Masahiro Nakaoka

Nonlethal effects of predators on prey populations : predator-mediated change in bivalve growth

Predators affect dynamics of prey populations not only by decreasing their survivorship, but also by changing other vital rates such as growth and reproduction. However, such nonlethal effects have rarely been examined experimentally in the field. In North Carolina estuaries, the hard clam Mercenaria mercenaria generally shows lower growth rates in habitats where predation rates are higher. Predator avoidance behavior of the clams (feeding inhibition) might be one of the factors responsible for this correlation. To examine possible effects of predators on clam growth, I first compared variation in clam growth rate among habitats with different predation pressure, and between predator-exclusion cages and uncaged controls. I then carried out a predator-enclosure experiment using the clam and the whelk Busycon carica, a major predator on adult clams. Among-habitat variation in survivorship, predation rate, and growth rate was generally small or not detected in winter, while large variation was found for adul...

Biodiversity mediates top–down control in eelgrass ecosystems: a global comparative‐experimental approach

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors.

Re-Structuring of Marine Communities Exposed to Environmental Change: A Global Study on the Interactive Effects of Species and Functional Richness

Species richness is the most commonly used but controversial biodiversity metric in studies on aspects of community stability such as structural composition or productivity. The apparent ambiguity of theoretical and experimental findings may in part be due to experimental shortcomings and/or heterogeneity of scales and methods in earlier studies. This has led to an urgent call for improved and more realistic experiments. In a series of experiments replicated at a global scale we translocated several hundred marine hard bottom communities to new environments simulating a rapid but moderate environmental change. Subsequently, we measured their rate of compositional change (re-structuring) which in the great majority of cases represented a compositional convergence towards local communities. Re-structuring is driven by mortality of community components (original species) and establishment of new species in the changed environmental context. The rate of this re-structuring was then related to various system properties. We show that availability of free substratum relates negatively while taxon richness relates positively to structural persistence (i.e., no or slow re-structuring). Thus, when faced with environmental change, taxon-rich communities retain their original composition longer than taxon-poor communities. The effect of taxon richness, however, interacts with another aspect of diversity, functional richness. Indeed, taxon richness relates positively to persistence in functionally depauperate communities, but not in functionally diverse communities. The interaction between taxonomic and functional diversity with regard to the behaviour of communities exposed to environmental stress may help understand some of the seemingly contrasting findings of past research.

Global analysis of seagrass restoration: the importance of large-scale planting

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 trial survival - large numbers ensure the spread of risks, which is needed to overcome high natural variability. Secondly, a large-scale trial increases population growth rate by enhancing self-sustaining feedback, which is generally found in foundation species in stressful environments such as seagrass beds. Thus, by careful site selection and applying appropriate techniques, spreading of risks and enhancing self-sustaining feedback in concert increase success of seagrass restoration.

Plant–animal interactions in seagrass beds: ongoing and future challenges for understanding population and community dynamics

Seagrass beds are some of the most productive parts of coastal ecosystems, hosting a wide variety of associated fauna. This paper reviews recent studies of animal–plant interactions in seagrass beds, focusing particularly on studies conducted in Japan and Thailand. Although the positive effect of seagrass habitat structure on animals has been widely acknowledged, the magnitude of this effect varies greatly among studies. A comparative study on epifaunal communities and a manipulative experiment using an infaunal bivalve revealed that behavioral and life-history traits of component species and their interactions influence the observed variation. Some recent studies have challenged the previously accepted view that direct herbivory on seagrasses is rare, and has a minor effect on the seagrass community. A series of studies of dugong herbivory revealed that the marine mammal has great impacts not only on seagrass productivity but also on the infaunal community. Furthermore, it has been found that seed predators have a negative influence on seed production and the subsequent recruitment of seagrass. Recent studies have also demonstrated significant effects of fine-scale landscape patterns in seagrass vegetation on productivity, species interactions and community structure in seagrass beds. Future research integrating new concepts and theories in ecology, such as metapopulation and hierarchy theories, with new research tools, such as molecular-genetic analyses and remote-sensing techniques, may aid in developing a more comprehensive understanding of population and community dynamics in seagrass beds.

Latitudinal gradient of species diversity: multi-scale variability in rocky intertidal sessile assemblages along the Northwestern Pacific coast

This study examined the latitudinal gradient of species diversity of rocky intertidal sessile assemblages on the slopes of rocks along the Northwestern Pacific coast of Japan, located between 31°N and 43°N, by explicitly incorporating an hierarchical spatial scale into the monitoring design. The specific questions were to examine: (1) whether there is a latitudinal gradient of regional diversity, (2) how spatial components of the regional diversity (local diversity and turnover diversity) vary with latitude depending on spatial scale, and (3) whether the latitudinal gradient differs between different measures of species diversity, i.e. species richness and Simpson’s diversity index. We measured coverage and the presence or absence of all sessile organisms in a total of 150 census plots established at five shores in each of six regions. The results showed that there were clear latitudinal gradients in regional species richness and in species turnover among shores. However, these patterns were not reflected in smaller-scale local species richness. For Simpson’s diversity index, there was no evidence of latitudinal clines either in regional diversity or in spatial components. These results suggest that relative abundance of common species does not vary along latitude, while the number of rare species increases with decreasing latitude.

Predation on seeds of seagrasses Zostera marina and Zostera caulescens by a tanaid crustacean Zeuxo sp.

