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Although Southeast Asia is a hotspot of global seagrass diversity, there are considerable information gaps in the distribution of seagrass beds. Broad-scale seagrass distribution has not been updated in the global seagrass database by UNEP-WCMC since 2000, although studies on seagrasses have been undertaken intensively in each region. Here we analyze the recent distribution of tropical seagrass beds, their temporal changes, causes of decline and conservation status in Southeast Asia (plus southern mainland China, Taiwan and Ryukyu Island of Japan) using data collected after 2000. Based on the 195 literature published since 2000, we identified 1,259 point data and 1,461 polygon data showing the distribution of seagrass beds. A large discrepancy was found in the seagrass bed distribution between our updated data and the UNEP-WCMC database, mostly due to inaccurate and low resolution location information in the latter. Temporal changes in seagrass bed area analyzed for 68 sites in nine countries/regions demonstrated that more than 60% of seagrass beds declined at an average rate of 10.9% year–1, whereas 20% of beds increased at an average rate of 8.1% year–1, leading to an overall average decline of 4.7% year–1. Various types of human-induced threats were reported as causes for the decline, including coastal development, fisheries/aquaculture, and natural factors such as typhoons and tsunamis. The percentage of seagrass beds covered with existing marine protected areas (MPAs) varied greatly among countries/regions, from less than 1% in Brunei Darussalam and Singapore to 100% in southern Japan. However, the degree of conservation regulation was not sufficient even in regions with higher MPA coverage. The percentage of seagrass beds within EBSAs (Ecologically and Biologically Significant Area determined by the Convention of Biological Diversity) was higher than that within MPAs becau

In coastal waters around the world, the dominant primary producers are benthic macrophytes, including seagrasses and macroalgae, that provide habitat structure and food for diverse and abundant biological communities and drive ecosystem processes. Seagrass meadows and macroalgal forests play key roles for coastal societies, contributing to fishery yields, storm protection, biogeochemical cycling and storage, and important cultural values. These socio-economically valuable services are threatened worldwide by human activities, with substantial areas of seagrass and macroalgal forests lost over the last half-century. Tracking the status and trends in marine macrophyte cover and quality is an emerging priority for ocean and coastal management, but doing so has been challenged by limited coordination across the numerous efforts to monitor macrophytes, which vary widely in goals, methodologies, scales, capacity, governance approaches, and data availability. Here, we present a consensus assessment and recommendations on the current state of and opportunities for advancing global marine macrophyte observations, integrating contributions from a community of researchers with broad geographic and disciplinary expertise. With the increasing scale of human impacts, the time is ripe to harmonize marine macrophyte observations by building on existing networks and identifying a core set of common metrics and approaches in sampling design, field measurements, governance, capacity building, and data management. We recommend a tiered observation system, with improvement of remote sensing and remote underwater imaging to expand capacity to capture broad-scale extent at intervals of several years, coordinated with stratified in situ sampling annually to characterize the key variables of cover and taxonomic or functional group composition, and to provide ground-truth. A robust networked system of macrophyte observations will be facilitated by establishing best practices, including stand

In coastal waters around the world, the dominant primary producers are benthic macrophytes, including seagrasses and macroalgae, that provide habitat structure and food for diverse and abundant biological communities and drive ecosystem processes. Seagrass meadows and macroalgal forests play key roles for coastal societies, contributing to fishery yields, storm protection, biogeochemical cycling and storage, and important cultural values. These socio-economically valuable services are threatened worldwide by human activities, with substantial areas of seagrass and macroalgal forests lost over the last half-century. Tracking the status and trends in marine macrophyte cover and quality is an emerging priority for ocean and coastal management, but doing so has been challenged by limited coordination across the numerous efforts to monitor macrophytes, which vary widely in goals, methodologies, scales, capacity, governance approaches, and data availability. Here, we present a consensus assessment and recommendations on the current state of and opportunities for advancing global marine macrophyte observations, integrating contributions from a community of researchers with broad geographic and disciplinary expertise. With the increasing scale of human impacts, the time is ripe to harmonize marine macrophyte observations by building on existing networks and identifying a core set of common metrics and approaches in sampling design, field measurements, governance, capacity building, and data management. We recommend a tiered observation system, with improvement of remote sensing and remote underwater imaging to expand capacity to capture broad-scale extent at intervals of several years, coordinated with stratified in situ sampling annually to characterize the key variables of cover and taxonomic or functional group composition, and to provide ground-truth. A robust networked system of macrophyte observations will be facilitated by establishing best practices, including stand

