Human impacts have depleted >90% of formerly important species, destroyed >65% of seagrass and wetland habitat, degraded water quality, and accelerated species invasions. Twentieth-century conservation efforts achieved partial recovery of upper trophic levels but have so far failed to restore former ecosystem structure and function. Our results provide detailed historical baselines and quantitative targets for ecosystem-based management and marine conservation.

Eutrophication can profoundly change rocky shore communities. These changes often cause the replacement of perennial, canopy-forming algae such as Fucus spp. with annual, bloom-forming algae such as Enteromorpha spp. Grazing, however, can counteract eutrophication by eliminating the annual algae’s susceptible recruits. We examine these generalizations across large scales. We use replicated ‘‘bioassay’’ experiments to compare the effects of eutrophication and grazing across four paired control versus eutrophied sites in the Northwest Atlantic and four eutrophied sites in the Baltic Sea in spring and summer. At each site, annual algal recruitment and grazing pressure were estimated using tiles seeded with Enteromorpha intestinalispropagules. Tiles were exposed for 3 weeks with grazers excluded or allowed access. Productivity of E. intestinalis recruits was strongly related to eutrophication (10-fold increase) and grazing (80% decrease) and was weakly related to season. While the absolute grazing rate increased in a linear fashion with algal productivity, the relative grazing rate remained surprisingly constant ( ;80%). Comparative field surveys showed that perennial algae decreased by 30‐60%, while annual algae, filter feeders, and grazers increased across a gradient of eutrophication. As eutrophication increased from control to eutrophied to point source sites, rocky shore communities became increasingly dominated by single species of annual algae or filter feeders, and community diversity declined consistently by 24‐46%. We conclude that grazers are important controllers of algal blooms but that, ultimately, they cannot override the effects of increasing eutrophication on rocky shore community structure and biodiversity. Rocky shores are among the most dynamic and productive ecosystems on the planet. Biomass and primary productivity are typically dominated by canopy-forming perennial macroalgae such as fucoids and laminarians. Together with seagrasses on soft-bottom habitats, these algae generate up to 40% of the primary productivity of the coastal zone (Char

Humans have interacted with the Wadden Sea since its origin 7,500 years ago. However, exploitation, habitat alteration and pollution have strongly increased since the Middle Ages, affecting abundance and distribution of many marine mammals, birds, fish, invertebrates and plants. Large whales and some large birds disappeared more than 500 years ago. Most small whales, seals, birds, large fish and oysters were severely reduced by the late 19th and early 20th centuries, leading to the collapse of several traditional fisheries. In the 20th century, conservation efforts have enabled some breeding birds and seals to recover. But other species declined further due to continuing exploitation, habitat destruction, pollution and eutrophication. Moreover, complex three-dimensional habitats such as oyster banks, Sabellaria reefs and subtidal eelgrass beds have been lost completely. In contrast, several opportunistic species such as gulls, polychaetes, green algae and exotic invaders increased during the 20th century. Taken together, multiple human impacts have caused dramatic losses of large predators and habitat-building species in the Wadden Sea over the last 500 years. Although still of high natural value and global importance, the Wadden Sea is a fundamentally changed ecosystem. On the other hand, reduced hunting pressure, increased habitat protection and reduced river pollution have enabled the recent recovery of several species and an increase in environmental quality. These successes, together with a historical vision of what was once possible, should guide current and future conservation, restoration and management efforts towards a more sustainable interaction between man and the sea.

Concentrations of biodiversity, or hotspots, represent conservation priorities in terrestrial ecosystems but remain largely unexplored in marine habitats. In the open ocean, many large predators such as tunas, sharks, billfishes, and sea turtles are of current conservation concern because of their vulnerability to overfishing and ecosystem role. Here we use scientific-observer records from pelagic longline fisheries in the Atlantic and Pacific Oceans to show that oceanic predators concentrate in distinct diversity hotspots. Predator diversity consistently peaks at intermediate latitudes (20–30° N and S), where tropical and temperate species ranges overlap. Individual hotspots are found close to prominent habitat features such as reefs, shelf breaks, or seamounts and often coincide with zooplankton and coral reef hotspots. Closed-area models in the northwest Atlantic predict that protection of hotspots outperforms other area closures in safeguarding threatened pelagic predators from ecological extinction. We conclude that the seemingly monotonous landscape of the open ocean shows rich structure in species diversity and that these features should be used to focus future conservation efforts.

