Seagrass Colonization Research Articles

Patch age alters seagrass response mechanisms to herbivory damage

Natural disturbances can produce a mosaic of seagrass patches of different ages, which may affect the response to herbivory. These pressures can have consequences for plant performance. To assess how seagrass patch age affects the response to herbivory, we simulated the effect of herbivory by clipping leaves of Halodule wrightii in patches of 2, 4 and 6 years. All clipped plants showed ability to compensate herbivory by increasing leaf growth rate (on average 4.5-fold). The oldest patches showed resistance response by increasing phenolic compounds (1.2-fold). Contrastingly, the concentration of phenolics decreased in the youngest patches (0.26-fold), although they had a similar leaf carbon content to controls. These results suggest that younger plants facing herbivory pressure reallocate their phenolic compounds towards primary metabolism. Results confirm the H. wrightii tolerance to herbivory damage and provides evidence of age-dependent compensatory responses, which may have consequences for seagrass colonization and growth in perturbed habitats.

A 2,000‐Year Record of Eelgrass (Zostera marina L.) Colonization Shows Substantial Gains in Blue Carbon Storage and Nutrient Retention

AbstractAssessing historical environmental conditions linked to habitat colonization is important for understanding long‐term resilience and improving conservation and restoration efforts. Such information is lacking for the seagrass Zostera marina, an important foundation species across cold‐temperate coastal areas of the Northern Hemisphere. Here, we reconstructed environmental conditions during the last 14,000 years from sediment cores in two eelgrass (Z. marina) meadows along the Swedish west coast, with the main aims to identify the time frame of seagrass colonization and describe subsequent biogeochemical changes following establishment. Based on vegetation proxies (lipid biomarkers), eelgrass colonization occurred about 2,000 years ago after geomorphological changes that resulted in a shallow, sheltered environment favoring seagrass growth. Seagrass establishment led to up to 20‐ and 24‐fold increases in sedimentary carbon and nitrogen accumulation rates, respectively. This demonstrates the capacity of seagrasses as efficient ecosystem engineers and their role in global change mitigation and adaptation through CO2 removal, and nutrient and sediment retention. By combining regional climate projections and landscape models, we assessed potential climate change effects on seagrass growth, productivity and distribution until 2100. These predictions showed that seagrass meadows are mostly at risk from increased sedimentation and hydrodynamic changes, while the impact from sea level rise alone might be of less importance in the studied area. This study showcases the positive feedback between seagrass colonization and environmental conditions, which holds promise for successful conservation and restoration efforts aimed at supporting climate change mitigation and adaptation, and the provision of several other crucial ecosystem services.

Mangrove ghost forests provide opportunities for seagrass

Mangrove forests are degraded by extreme climatic events worldwide, often leaving behind dead standing stems called “ghost forests”. Ghost forests may provide opportunities for seagrass colonization but there is limited research into the conditions found within these ecosystems, or whether they provide a suitable habitat for seagrasses. This study aimed to characterize the environmental conditions within mangrove ghost forests, determine whether conditions are suitable for seagrass survival, and identify whether seagrass was present within the ghost forests of Moreton Bay, Queensland. Six study locations within mangrove ghost forests adjacent to live mangrove forests and seagrass meadows were selected and biophysical conditions within these habitats over the six sites were assessed. Two of the six study sites were found to have live seagrass present within the ghost forests (Godwin Beach and South Stradbroke). Suitable water temperature was linked to the presence and abundance of seagrass within mangrove ghost forests. Mangrove characteristics, including canopy cover, diameter at breast height, and stem densities, contributed to variation in the environment among the three habitats, suggesting that light is a key factor limiting seagrass colonization into live or ghost mangrove forests. Overall, these results suggest that ghost forests can provide suitable habitats for seagrasses, and degraded mangroves may transition to seagrass under future sea level rise scenarios.

