Climate change mitigation by coral reefs and seagrass beds at risk: How global change compromises coastal ecosystem services.

Seagrass meadows provide valuable ecosystem services of coastal protection and chemical habitat formation that could help mitigate the impact of sea level rise and ocean acidification. However, the intensification of hydrodynamic forces caused by sea level rise, in addition to habitat degradation threaten the provision of these ecosystem services. With quantitative field measurements of the coastal protection and chemical habitat formation services of seagrass meadows, we statistically model the relationships between hydrodynamic forces, vegetation density and the provision of these ecosystem services. Utilising a high-resolution hydrodynamic model that simulates end of the century hydrodynamic conditions and three scenarios of coral reef degradation (i.e., keep up, remain or loss) we quantify how the environmental conditions within a tropical bay will change given changes to the provision of ecosystem services. Our study shows that increasing hydrodynamic forces lead to a seafloor made up of a larger grain size that is increasingly unstable and more vulnerable to erosion. The loss of a fringing reef leads to larger hydrodynamic forces entering the bay, however, the 0.87 m increase in depth due to sea-level rise reduces the bed shear stress in shallower areas, which limits the change in the ecosystem services provided by the current benthic seagrass meadow. Loss of seagrass constitutes the greatest change in a bay ecosystem, resulting in the sediment surface where seagrass existed becoming unstable and the median sediment grain size increasing by 5-7 %. The loss of seagrass also leads to the disappearance of the unique fluctuating chemical habitat, which leaves the surrounding community vulnerable to ocean acidification. A reduction or complete loss of these ecosystem services would impact the entire community assemblage while also leaving the surrounding coastline vulnerable to erosion, thus exacerbating negative effects brought about by climate change.

Salt marsh fragmentation in a mesotidal estuary: Implications for medium to long-term management.

During the last decades many salt marshes worldwide have suffered important losses in their extent and associated ecosystem services. The salt marshes of San Vicente de la Barquera estuary (N Spain) are a clear example of this, with a drastic reduction in vegetation surface over the last 60 years. This paper provides insights into the main factors controlling salt marsh functioning in sheltered estuarine areas. Regional and local factors have been disaggregated to identify the main drivers controlling the functioning of the salt marsh to develop appropriate management measures according to the evolution of the system. These factors have been studied in their spatial context through detailed maps of change in vegetation cover combined with topographic data obtained from UAV and RTK-DGPS surveys. The results demonstrate that in this estuary the salt marsh area is declining following a fragmentation process. No clear pattern of vegetation loss/gain with elevation has been identified. However, the results point to increased hydrodynamic stress in the area, with stronger currents inside the estuary. This is probably the major factor responsible for the decline of the salt marshes in the San Vicente de la Barquera estuary. Furthermore, several human interventions during the 20th century (local drivers) have also probably contributed to a lower resilience against SLR (regional driver). This work demonstrates that both natural and human drivers of change need to be considered when characterizing the evolution of salt marshes, wherever efficient management strategies need to be designed.

How to cope in heterogeneous coastal environments: Spatio-temporally endogenous circadian rhythm of valve gaping by mussels.

Transitional coastal zones are subject to high degrees of temporal fluctuation in environmental conditions, with these patterns varying in space. Gaining an in depth understanding of how sessile organisms cope with and respond to such environmental changes at multiple scales is needed to i) advance fundamental knowledge, ii) predict how organisms may react to stressors and iii) support the management of halieutic resources in transitional coastal areas. We addressed this question using mussels (Mytilus galloprovincialis) as model system. Valve-gaping sensor were deployed at multiple sites within the southern Venice Lagoon over a period of 6 months, to investigate the existence of periodicity in valve-gaping and its relationship with environmental variables, such as temperature and chlorophyll-a. Gaping behaviour was found to have periodic rhythms, of ~12 h and ~ 24 h, which were most pronounced in the inner part of lagoon part and were strongest during summer months. In autumn, the dual periodicity became weaker and mostly the 12 h remained. Gaping was closely linked with tide, but the relationship in terms of phasing varied upon location. Surprisingly, no clear direct relationships were found with chlorophyll-a, but food delivery may be mediated by tide itself. The results highlight the heterogeneity of behaviour and the endogenic nature of circadian rhythms in space and time. These findings have important implications for management of transitional areas where tidal alteration may have impacts on key behaviours, and emphasize the importance of characterizing their rhythms before using these as stress indicator. Moreover, the described tidal relationships should be included in growth models of bivalves in these systems.

Interactive effects of vegetation and sediment properties on erosion of salt marshes in the Northern Adriatic Sea.