Despite the importance of coastal ecosystems for the global carbon budgets, knowledge of their carbon storage capacity and the factors driving variability in storage capacity is still limited. Here we provide an estimate on the magnitude and variability of carbon stocks within a widely distributed marine foundation species throughout its distribution area in temperate Northern Hemisphere. We sampled 54 eelgrass (Zostera marina) meadows, spread across eight ocean margins and 36° of latitude, to determine abiotic and biotic factors influencing organic carbon (Corg) stocks in Zostera marina sediments. The Corg stocks (integrated over 25-cm depth) showed a large variability and ranged from 318 to 26,523 g C/m2 with an average of 2,721 g C/m2. The projected Corg stocks obtained by extrapolating over the top 1 m of sediment ranged between 23.1 and 351.7 Mg C/ha, which is in line with estimates for other seagrasses and other blue carbon ecosystems. Most of the variation in Corg stocks was explained by five environmental variables (sediment mud content, dry density and degree of sorting, and salinity and water depth), while plant attributes such as biomass and shoot density were less important to Corg stocks. Carbon isotopic signatures indicated that at most sites <50% of the sediment carbon is derived from seagrass, which is lower than reported previously for seagrass meadows. The high spatial carbon storage variability urges caution in extrapolating carbon storage capacity between geographical areas as well as within and between seagrass species.

Despite the importance of coastal ecosystems for the global carbon budgets, knowledge of their carbon storage capacity and the factors driving variability in storage capacity is still limited. Here we provide an estimate on the magnitude and variability of carbon stocks within a widely distributed marine foundation species throughout its distribution area in temperate Northern Hemisphere. We sampled 54 eelgrass (Zostera marina) meadows, spread across eight ocean margins and 36° of latitude, to determine abiotic and biotic factors influencing organic carbon (Corg) stocks in Zostera marina sediments. The Corg stocks (integrated over 25-cm depth) showed a large variability and ranged from 318 to 26,523 g C/m2 with an average of 2,721 g C/m2. The projected Corg stocks obtained by extrapolating over the top 1 m of sediment ranged between 23.1 and 351.7 Mg C/ha, which is in line with estimates for other seagrasses and other blue carbon ecosystems. Most of the variation in Corg stocks was explained by five environmental variables (sediment mud content, dry density and degree of sorting, and salinity and water depth), while plant attributes such as biomass and shoot density were less important to Corg stocks. Carbon isotopic signatures indicated that at most sites <50% of the sediment carbon is derived from seagrass, which is lower than reported previously for seagrass meadows. The high spatial carbon storage variability urges caution in extrapolating carbon storage capacity between geographical areas as well as within and between seagrass species.

Rapid evolution of non-native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction and magnitude of phenotypic evolution during invasion has been underestimated. We explored the utility of niche-shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe and northwestern Africa within the last 100 years. A genetically-informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non-native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non-native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field-collected and 325 common-garden thalli from 40 locations and as predicted, non-native populations had greater tolerance for ecologically-relevant extreme heat (40°C) than did Japanese source populations. Non-native populations also had greater tolerance for cold and low-salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: populations from warmer, lower-latitude estuaries had greater heat tolerance than did populations from colder, higher-latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non-native marine species. Genetically-informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions, and may help