Genome-resolved metagenomics provides insights into the microbial-mediated sulfur and nitrogen cycling in temperate seagrass meadows

The presence of seagrasses facilitates numerous microbial-mediated biogeochemical cycles, with sulfur- and nitrogen-cycling microorganisms playing crucial roles as regulators. Despite efforts to comprehend the diversity of microbes in seagrass ecosystems, the metabolic functions of these benthic microorganisms in seagrass sediments remain largely unknown. Using metagenomics, we provide insights into the sulfur- and nitrogen-cycling pathways and key metabolic capacities of microorganisms in both Z. japonica-colonized and unvegetated sediments over a seasonal period. Taxonomic analysis of N and S cycling genes revealed that δ- and γ- proteobacteria dominated the benthic sulfate-reducing bacteria, while α- and γ-proteobacteria played a significant role in the sulfur-oxidation processes. The proteobacterial lineages were also major contributors to the benthic nitrogen cycling. However, at a finer taxonomic resolution, microbial participants in different processes were observed to be highly diverse and mainly driven by environmental factors such as temperature and salinity. The gene pools of sulfur and nitrogen cycles in the seagrass sediments were dominated by genes involved in sulfide oxidation (fccA) and hydroxylamine oxidation (hao), respectively. Seagrass colonization elevated the relative abundance of genes responsible for sulfite production (phsC), hydroxylamine oxidation (hao), and nitrogen fixation (nifK), but suppressed sulfur oxidation (soxXYZ) and denitrification (nosZ and nirS). The prevalence of proteobacterial lineages functioned with versatile capabilities in both sulfur and nitrogen cycles in seagrass ecosystems, highlighting tight couplings between these processes, which was further supported by the recovery of 83 metagenome-assembled genomes (MAGs). These findings broaden our understanding of the biogeochemical processes that are mediated by microorganisms in seagrass ecosystems.

Dark waters: Evaluating seagrass community response to optical water quality and freshwater discharges in a highly managed subtropical estuary

Seagrass communities are vitally important ecosystems that support and foster organisms across trophic levels. Understanding the drivers of seagrass distribution and extent in estuary systems is valuable for conservation and restoration efforts. A critical driver of seagrass distribution is how light moves through the water column (i.e. light attenuation) and the availability of light to reach the estuary bottom. Light attenuation combined with the depth of seagrass colonization is used to estimate the percent surface irradiance and is an indicator of seagrass light requirements The objective of this study was to evaluate optical water quality parameters within the Caloosahatchee River Estuary (CRE) relative to changes in freshwater discharge conditions. During the study period, freshwater discharge significantly increased resulting in an increase of annual stressful and damaging discharge events to marine seagrass species. Concurrent with changes in freshwater discharge conditions, changes to optical water quality parameters including color, chlorophyll-a, total suspended solids, and light attenuation were detected along the estuary. Using photo-interpreted seagrass coverage data combined with bathymetric data, the depth of colonization was estimated for three survey years (2008, 2014, and 2020). This information combined with a spatial and temporal light attenuation generalized additive model percent surface irradiance (%SI) was estimated. Over the study period, %SI significantly increased across the lower CRE indicating an improvement in light attenuation. The current freshwater management plan has increased stressful and damaging discharge events, however recent changes in operations has minimized these damaging events which could be linked to the precipitous increase in %SI across the lower CRE. Seagrass density data collected from transects within the estuary suggest that prolonged stress and damaging (>5.13 hm3d−1) has significantly reduce species density. Future water management strategies are expected to reduce the frequency and volume of discharges to the CRE, effectively reducing stress and damaging freshwater discharge event.

Seagrass Colonization Alters Diversity, Abundance, Taxonomic, and Functional Community Structure of Benthic Microbial Eukaryotes.