Diverse coastal seaweed communities dominated by perennial fucoids become replaced by species-poor turfs of annual algae throughout the Baltic Sea. A large-scale field survey and factorial field experiments indicated that grazers maintain the fucoid community through selective consumption of annual algae. Interactive effects between grazers and dormant propagules of annual algae, stored in a 'marine seed bank', determine the response of this system to anthropogenic nutrient loading. Nutrients override grazer control and accelerate the loss of algal diversity in the presence but not in the absence of a propagule bank. This implies a novel role of propagule banks for community regulation and ecosystem response to marine eutrophication.

13 pages, 8 figures, 4 tables, Seagrass beds provide important habitat for a wide range of marine species but are threatened by multiple human impacts in coastal waters. Although seagrass communities have been well-studied in the field, a quantification of their food-web structure and functioning, and how these change across space and human impacts has been lacking. Motivated by extensive field surveys and literature information, we analyzed the structural features of food webs associated with Zostera marina across 16 study sites in 3 provinces in Atlantic Canada. Our goals were to (i) quantify differences in food-web structure across local and regional scales and human impacts, (ii) assess the robustness of seagrass webs to simulated species loss, and (iii) compare food-web structure in temperate Atlantic seagrass beds with those of other aquatic ecosystems. We constructed individual food webs for each study site and cumulative webs for each province and the entire region based on presence/absence of species, and calculated 16 structural properties for each web. Our results indicate that food-web structure was similar among low impact sites across regions. With increasing human impacts associated with eutrophication, however, food-web structure show evidence of degradation as indicated by fewer trophic groups, lower maximum trophic level of the highest top predator, fewer trophic links connecting top to basal species, higher fractions of herbivores and intermediate consumers, and higher number of prey per species. These structural changes translate into functional changes with impacted sites being less robust to simulated species loss. Temperate Atlantic seagrass webs are similar to a tropical seagrass web, yet differed from other aquatic webs, suggesting consistent food-web characteristics across seagrass ecosystems in different regions. Our study illustrates that food-web structure and functioning of seagrass habitats change with human impacts and that the spatial scale of food-web analysis is critical for determining results, This work was funded by an NSERC Discovery grant to HKL. AS was supported by NSERC PGSD and Killam Trusts Predoctoral Scholarships, and MC by a postdoctoral fellowship from the Spanish Ministry of Science and Technology, and the European Community Marie-Curie Program through the International Outgoing Fellowships (Call: FP7-PEOPLE-2007-4-1-IOF) to the ECOFUN project. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript

BackgroundWorldwide, finfish fisheries are receiving increasing assessment and regulation, slowly leading to more sustainable exploitation and rebuilding. In their wake, invertebrate fisheries are rapidly expanding with little scientific scrutiny despite increasing socio-economic importance.Methods and findingsWe provide the first global evaluation of the trends, drivers, and population and ecosystem consequences of invertebrate fisheries based on a global catch database in combination with taxa-specific reviews. We also develop new methodologies to quantify temporal and spatial trends in resource status and fishery development. Since 1950, global invertebrate catches have increased 6-fold with 1.5 times more countries fishing and double the taxa reported. By 2004, 34% of invertebrate fisheries were over-exploited, collapsed, or closed. New fisheries have developed increasingly rapidly, with a decrease of 6 years (3 years) in time to peak from the 1950s to 1990s. Moreover, some fisheries have expanded further and further away from their driving market, encompassing a global fishery by the 1990s. 71% of taxa (53% of catches) are harvested with habitat-destructive gear, and many provide important ecosystem functions including habitat, filtration, and grazing.ConclusionsOur findings suggest that invertebrate species, which form an important component of the basis of marine food webs, are increasingly exploited with limited stock and ecosystem-impact assessments, and enhanced management attention is needed to avoid negative consequences for ocean ecosystems and human well-being.

We show that globally declining fisheries catch trends cannot be explained by random processes and are consistent with declining stock abundance trends. Future projections are inherently uncertain but may provide a benchmark against which to assess the effectiveness of conservation measures. Marine reserves and fisheries closures are among those measures and can be equally effective in tropical and temperate areas—but must be combined with catch-, effort-, and gear restrictions to meet global conservation objectives.