Rapid evolution of non-native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction and magnitude of phenotypic evolution during invasion has been underestimated. We explored the utility of niche-shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe and northwestern Africa within the last 100 years. A genetically-informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non-native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non-native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field-collected and 325 common-garden thalli from 40 locations and as predicted, non-native populations had greater tolerance for ecologically-relevant extreme heat (40°C) than did Japanese source populations. Non-native populations also had greater tolerance for cold and low-salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: populations from warmer, lower-latitude estuaries had greater heat tolerance than did populations from colder, higher-latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non-native marine species. Genetically-informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions, and may help

Rapid evolution of non-native species can facilitate invasion success, but recent reviews indicate that such microevolution rarely yields expansion of the climatic niche in the introduced habitats. However, because some invasions originate from a geographically restricted portion of the native species range and its climatic niche, it is possible that the frequency, direction and magnitude of phenotypic evolution during invasion has been underestimated. We explored the utility of niche-shift analyses in the red seaweed Gracilaria vermiculophylla, which expanded its range from the northeastern coastline of Japan to North America, Europe and northwestern Africa within the last 100 years. A genetically-informed climatic niche shift analysis indicates that native source populations occur in colder and highly seasonal habitats, while most non-native populations typically occur in warmer, less seasonal habitats. This climatic niche expansion predicts that non-native populations evolved greater tolerance for elevated heat conditions relative to native source populations. We assayed 935 field-collected and 325 common-garden thalli from 40 locations and as predicted, non-native populations had greater tolerance for ecologically-relevant extreme heat (40°C) than did Japanese source populations. Non-native populations also had greater tolerance for cold and low-salinity stresses relative to source populations. The importance of local adaptation to warm temperatures during invasion was reinforced by evolution of parallel clines: populations from warmer, lower-latitude estuaries had greater heat tolerance than did populations from colder, higher-latitude estuaries in both Japan and eastern North America. We conclude that rapid evolution plays an important role in facilitating the invasion success of this and perhaps other non-native marine species. Genetically-informed ecological niche analyses readily generate clear predictions of phenotypic shifts during invasions, and may help

The susceptibility of native and non-native populations of the red alga Gracilaria vermiculophylla to fouling was compared in common garden experiments. Native and non-native algae were enclosed into dialysis membrane tubes, and the tubes were exposed to natural fouling. Fouling on the outside of the tubes was mediated by chemical compounds excreted by G. vermiculophylla that diffused through the membranes. Fouling pressure was significantly higher in the Kiel Fjord (non-native range) than in Akkeshi Bay (native range), but, at both sites, tubes containing non-native G. vermiculophylla were less fouled than those with native conspecifics. This is the first in situ evidence that susceptibility to fouling differs between native and non-native populations of an aquatic organism. The technique of enclosing organisms into dialysis tubes represents a simple, efficient and accurate way to test chemical antifouling defenses and could possibly be applied to other organisms.

Fouling is a stressor that might determine the fate of seaweeds, but reports of algal adaptation to epibiosis are scarce. Previous comparisons have shown resistance to epibionts can be higher in non-native than in resident seaweed species, but we do not know whether it is an intrinsic trait of the non-natives or it has been acquired during the invasion process. We here compared native and non-native populations of the same algal species to elucidate this question. Resistance against two groups of epiphytes was assessed in living thalli and in artificial substrata coated with surface extracts, both gained from four Asian (native) and four European (non-native) populations of the red alga Gracilaria vermiculophylla. Two diatom species and two filamentous macroalgae were used as micro- and macro-epiphytes, and one of each type was collected in Asia, while the other came from Europe. Laboratory assays were done in both distributional ranges of G. vermiculophylla and in different seasons. We used a fully crossed design with the factors (i) ‘Origin of Gracilaria’, (ii) ‘Origin of epiphytes’, (iii) ‘Season’ and (iv) ‘Solvent used for extraction’. Both groups of epiphytes, regardless of their origin, attached less to living thalli and to surface extracts from non-native G. vermiculophylla. Fewer diatoms attached to hexane-based extracts, while fewer Ceramium filaments settled on extracts gained with dichloromethane. Our results show for the first time that non-native individuals of a seaweed are better defended against epiphytes than native conspecifics. Furthermore, we found evidence that at least a part of the defence is based on extractable secondary metabolites. We suggest that an enhanced defence against epiphytes after introduction is one reason for G. vermiculophylla’s invasion success. Our observation may also apply to other basibiont–epibiont interactions and could be a key feature of seaweed bioinvasions.