Seagrass form high productive ecosystems in coastal environments. However, the effects of these coastal plants on the structure and function of the belowground eukaryotic microbiome remain elusive. In this study, we characterized the community of microbial eukaryotes (microeukaryotes) in both vegetated and unvegetated sediments using 18S rRNA gene amplicon sequencing and quantitative PCR. Analysis of sequencing data showed that the eelgrass (Zostera marina) colonization decreased the alpha diversity indices of benthic microeukaryotes. Apicomplexa represented an average of 83% of reads across all samples, with a higher proportion at the vegetated sites. The taxonomic community structure was significantly different between these two types of sediments, for which the concentration of in sediment porewater and salinity could account. Phylogenetic analyses of long 18S rRNA genes (around 1,030 bp) indicated these apicomplexan parasites are closely related to gregarine Lecudina polymorpha. Determination of 18S rRNA gene abundances provided evidence that the eelgrass markedly promoted the biomass of the gregarine and all microeukaryotes in the seagrass-colonized sediments and confirmed that the gregarine was hosted by a polychaete species. Significantly higher gene abundances of heterotrophs and mixotrophs were found at the vegetated sites, which could be explained by the finer sediments and short supply of dissolved inorganic nitrogen, respectively. The pigmented protists were more abundant in 18S rRNA gene copies at the lower and higher pH levels than at the intermediate. Nevertheless, the fractions of heterotrophs and phototrophs in the community were significantly related to porewater N:P ratio. These results indicate that seagrass colonization significantly induces an increase in overall biomass and a decrease in diversity of benthic microeukaryotes, making them more heterotrophic. This study also highlights that the hotspot of eukaryotic parasites could be linked with the high productivity of a natural ecosystem.

Effects of Organic Phosphorus on Methylotrophic Methanogenesis in Coastal Lagoon Sediments With Seagrass (Zostera marina) Colonization.

Methanogens are the major contributors of greenhouse gas methane and play significant roles in the degradation and transformation of organic matter. These organisms are particularly abundant in Swan Lake, which is a shallow lagoon located in Rongcheng Bay, Yellow Sea, northern China, where eutrophication from overfertilization commonly results in anoxic environments. High organic phosphorus content is a key component of the total phosphorus in Swan Lake and is possibly a key factor affecting the eutrophication and carbon and nitrogen cycling in Swan Lake. The effects of organic phosphorus on eutrophication have been well-studied with respect to bacteria, such as cyanobacteria, unlike the effects of organic phosphorus on methanogenesis. In this study, different sediment layer samples of seagrass-vegetated and unvegetated areas in Swan Lake were investigated to understand the effects of organic phosphorus on methylotrophic methanogenesis. The results showed that phytate phosphorus significantly promoted methane production in the deepest sediment layer of vegetated regions but suppressed it in unvegetated regions. Amplicon sequencing revealed that methylotrophic Methanococcoides actively dominated in all enrichment samples from both regions with additions of trimethylamine or phytate phosphorus, whereas methylotrophic Methanolobus and Methanosarcina predominated in the enrichments obtained from vegetated and unvegetated sediments, respectively. These results prompted further study of the effects of phytate phosphorus on two methanogen isolates, Methanolobus psychrophilus, a type strain, Methanosarcina mazei, an isolate from Swan Lake sediments. Cultivation experiments showed that phytate phosphorus could inhibit methane production by M. psychrophilus but promote methane production by M. mazei. These culture-based studies revealed the effects of organic phosphorus on methylotrophic methanogenesis in coastal lagoon sediments and improves our understanding of the mechanisms of organic carbon cycling leading to methanogenesis mediated by organic phosphorus dynamics in coastal wetlands.

Community Structure and Abundance of Archaea in a Zostera marina Meadow: A Comparison between Seagrass-Colonized and Bare Sediment Sites.

Seagrass colonization alters sediment physicochemical properties by depositing seagrass fibers and releasing organic carbon and oxygen from the roots. How this seagrass colonization-induced spatial heterogeneity affects archaeal community structure and abundance remains unclear. In this study, we investigated archaeal abundance, diversity, and composition in both vegetated and adjacent bare surface sediments of a Zostera marina meadow. High-throughput sequencing of 16S rDNA showed that Woesearchaeota, Bathyarchaeota, and Thaumarchaeota were the most abundant phyla across all samples, accounting for approximately 42%, 21%, and 17% of the total archaeal communities, respectively. In terms of relative abundance, Woesearchaeota and Bathyarchaeota were not significantly different between these two niches; however, specific subclades (Woese-3, Woese-21, Bathy-6, Bathy-18) were significantly enriched in vegetated sediments (P < 0.05), while Thaumarchaeota was favored in unvegetated sites (P = 0.02). The quantification of archaeal 16S rRNA genes showed that the absolute abundance of the whole archaeal community, Bathyarchaeota, and Woese-3, Woese-10, Woese-13, and Woese-21 was significantly more abundant in vegetated sediments than in bare sediments (P < 0.05). Our study expands the available knowledge of the distribution patterns and niche preferences of archaea in seagrass systems, especially for the different subclades of Woesearchaeota and Bathyarchaeota, in terms of both relative proportions and absolute quantities.