We investigated how lateral erosion control, measured by novel photogrammetry techniques, is modified by the presence of Spartina spp. vegetation, sediment grain size, and the nutrient status of salt marshes across 230 km of the Italian Northern Adriatic coastline. Spartina spp. vegetation reduced erosion across our study sites. The effect was more pronounced in sandy soils, where erosion was reduced by 80% compared to 17% in silty soils. Erosion resistance was also enhanced by Spartina spp. root biomass. In the absence of vegetation, erosion resistance was enhanced by silt content, with mean erosion 72% lower in silty vs. sandy soils. We found no relevant relationships with nutrient status, likely due to overall high nutrient concentrations and low C:N ratios across all sites. Our results contribute to quantifying coastal protection ecosystem services provided by salt marshes in both sandy and silty sediments.

Pollen limitation may be a common Allee effect in marine hydrophilous plants: implications for decline and recovery in seagrasses.

Pollen limitation may be an important factor in accelerated decline of sparse or fragmented populations. Little is known whether hydrophilous plants (pollen transport by water) suffer from an Allee effect due to pollen limitation or not. Hydrophilous pollination is a typical trait of marine angiosperms or seagrasses. Although seagrass flowers usually have high pollen production, floral densities are highly variable. We evaluated pollen limitation for intertidal populations of the seagrass Zostera noltei in The Netherlands and found a significant positive relation between flowering spathe density and fruit-set, which was suboptimal at <1200 flowering spathes m(-2) (corresponding to <600 reproductive shoots m(-2)). A fragmented population had ≈35 % lower fruit-set at similar reproductive density than a continuous population. 75 % of all European populations studied over a large latitudinal gradient had flowering spathe densities below that required for optimal fruit-set, particularly in Southern countries. Literature review of the reproductive output of hydrophilous pollinated plants revealed that seed- or fruit-set of marine hydrophilous plants is generally low, as compared to hydrophilous freshwater and wind-pollinated plants. We conclude that pollen limitation as found in Z. noltei may be a common Allee effect for seagrasses, potentially accelerating decline and impairing recovery even after environmental conditions have improved substantially.

Early warning indicators for river nutrient and sediment loads in tropical seagrass beds: a benchmark from a near-pristine archipelago in Indonesia.

In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements.

Limited toxicity of NH(x) pulses on an early and late successional tropical seagrass species: interactions with pH and light level.

Seagrasses have declined at a global scale due to light reduction and toxicity events, caused by eutrophication and increased sediment loading. Although several studies have tested effects of light reduction and toxicants on seagrasses, there is at present no information available on their interacting effects. In a full-factorial 5-day laboratory experiment, we studied short-term interactive effects of light conditions, pH and reduced nitrogen (NH(x)) in the water layer, mimicking pulses of river discharge, on the tropical early successional species Halodule uninervis and the late successional species Thalassia hemprichii. In contrast to recent results reported for the temperate species Zostera marina, increased NH(x) supply did not affect leaf mortality or photochemical efficiency in H. uninervis and in 7 out of 8 treatments for T. hemprichii. However, both tropical species demonstrated striking differences in nitrogen accumulation, free amino acid composition and free NH3 accumulation. The increase in tissue nitrogen content was two times higher for H. uninervis than for T. hemprichii. Nitrogen stored as free amino acids (especially asparagine) only increased in H. uninervis. High pH only affected T. hemprichii, but only when not shaded, by doubling its free NH3 concentrations, concomitantly decreasing its photosynthetic efficiency. Our results indicate that the early successional H. uninervis has higher tolerance to high NH(x) loads as compared to the late successional T. hemprichii. H. uninervis was better able to avoid toxic internal NH(x) levels by further assimilating glutamine into asparagine in contrast to T. hemprichii. Moreover, both tropical species seem to cope much better with high NH(x) than the temperate Z. marina. The implications for the distribution and succession of seagrass species under high nutrient loads are discussed.

Comparing ecosystem engineering efficiency of two plant species with contrasting growth strategies.

Many ecosystems are greatly affected by ecosystem engineering, such as coastal salt marshes, where macrophytes trap sediment by reducing hydrodynamic energy. Nevertheless, little is known about the costs and benefits that are imposed on engineering species by the traits that underlie their ecosystem engineering capacity. We addressed this topic by comparing ecosystem engineering efficiency defined as the benefit-cost ratio per unit of biomass investment for two species from the intertidal habitat: the stiff grass Spartina anglica and the flexible grass Puccinellia maritima. These species were selected for their ability to modify their habitat by trapping large quantities of sediment despite their contrasting growth form. On a biomass basis, dissipation of hydrodynamic energy from waves (a proxy for benefits associated with ecosystem engineering capability as it relates to the sediment trapping capability) was strikingly similar for both salt marsh species, indicating that both species are equally effective in modifying their habitat. The drag forces per unit biomass (a proxy for costs associated with ecosystem engineering ability as it relates to the requirements on tissue construction and shoot anchoring to prevent breaking and/or washing away) were slightly higher in the species with flexible shoots. As a result, stiff Spartina vegetation had slightly higher ecosystem engineering efficiency, due to lower engineering costs rather than to a higher engineering effect. Thus, Spartina is a slightly more efficient rather than a more effective ecosystem engineer. Ecosystem engineering efficiency was found to be a species-specific characteristic, independent of vegetation density and relatively constant in space. Analyzing ecosystem engineering by quantifying trade-offs offers a useful way toward developing a better understanding of different engineering strategies.