Fouling is a stressor that might determine the fate of seaweeds, but reports of algal adaptation to epibiosis are scarce. Previous comparisons have shown resistance to epibionts can be higher in non-native than in resident seaweed species, but we do not know whether it is an intrinsic trait of the non-natives or it has been acquired during the invasion process. We here compared native and non-native populations of the same algal species to elucidate this question. Resistance against two groups of epiphytes was assessed in living thalli and in artificial substrata coated with surface extracts, both gained from four Asian (native) and four European (non-native) populations of the red alga Gracilaria vermiculophylla. Two diatom species and two filamentous macroalgae were used as micro- and macro-epiphytes, and one of each type was collected in Asia, while the other came from Europe. Laboratory assays were done in both distributional ranges of G. vermiculophylla and in different seasons. We used a fully crossed design with the factors (i) ‘Origin of Gracilaria’, (ii) ‘Origin of epiphytes’, (iii) ‘Season’ and (iv) ‘Solvent used for extraction’. Both groups of epiphytes, regardless of their origin, attached less to living thalli and to surface extracts from non-native G. vermiculophylla. Fewer diatoms attached to hexane-based extracts, while fewer Ceramium filaments settled on extracts gained with dichloromethane. Our results show for the first time that non-native individuals of a seaweed are better defended against epiphytes than native conspecifics. Furthermore, we found evidence that at least a part of the defence is based on extractable secondary metabolites. We suggest that an enhanced defence against epiphytes after introduction is one reason for G. vermiculophylla’s invasion success. Our observation may also apply to other basibiont–epibiont interactions and could be a key feature of seaweed bioinvasions.

The susceptibility of native and non-native populations of the red alga Gracilaria vermiculophylla to fouling was compared in common garden experiments. Native and non-native algae were enclosed into dialysis membrane tubes, and the tubes were exposed to natural fouling. Fouling on the outside of the tubes was mediated by chemical compounds excreted by G. vermiculophylla that diffused through the membranes. Fouling pressure was significantly higher in the Kiel Fjord (non-native range) than in Akkeshi Bay (native range), but, at both sites, tubes containing non-native G. vermiculophylla were less fouled than those with native conspecifics. This is the first in situ evidence that susceptibility to fouling differs between native and non-native populations of an aquatic organism. The technique of enclosing organisms into dialysis tubes represents a simple, efficient and accurate way to test chemical antifouling defenses and could possibly be applied to other organisms.

Contains fulltext : 161998.pdf (publisher's version ) (Closed access)

Contains fulltext : 161998.pdf (publisher's version ) (Closed access)

Contains fulltext : 161998.pdf (publisher's version ) (Closed access)

In coastal and estuarine systems, foundation species like seagrasses, mangroves, saltmarshes or corals provide important ecosystem services. Seagrasses are globally declining and their reintroduction has been shown to restore ecosystem functions. However, seagrass restoration is often challenging, given the dynamic and stressful environment that seagrasses often grow in. From our world-wide meta-analysis of seagrass restoration trials (1786 trials), we describe general features and best practice for seagrass restoration. We confirm that removal of threats is important prior to replanting. Reduced water quality (mainly eutrophication), and construction activities led to poorer restoration success than, for instance, dredging, local direct impact and natural causes. Proximity to and recovery of donor beds were positively correlated with trial performance. Planting techniques can influence restoration success. The meta-analysis shows that both trial survival and seagrass population growth rate in trials that survived are positively affected by the number of plants or seeds initially transplanted. This relationship between restoration scale and restoration success was not related to trial characteristics of the initial restoration. The majority of the seagrass restoration trials have been very small, which may explain the low overall trial survival rate (i.e. estimated 37%). Successful regrowth of the foundation seagrass species appears to require crossing a minimum threshold of reintroduced individuals. Our study provides the first global field evidence for the requirement of a critical mass for recovery, which may also hold for other foundation species showing strong positive feedback to a dynamic environment. Synthesis and applications. For effective restoration of seagrass foundation species in its typically dynamic, stressful environment, introduction of large numbers is seen to be beneficial and probably serves two purposes. First, a large-scale planting increases