Seagrass (Zostera marina) Colonization Promotes the Accumulation of Diazotrophic Bacteria and Alters the Relative Abundances of Specific Bacterial Lineages Involved in Benthic Carbon and Sulfur Cycling.

Seagrass colonization changes the chemistry and biogeochemical cycles mediated by microbes in coastal sediments. In this study, we molecularly characterized the diazotrophic assemblages and entire bacterial community in surface sediments of a Zostera marina-colonized coastal lagoon in northern China. Higher nitrogenase gene (nifH) copy numbers were detected in the sediments from the vegetated region than in the sediments from the unvegetated region nearby. The nifH phylotypes detected were mostly affiliated with the Geobacteraceae, Desulfobulbus, Desulfocapsa, and Pseudomonas. Redundancy analysis based on terminal restriction fragment length polymorphism analysis showed that the distribution of nifH genotypes was mostly shaped by the ratio of total organic carbon to total organic nitrogen, the concentration of cadmium in the sediments, and the pH of the overlying water. High-throughput sequencing and phylogenetic analyses of bacterial 16S rRNA genes also indicated the presence of Geobacteraceae and Desulfobulbaceae phylotypes in these samples. A comparison of these results with those of previous studies suggests the prevalence and predominance of iron(III)-reducing Geobacteraceae and sulfate-reducing Desulfobulbaceae diazotrophs in coastal sedimentary environments. Although the entire bacterial community structure was not significantly different between these two niches, Desulfococcus (Deltaproteobacteria) and Anaerolineae (Chloroflexi) presented with much higher proportions in the vegetated sediments, and Flavobacteriaceae (Bacteroidetes) occurred more frequently in the bare sediments. These data suggest that the high bioavailability of organic matter (indicated by relatively lower carbon-to-nitrogen ratios) and the less-reducing anaerobic condition in vegetated sediments may favor Desulfococcus and Anaerolineae lineages, which are potentially important populations in benthic carbon and sulfur cycling in the highly productive seagrass ecosystem.

Assessing Posidonia oceanica seedling substrate preference: an experimental determination of seedling anchorage success in rocky vs. sandy substrates.

In the last decades the growing awareness of the ecological importance of seagrass meadows has prompted increasing efforts to protect existing beds and restore degraded habitats. An in-depth knowledge of factors acting as major drivers of propagule settlement and recruitment is required in order to understand patterns of seagrass colonization and recovery and to inform appropriate management and conservation strategies. In this work Posidonia oceanica seedlings were reared for five months in a land-based culture facility under simulated natural hydrodynamic conditions to identify suitable substrates for seedling anchorage. Two main substrate features were investigated: firmness (i.e., sand vs. rock) and complexity (i.e., size of interstitial spaces between rocks). Seedlings were successfully grown in culture tanks, obtaining overall seedling survival of 93%. Anchorage was strongly influenced by substrate firmness and took place only on rocks, where it was as high as 89%. Anchorage occurred through adhesion by sticky root hairs. The minimum force required to dislodge plantlets attached to rocky substrates reached 23.830 N (equivalent to 2.43 kg), which would potentially allow many plantlets to overcome winter storms in the field. The ability of rocky substrates to retain seedlings increased with their complexity. The interstitial spaces between rocks provided appropriate microsites for seedling settlement, as seeds were successfully retained, and a suitable substrate for anchorage was available. In conclusion P. oceanica juveniles showed a clear-cut preference for hard substrates over the sandy one, due to the root system adhesive properties. In particular, firm and complex substrates allowed for propagule early and strong anchorage, enhancing persistence and establishment probabilities. Seedling substrate preference documented here leads to expect a more successful sexual recruitment on hard bottoms compared with soft ones. This feature could have influenced P. oceanica patterns of colonization in past and present time.