Limited costs of wrong root placement in Rumex palustris in heterogeneous soils.

Nutrient hot spots in the soil have a limited life span, but the costs and benefits for root foraging are still underexposed. We assessed short-term costs that may arise when a nutrient-rich patch induces root proliferation, but then rapidly disappears. Rumex palustris plants were grown with a homogeneous or a heterogeneous nutrient application. After root proliferation in a nutrient-rich patch, nutrient supply was switched from homogeneous to heterogeneous, and vice versa, or the patch location was changed. R. palustris proliferated its roots in the rich patch. After switching, the relative growth rates of the roots were adjusted to the novel pattern of nutrient availability. However, the changes in local root biomass lagged behind the rapid shift in nutrient supply, because the root mass realized in specific sectors could not be rapidly relocated. Despite this, R. palustris did not exhibit costs of switching in terms of biomass or nitrogen uptake. Our data suggest that rapid shifts in uptake rate and redistribution of nitrogen within the plant may have lowered the costs of incorrect root placement.

Linking root traits to potential growth rate in six temperate tree species.

There is an extremely limited understanding of how plants of different potential growth rate vary in root traits, especially in woody species. We contrasted fine root morphology, physiology, and elemental construction between a fast- and a slow-growing species in each of three families: Aceraceae (maple), Fagaceae (oak), and Pinaceae (pine). Measurements were primarily made on 1-year-old seedlings growing in a growth chamber. Across all three families, first- and second-order roots of fast-growing species had greater specific root length, thinner diameters, and faster respiration rates than those of slow-growing species. These fine roots of fast-growing species in Aceraceae and Fagaceae also had faster phosphorus (P) uptake on a surface area basis than those of slow-growing species, whereas little difference in P uptake was found between Pinaceae species. On a dry weight basis, roots of fast-growing species in Aceraceae and Fagaceae had higher nitrogen concentrations, lower carbon:nitrogen ratios and higher tissue construction costs than roots of slow-growing species (data were unavailable for Pinaceae). Tissue density did not vary in a consistent pattern between fast- and slow-growing species across all three families (P=0.169). To better understand the ecological significance of differences in these root characteristics, a root efficiency model was used to compare P uptake and root carbon (C) cost of each species in simulated field situations in two soils, one with low P buffering capacity (loamy sand) and another with relatively high P buffering capacity (silt loam). For the soil conditions modeled, fast-growing species of Aceraceae and Fagaceae were 17-70% more efficient (defined as cumulative P gain divided by cumulative C cost) at nutrient capture than slow-growing species while the fast-growing Pinaceae species was 20-24% less efficient than the slow-growing species. However, among all three families, roots of fast-growing species reached maximum lifetime efficiency 5-120 days sooner, depending on soil type. Thus, modeling results indicated that root traits of fast- and slow-growing species affected P acquisition in simulated field soil although soil type also had a strong impact.

Protein targeting towards the thylakoid lumen of chloroplasts: proper localization of fusion proteins is only observed in vivo.

Routeing of fusion proteins to the thylakoid lumen of the chloroplast was compared in vitro and in vivo. The Escherichia coli protein beta-lactamase was used as a passenger to study this intraorganellar sorting process. The first step, translocation of beta-lactamase into the chloroplast stroma, occurs properly both in vitro and in vivo and is dependent on the presence of a transit peptide in the protein construct. The second step, targeting towards the thylakoid lumen, is more complicated as was also observed previously when other passenger proteins were used. In vitro, the presence of a thylakoid transfer domain is not enough for routeing and proper processing. Only when the complete thylakoid lumen precursor plastocyanin was fused to beta-lactamase was the fusion protein processed adequately, but routeing was still incomplete. However, in vivo, the information present in the thylakoid transfer domain was the only requirement for proper transport towards the thylakoid lumen. These data show that in vivo, the only requirement for targeting of passenger proteins towards the thylakoid lumen is the presence of a transit peptide and a thylakoid transfer domain. Furthermore, we demonstrate that the in vitro import system does not necessarily reflect the in vivo situation with respect to intraorganellar sorting.