In coastal and estuarine systems, foundation species like seagrasses, mangroves, saltmarshes or corals provide important ecosystem services. Seagrasses are globally declining and their reintroduction has been shown to restore ecosystem functions. However, seagrass restoration is often challenging, given the dynamic and stressful environment that seagrasses often grow in. From our world-wide meta-analysis of seagrass restoration trials (1786 trials), we describe general features and best practice for seagrass restoration. We confirm that removal of threats is important prior to replanting. Reduced water quality (mainly eutrophication), and construction activities led to poorer restoration success than, for instance, dredging, local direct impact and natural causes. Proximity to and recovery of donor beds were positively correlated with trial performance. Planting techniques can influence restoration success. The meta-analysis shows that both trial survival and seagrass population growth rate in trials that survived are positively affected by the number of plants or seeds initially transplanted. This relationship between restoration scale and restoration success was not related to trial characteristics of the initial restoration. The majority of the seagrass restoration trials have been very small, which may explain the low overall trial survival rate (i.e. estimated 37%). Successful regrowth of the foundation seagrass species appears to require crossing a minimum threshold of reintroduced individuals. Our study provides the first global field evidence for the requirement of a critical mass for recovery, which may also hold for other foundation species showing strong positive feedback to a dynamic environment. Synthesis and applications. For effective restoration of seagrass foundation species in its typically dynamic, stressful environment, introduction of large numbers is seen to be beneficial and probably serves two purposes. First, a large-scale planting increases

Among the numerous anti-herbivore defences developed by macroalgae, chemical and morphological traits are best documented and understood. Plant defence theory suggests that these resistances, which can either be constitutive or inducible, are associated with metabolic costs. They should therefore be impaired under conditions of energy limitation, but evidence for this prediction is scarce. In two subsequent experiments, we tested whether a reduction of light availability is changing feeding rates of the two mesoherbivores Idotea ochotensis and Lacuna smithii on the red alga Chondrus yendoi. Algal individuals were kept in outdoor mesocosm facilities for 10days, during which we manipulated the amount of incoming sunlight at 6 levels (0%99% reduction, i.e. 200020 mols1m2). Orthogonal to this, we established the presence or absence of one of the herbivores to test whether C. yendoi can generate a defence. Algal palatability was investigated afterwards in no-choice feeding assays using na ve grazer individuals. The consumption of algal tissue in L. smithii increased with decreasing light, while this was not the case for I. ochotensis. However, we found a defence induced as a reaction to herbivory only by the highly mobile isopod but not when the slowly moving snail grazed on C. yendoi. Isopod total consumption rates in our experiments were 40 times higher than those of the gastropods. We therefore suggest that C. yendoi exhibits grazer-specific reactions to herbivory, depending on the mobility and voracity of the consumers. Interestingly, only for one of the grazers, i.e. the snail, short-term light reduction influenced the palatability of algal tissue. We discuss different but not mutually exclusive models that could explain this pattern. In conclusion, we view this three-species system as an illustrative example for specificity in grazeralgal interactions and their modification by environmental stress.