Ecosystem metabolism along a colonization gradient of eelgrass (Zostera marina) measured by eddy correlation

The Virginia coastal bays experienced local extinction of eelgrass (Zostera marina) during the early 1930s, and restoration beginning in 2001 has generated an ecosystem state change from bare to vegetated sediments. Oxygen fluxes were measured seasonally using the eddy correlation technique at three sites representing different stages of seagrass colonization: unvegetated (bare), 5 yr, and 11 yr since seeding. Derived seasonal ecosystem respiration (R) and gross primary production (GPP) increased up to 10‐fold and 25‐fold, respectively, with meadow age. Although hourly oxygen (O2) fluxes were highly correlated with light at the vegetated sites, no identifiable trends with light were observed at the bare site. The light compensation point where O2 production and respiration are in balance increased from 46 µmol photons m−2 s−1 to 257 µmol photons m−2 s−1 and 63 µmol photons m−2 s−1 to 472 µmol photons m−2 s−1 at the 5 yr and 11 yr seagrass sites, respectively, with increasing seasonal temperatures from 12.3°C to 27.9°C and 9.3°C to 30.5°C, respectively. This suggests that more light, and thus more O2 production, is required to offset increasing respiration with both temperature and meadow age. Photosynthesis—irradiance curves generated from hourly O2 fluxes throughout the seasons were used to estimate annual net ecosystem metabolism (NEM). Annual NEM rates at the bare, 5 yr, and 11 yr sites were −7.6, 8.6, and −7.0 mol O2 m−2 yr−1, respectively. Although the system went through a period of net autotrophy during early stages of colonization, the ecosystem state change from unvegetated sediments to dense seagrass meadows changed the magnitude of both GPP and R, but not the overall metabolic balance of the system.

Spatial Structure of Seagrass Suggests That Size-Dependent Plant Traits Have a Strong Influence on the Distribution and Maintenance of Tropical Multispecies Meadows

BackgroundSeagrass species in the tropics occur in multispecies meadows. How these meadows are maintained through species co-existence and what their ecological drivers may be has been an overarching question in seagrass biogeography. In this study, we quantify the spatial structure of four co-existing species and infer potential ecological processes from these structures.Methods and ResultsSpecies presence/absence data were collected using underwater towed and dropped video cameras in Pulau Tinggi, Malaysia. The geostatistical method, utilizing semivariograms, was used to describe the spatial structure of Halophila spp, Halodule uninervis, Syringodium isoetifolium and Cymodocea serrulata. Species had spatial patterns that were oriented in the along-shore and across-shore directions, nested with larger species in meadow interiors, and consisted of multiple structures that indicate the influence of 2–3 underlying processes. The Linear Model of Coregionalization (LMC) was used to estimate the amount of variance contributing to the presence of a species at specific spatial scales. These distances were <2.5 m (micro-scale), 2.5–50 m (fine-scale) and >50 m (broad-scale) in the along-shore; and <2.5 m (micro-scale), 2.5–140 m (fine-scale) and >140 m (broad-scale) in the across-shore. The LMC suggests that smaller species (Halophila spp and H. uninervis) were most influenced by broad-scale processes such as hydrodynamics and water depth whereas large, localised species (S. isoetifolium and C. serrulata) were more influenced by finer-scale processes such as sediment burial, seagrass colonization and growth, and physical disturbance.ConclusionIn this study, we provide evidence that spatial structure is distinct even when species occur in well-mixed multispecies meadows, and we suggest that size-dependent plant traits have a strong influence on the distribution and maintenance of tropical marine plant communities. This study offers a contrast from previous spatial models of seagrasses which have largely focused on monospecific temperate meadows.