This paper reports the first evidence of seed predation by a tanaid crustacean on seagrasses Zostera marina and Z. caulescens in Otsuchi Bay, northeastern Japan. A tanaid, Zeuxo sp., was found to live in spathes of the seagrasses. Spathes with the tanaid were observed to have bored seeds. Laboratory observations revealed that the tanaid consumes seeds by drilling a hole. For both seagrass species, tanaids and bored ovules were not observed for spathes before anthesis, whereas spathes with bored seeds and density of the tanaid increased after anthesis, from June to August. In August, 14 and 27% of seeds were found bored in Z. marina and Z. caulescens, respectively, suggesting that seed predation by the tanaid has considerable negative impact on seed production of the seagrasses. Some adult individuals of Zeuxo sp. in spathes had brood pouches containing eggs and juveniles. A large number of juveniles were found in spathes in August. Thus, the tanaid utilizes seagrass spathes not only as feeding site, but also as place of reproduction and a nursery.

Temporal and spatial macrofaunal community changes along a salinity gradient in seagrass meadows of Akkeshi-ko estuary and Akkeshi Bay, northern Japan

Temporal and spatial variation of the macrofaunal community was investigated in seagrass meadows in Akkeshi-ko estuary and coastal area of Akkeshi Bay, northern Japan. We specifically addressed the question of how the salinity gradient in seagrass meadows affects the species richness, abundance and similarity of faunal groups classified based on the degree of association with the seagrasses, i.e. highly motile species that drift in the water column among seagrass blades (drift-fauna, DF group) and less motile species that are tightly associated with seagrass substrates (seagrass-associated fauna, SA group). A total of 70 species were collected semi-quantitatively using an epibenthic sledge, among which more than one third of the species were captured in all areas, and a quarter of species only in the marine area. Significant spatial variation in species richness, as well as a positive relationship between salinity and species richness was found for most sampling occasions and for both functional groups. Whereas, relationship between salinity and abundance of macrofauna was not clear although significant time and site interactions were found for both functional groups. Patterns of similarity of assemblages varied between the functional groups: clear differences by sampling sites were discerned for DF group but not for SA group. These results provided evidence that the macrofaunal community structures in seagrass beds varied with the salinity gradient, but the pattern differed with time and between functional groups, possibly due to the effect of biotic and abiotic factors that also changed with salinity.

Habitat Use by Fishes in Coral Reefs, Seagrass Beds and Mangrove Habitats in the Philippines

Understanding the interconnectivity of organisms among different habitats is a key requirement for generating effective management plans in coastal ecosystems, particularly when determining component habitat structures in marine protected areas. To elucidate the patterns of habitat use by fishes among coral, seagrass, and mangrove habitats, and between natural and transplanted mangroves, visual censuses were conducted semiannually at two sites in the Philippines during September and March 2010–2012. In total, 265 species and 15,930 individuals were recorded. Species richness and abundance of fishes were significantly higher in coral reefs (234 species, 12,306 individuals) than in seagrass (38 species, 1,198 individuals) and mangrove (47 species, 2,426 individuals) habitats. Similarity tests revealed a highly significant difference among the three habitats. Fishes exhibited two different strategies for habitat use, inhabiting either a single (85.6% of recorded species) or several habitats (14.4%). Some fish that utilized multiple habitats, such as Lutjanus monostigma and Parupeneus barberinus, showed possible ontogenetic habitat shifts from mangroves and/or seagrass habitats to coral reefs. Moreover, over 20% of commercial fish species used multiple habitats, highlighting the importance of including different habitat types within marine protected areas to achieve efficient and effective resource management. Neither species richness nor abundance of fishes significantly differed between natural and transplanted mangroves. In addition, 14 fish species were recorded in a 20-year-old transplanted mangrove area, and over 90% of these species used multiple habitats, further demonstrating the key role of transplanted mangroves as a reef fish habitat in this region.

Geographic variability in organic carbon stock and accumulation rate in sediments of East and Southeast Asian seagrass meadows

Organic carbon (OC) stored in the sediments of seagrass meadows has been considered a globally significant OC reservoir. However, the sparsity and regional bias of studies on long-term OC accumulation in coastal sediments have limited reliable estimation of the capacity of seagrass meadows as a global OC sink. We evaluated the amount and accumulation rate of OC in sediment of seagrass meadows and adjacent areas in East and Southeast Asia. In temperate sites, the average OC concentration in the top 30 cm of sediment was higher in seagrass meadows (780–1080 μmol g−1) than in sediments without seagrass cover (52–430 μmol g−1). The average OC in the top 30 cm of subtropical and tropical seagrass meadow sediments ranged from 140 to 440 μmol g−1. Carbon isotope mass balancing suggested that the contribution of seagrass-derived carbon to OC stored in sediments was often relatively minor (temperate: 10–40%; subtropical: 35–82%; tropical: 4–34%) and correlated to the habitat type, being particularly low in estuarine habitats. Stock of OC in the top meter of sediment of all the studied meadows ranged from 38 to 120 Mg ha−1. The sediment accumulation rates were estimated by radiocarbon dating of six selected cores (0.32–1.34 mm yr−1). The long-term OC accumulation rates calculated from the sediment accumulation rate and the top 30 cm average OC concentration for the seagrass meadows (24–101 kg ha−1 yr−1) were considerably lower than the OC accumulation rates previously reported for Mediterranean Posidonia oceanica meadows (580 kg ha−1 yr−1 on average). Current estimates for the global carbon sink capacity of seagrass meadows, which rely largely on Mediterranean studies, may be considerable overestimations.