Among the numerous anti-herbivore defences developed by macroalgae, chemical and morphological traits are best documented and understood. Plant defence theory suggests that these resistances, which can either be constitutive or inducible, are associated with metabolic costs. They should therefore be impaired under conditions of energy limitation, but evidence for this prediction is scarce. In two subsequent experiments, we tested whether a reduction of light availability is changing feeding rates of the two mesoherbivores Idotea ochotensis and Lacuna smithii on the red alga Chondrus yendoi. Algal individuals were kept in outdoor mesocosm facilities for 10days, during which we manipulated the amount of incoming sunlight at 6 levels (0%99% reduction, i.e. 200020 mols1m2). Orthogonal to this, we established the presence or absence of one of the herbivores to test whether C. yendoi can generate a defence. Algal palatability was investigated afterwards in no-choice feeding assays using na ve grazer individuals. The consumption of algal tissue in L. smithii increased with decreasing light, while this was not the case for I. ochotensis. However, we found a defence induced as a reaction to herbivory only by the highly mobile isopod but not when the slowly moving snail grazed on C. yendoi. Isopod total consumption rates in our experiments were 40 times higher than those of the gastropods. We therefore suggest that C. yendoi exhibits grazer-specific reactions to herbivory, depending on the mobility and voracity of the consumers. Interestingly, only for one of the grazers, i.e. the snail, short-term light reduction influenced the palatability of algal tissue. We discuss different but not mutually exclusive models that could explain this pattern. In conclusion, we view this three-species system as an illustrative example for specificity in grazeralgal interactions and their modification by environmental stress.

Factors and processes affecting community structures operate at various spatial and temporal scales. We analyzed how similarities of rocky intertidal assemblages vary at different spatial scales using a nested, hierarchical sampling design. Intertidal assemblages consisting of algae, sessile animals, and mobile animals were censused on five rock walls at each of five shores chosen within each of six regions along the Pacific coast of Japan, encompassing 1,800 km of coastlines. Based on this sampling design, similarities in assemblages were calculated using both qualitative (presence/absence) and quantitative (percent cover and density) data, and compared at three spatial levels: (1) rock level (the finest spatial scale, encompassing several to hundreds of meters), (2) shore level (the intermediate spatial scale, encompassing several to tens of kilometers), and (3) region level (the broadest spatial scale, encompassing hundreds to thousands of kilometers). Cluster analysis showed that assemblages were categorized into distinct regional groups except for the two southern regions, but they did not separate clearly from each shore. A nested analysis of similarities revealed significant variation in similarities among regions and among shores within each region, with the former showing greater variation. Similarity was negatively correlated with geographic distance at the regional level but not at the shore or the rock levels. At the regional level, similarity decreased more rapidly with distance for mobile animals than sessile organisms. The analyses highlighted the importance of broad-scale abiotic/biotic factors such as oceanic current conditions and biogeographic factors in determining observed patterns in similarity of rocky intertidal assemblages.

Factors and processes affecting community structures operate at various spatial and temporal scales. We analyzed how similarities of rocky intertidal assemblages vary at different spatial scales using a nested, hierarchical sampling design. Intertidal assemblages consisting of algae, sessile animals, and mobile animals were censused on five rock walls at each of five shores chosen within each of six regions along the Pacific coast of Japan, encompassing 1,800 km of coastlines. Based on this sampling design, similarities in assemblages were calculated using both qualitative (presence/absence) and quantitative (percent cover and density) data, and compared at three spatial levels: (1) rock level (the finest spatial scale, encompassing several to hundreds of meters), (2) shore level (the intermediate spatial scale, encompassing several to tens of kilometers), and (3) region level (the broadest spatial scale, encompassing hundreds to thousands of kilometers). Cluster analysis showed that assemblages were categorized into distinct regional groups except for the two southern regions, but they did not separate clearly from each shore. A nested analysis of similarities revealed significant variation in similarities among regions and among shores within each region, with the former showing greater variation. Similarity was negatively correlated with geographic distance at the regional level but not at the shore or the rock levels. At the regional level, similarity decreased more rapidly with distance for mobile animals than sessile organisms. The analyses highlighted the importance of broad-scale abiotic/biotic factors such as oceanic current conditions and biogeographic factors in determining observed patterns in similarity of rocky intertidal assemblages.