Seagrass colonization: Knock-on effects on zoobenthic community, populations and individual health

This study provided evidence that Zostera noltii presence affects macrofauna community structure independently from median sediment grain-size and that the notion of ecosystem health is rather subjective: in the present case, we recorded “good health” in terms of seagrass development, “no impact” in terms of macrobenthic biotic indices and “negative effect” for a given key-population. The occurrence and development of a Z. noltii seagrass bed was surveyed at Banc d’Arguin, Arcachon Bay (France), to estimate the modification of the macrozoobenthic community and of the dynamics of a key-population for the local ecosystem, – the cockle Cerastoderma edule. Even though median grain-size of the sediment decreased only at the very end of the survey, i.e. when seagrass totally invaded the area, most of the macrofauna community characteristics (such as abundance and biomass) increased as soon as Z. noltii patches appeared. The structure of the macrofauna community also immediately diverged between sand and seagrass habitats, without however modifying the tested biotic indices (BENTIX, BOPA, AMBI). The health of the cockle population (growth, abundance, recruitment) was impacted by seagrass development. Related parasite communities slowly diverged between habitats, with more parasites in the cockles from seagrass areas. However, the number of parasites per cockle was always insufficient to alter cockle fitness.

Understanding uncertainty in seagrass injury recovery: an information-theoretic approach

Vessel groundings cause severe, persistent gaps in seagrass beds. Varying degrees of natural recovery have been observed for grounding injuries, limiting recovery prediction capabilities, and therefore, management's ability to focus restoration efforts where natural recovery is unlikely. To improve our capacity for predicting seagrass injury recovery, we used an information-theoretic approach to evaluate the relative contribution of specific injury attributes to the natural recovery of 30 seagrass groundings in Florida Keys National Marine Sanctuary, Florida, USA. Injury recovery was defined by three response variables examined independently: (1) initiation of seagrass colonization, (2) areal contraction, and (3) sediment in-filling. We used a global model and all possible subsets for four predictor variables: (1) injury age, (2) original injury volume, (3) original injury perimeter-to-area ratio, and (4) wave energy. Successional processes were underway for many injuries with fast-growing, opportunistic seagrass species contributing most to colonization. The majority of groundings that exhibited natural seagrass colonization also exhibited areal contraction and sediment in-filling. Injuries demonstrating colonization, contraction, and in-filling were on average older and smaller, and they had larger initial perimeter-to-area ratios. Wave energy was highest for colonizing injuries. The information-theoretic approach was unable to select a single "best" model for any response variable. For colonization and contraction, injury age had the highest relative importance as a predictor variable; wave energy appeared to be associated with second-order effects, such as sediment in-filling, which in turn, facilitated seagrass colonization. For sediment in-filling, volume and perimeter-to-area ratio had similar relative importance as predictor variables with age playing a lesser role than seen for colonization and contraction. Our findings confirm that these injuries naturally initiate seagrass colonization with the potential to recover to pre-injury conditions, but likely on a decadal scale given the slow growth of the climax species (Thalassia testudinum), which is often the most severely injured. Our analysis supports current perceptions that sediment in-filling is critical to the recovery process and indicates that in order to stabilize injuries and facilitate seagrass recovery, managers should consider immediate restorative filling procedures for injuries having an original volume >14-16 m3.

Damage and recovery assessment of vessel grounding injuries on coral reef habitats by use of georeferenced landscape video mosaics

Vessel groundings are a major source of disturbance to coral reefs worldwide. Documenting the extent of damage caused by groundings is a crucial first step in the reef restoration process. Here, we describe the application of a novel survey methodology, landscape video mosaics, to assessment of the damage caused by vessel groundings. Video mosaics, created by merging thousands of video frames, combine quantitative and qualitative aspects of damage assessment and provide a georeferenced, landscape, high‐resolution, spatially accurate permanent record of an injury. The scar in a Florida reef impacted by a 49‐foot vessel, imaged in 2005 and 2006, covered an area of 150 m2 (total imaged area was &gt;600 m2). The impacted coral community showed limited signs of coral recovery more than 3 years after the initial impact; the cover of corals was still significantly higher in the undamaged areas compared to the scar. However, seagrass colonization of the scar was observed. Finally, no evidence of further physical impacts was documented even when four hurricanes passed near the grounding site in 2005. The video mosaics developed in this study proved to be ideal tools to survey the grounding scars. Mosaics provide a means to collect information on the size of the damage area and the status and trends of the impacted biological communities and provide a permanent visual record of the damage, thereby expanding the quality and diversity of information that can be collected during field surveys.

Facies analysis and palaeoenvironmental interpretation of the Late Oligocene Attard Member (Lower Coralline Limestone Formation), Malta

AbstractThe Oligocene represents a key interval during which coralline algae became dominant on carbonate ramps and luxuriant coral reefs emerged on a global scale. So far, few studies have considered the impact that these early reefs had on ramp development. Consequently, this study aimed at presenting a high‐resolution analysis of the Attard Member of the Lower Coralline Limestone Formation (Late Oligocene, Malta) in order to decipher the internal and external factors controlling the architecture of a typical Late Oligocene platform. Excellent exposures of the Lower Coralline Limestone Formation occurring along continuous outcrops adjacent to the Victoria Lines Fault reveal in detail the three‐dimensional distribution of the reef‐associated facies. A total of four sedimentary facies have been recognized and are grouped into two depositional environments that correspond to the inner and middle carbonate ramp. The inner ramp was characterized by a very high‐energy, shallow‐water setting, influenced by tide and wave processes. This setting passed downslope into an inner‐ramp depositional environment which was colonized by seagrass and interfingered with adjacent areas containing scattered corals. The middle ramp lithofacies were deposited in the oligophotic zone, the sediments being generated from combined in situ production and sediments swept from the shallower inner ramp by currents. Compositional characteristics and facies distributions of the Attard ramp are more similar to the Miocene ramps than to those of the Eocene. An important factor controlling this similarity may be the expansion of the seagrass colonization within the euphotic zone. This expansion may have commenced in the Late Oligocene and was associated with a concomitant reduction in the aerial extent of the larger benthonic foraminifera facies. Stacking‐pattern analysis shows that the depositional units (parasequences) at the study section are arranged into transgressive–regressive facies cycles. This cyclicity is superimposed on the overall regressive phase recorded by the Attard succession. Furthermore, a minor highstand (correlated with the Ru4/Ch1 sequence) and subsequent minor lowstand (Ch2 sequence) have been recognized. The biota assemblages of the Attard Member suggest that carbonate sedimentation took place in subtropical waters and oligotrophic to slightly mesotrophic conditions. The apparent low capacity of corals to form wave‐resistant reef structures is considered to have been a significant factor affecting substrate stability at this time. The resulting lack of resistant mid‐ramp reef frameworks left this zone exposed to wave and storm activity, thereby encouraging the widespread development of coralline algal associations dominated by rhodoliths.

Testing the predictive power of seagrass depth limit models

The power of equations predicting seagrass depth limit (Zc) from light extinction (K z) was tested on data on seagrass depth limits collected from the literature. The test data set comprised 424 reports of seagrass colonization depth and water transparency, including data for 10 seagrass species. This data set confirmed the strong negative relationship betweenZ c andK z. The regression equation in Duarte (1991) overestimated the realized seagrass colonization depths at colonization depths < 5 m, while there was no prediction bias above this threshold. These results indicated that seagrass colonizing turbid waters (K z 0.27 m-1) have higher apparent light requirements than those growing in clearer waters. The relationship between seagrass colonization depth and light attenuation shifts at a threshold of light attenuation of 0.27 m-1, requiring separate equations to predictZ c for seagrass growing in more turbid waters and clearer waters, and to set targets for seagrass restoration and conservation efforts.

Community metabolism and carbon budget along a gradient of seagrass(Cymodocea nodosa) colonization

We studied the effect of seagrass (Cymodocea nodosa) colonization on community metabolism and sediment conditions. The biomass of C. nodosa increased with time after seagrass colonization. The biomass increased steadily during the 6.1 yr of colonization to 9.1 mol C m−2. Gross primary production increased from 7 to 49.3 mmol C m−2 d−1 during the first stages of the colonization and then decreased to 20 mmol C m−2 d−1 at a time when the biomass was in excess of 6 mol C m−2. Net dissolved organic carbon (DOC) fluxes increased with time after colonization, shifting from a net uptake in patches younger than 2 yr to a net release in older patches. Community respiration (R) increased with the seagrass colonization, leading to a shift from net autotrophy in the unvegetated sediment community to net heterotrophy after C. nodosa colonization. The increase in net heterotrophy with seagrass colonization was reflected in the development of reducing conditions in the sediment. To maintain a net heterotrophic benthic metabolism and net DOC release, the C. nodosa community must receive an input of organic matter (OM) from an allochthonous source. OM inputs from sestonic material trapped by the seagrass canopy exceed 157 mmol C m−2 d−1 in developed C. nodosa communities. Thus, the seagrass community acts as an important link between the pelagic and benthic communities by trapping sestonic organic carbon.

The influence of burrowing thalassinid shrimps on the distribution of intertidal seagrasses in Willapa Bay, Washington, USA

Two species of seagrasses frequently co-occur with extensive thalassinid shrimp populations and aquaculture operations in the intertidal zone of estuaries along the west coast of North America. Although thalassinid shrimp are known to be strong bioturbators and affect both aquaculture operations and benthic intertidal community structure, few studies have investigated shrimp–seagrass interactions. Application of the pesticide carbaryl to control shrimp populations for oyster aquaculture in Willapa Bay, Washington provided us with an experimental tool to investigate one such interaction. We found that the seagrass Zostera japonica colonized areas where ghost shrimp ( Neotrypaea californiensis) had been removed via carbaryl application. We applied carbaryl to small (900 m 2) experimental plots and compared seagrass colonization on these to that on control plots where shrimp remained abundant (100 m −2). The cumulative proportion of Z. japonica seeds and sprouts was slightly higher in the surface layer of treated plots (presumably due to the lack of shrimp bioturbation distributing them to depth), but seedling abundance was not significantly different between treated (no shrimp) and untreated control plots when they first emerged in early spring. As the season progressed however, and shrimp became more active, fewer seedlings survived in the untreated areas, and those that did survive grew much more slowly than those in the treated plots. We suspect that this was due to the effects of shrimp bioturbation and either light limitation (shoots that survived were much smaller) or direct burial and loss. Although it is an introduced plant, the natural distribution of Z. japonica is high in the intertidal zone and it is often separated from its congener Zostera marina by an expansive sandflat that is dominated by the ghost shrimp N. californiensis in west coast estuaries. The treatment of intertidal oysterbeds with carbaryl clearly reduces abundance of shrimp in this zone and we documented the same pattern of seagrass colonization on a commercial oyster bed and lack of seagrass in an adjacent unsprayed area. Density of native seagrass Z. marina shoots was also enhanced in plots treated with carbaryl, but only at lower tidal elevations or in intertidal pools where it could survive. We believe the removal of shrimp will continue to broaden the distribution of Z. japonica in Washington coastal estuaries where carbaryl use is permitted and add an interesting perspective to this controversial management issue.

Do nutrient availability and plant density limit seagrass colonization in the Baltic Sea?

Seagrasses continue to decline at an alarming rate throughout the planet's temperate regions. After a decline recolonization or restoration starts from small patches of single shoots which then propagate vegetatively. We investigated the effects of plant density within a patch and nutrient resources on growth and survival of eelgrass (Zostera marina L.), the dominant seagrass species in the northern temperate zone. We created small (0.5 m') eelgrass patches by planting single shoots in circular plots at high (20 cm) and low (40 cm distance between shoots) density. In a factorial design, the sediment was nutrient-enriched (1) though biodeposition of transplanted mussels (Meus edulisL.) (2) by a slow- release NPK-fertilizer or (3) not enriched. The experiment was run over 1 growth period at a relatively nutrient-poor site (<30 pm01 NH4+ l-' porewater) in the Baltic Sea. Mussels increased NH4+ concen- trations and the fertihzer increased both NH4+ and Pod3- in the sediment porewater and the overlying water column, but this had only limited effects on eelgrass shoot growth rates and increase in shoot density, which were high overall (up to 75 mm shoot-' d-l, doubling shoot density every 3 mo). In contrast, increased plant density had clear positive effects on shoot growth, area1 expansion of patches and increase in shoot density. These results suggest that nutrient availability is not a major factor in eelgrass patch colonization or survival in the Baltic. Positive interactions among eelgrass shoots appear to be more important than competitive processes, during the early stages of recolonization.