Disturbance decreases genotypic diversity by reducing colonization: Implications for disturbance-diversity feedbacks.

One objective of eco-evolutionary dynamics is to understand how the interplay between ecology and evolution on contemporary timescales contributes to the maintenance of biodiversity. Disturbance is an ecological process that can alter species diversity through both ecological and evolutionary effects on colonization and extinction dynamics. While analogous mechanisms likely operate among genotypes within a population, empirical evidence demonstrating the relationship between disturbance and genotypic diversity remains limited. We experimentally tested how disturbance altered the colonization (gain) and extinction (loss) of genets within a population of the marine angiosperm Zostera marina (eelgrass). In a 2-year field experiment conducted in northern California, we mimicked grazing disturbance by migratory geese by clipping leaves at varying frequencies during the winter months. Surprisingly, we found the greatest rates of new colonization in the absence of disturbance and that clipping had negligible effects on extinction. We hypothesize that genet extinction was not driven by selective mortality from clipping or from any stochastic loss resulting from the reduced shoot densities in clipped plots. We also hypothesize that increased flowering effort and facilitation within and among clones drove the increased colonization of new genets in the undisturbed treatment. This balance between colonization and extinction resulted in a negative relationship between clipping frequency and net changes in genotypic richness. We interpret our results in light of prior work showing that genotypic diversity increased resistance to grazing disturbance. We suggest that both directions of a feedback between disturbance and diversity occur in this system with consequences for the maintenance of eelgrass genotypic diversity.

Response of eelgrass (Zostera marina) to an adjacent Olympia oyster restoration project.

Recent restoration efforts for the native Olympia oyster, Ostrea lurida, are commonly motivated by potential return of oyster-associated ecosystem services, including increased water filtration. The potential impact of such restoration on another species of ecological concern, eelgrass, Zostera marina, is unclear, but has been hypothesized to be positive if oyster filter feeding increases light penetration to eelgrass. For two years after construction of an oyster restoration project, we assessed the response of adjacent eelgrass (impact) compared to control and reference eelgrass beds by monitoring changes in light intensity, eelgrass shoot density, biomass, leaf morphometrics, and epiphyte load. We observed lower light intensity consistently over time, including prior to restoration, near the constructed oyster bed relative to the control and one of the reference locations. We also observed minor variations between control and impact eelgrass morphology and density. However, the changes observed were not outside the range of natural variation expected in this system, based upon comparisons to reference eelgrass beds, nor were they detrimental. This limited impact to eelgrass may be due in part to the incorporation of a buffer distance between the restored oyster bed and the existing eelgrass bed, which may have dampened both positive and negative impacts. These findings provide evidence that Olympia oyster restoration and eelgrass conservation goals can be compatible and occur simultaneously.

The amount of light reaching the leaves in seagrass (Zostera marina) meadows.

Seagrass meadows, and other submerged vegetated habitats, support a wide range of essential ecological services, but the true extents of these services are in many ways still not quantified. One important tool needed to assess and model many of these services is accurate estimations of the systems´ primary productivity. Such productivity estimations require an understanding of the underwater light field, especially regarding the amount of light that actually reaches the plants' photosynthetic tissue. In this study, we tested a simple practical approach to estimate leaf light exposure, relative to incoming light at the canopy, by attaching light sensitive film at different positions on leaves of Zostera marina, eelgrass, in four seagrass meadows composed of different shoot density and at two different depths. We found that the light reaching the leaves decreased linearly down through the canopy. While the upper parts of the leaves received approximately the same level of light (photosynthetic photon flux density, PPFD) as recorded with a PAR meter at the canopy top, the average light that the seagrass leaves were exposed to varied between 40 and 60% of the light on top of the canopy, with an overall average of 48%. We recommend that actual light interception is measured when assessing or modelling light depending processes in submerged vegetation, but if this is not achievable a rough estimation for vegetation similar to Z. marina would be to use a correction factor of 0.5 to compensate for the reduced light due to leaf orientation and internal shading.

Factors controlling the vertical zonation of the intertidal seagrass, Zostera japonica in its native range in the northwestern Pacific.

The desiccation tolerance of the intertidal seagrass Zostera japonica has been demonstrated in a number of studies; however, the factors limiting expansion of intertidal seagrass species into subtidal zones remain controversial. We transplanted Z. japonica shoots from the intermediate intertidal zone into the plots with and without Z. marina shoots in both the lower intertidal and shallow subtidal zones to investigate the factors controlling Z. japonica growth in these zones. Daily photon flux density at the Z. japonica canopy level was attenuated by both water depth and coexisting Z. marina shoots but more strongly by Z. marina shoots than water depth in the transplant plots. The shoot density and biomass of Z. japonica transplants were significantly lower in transplant plots in the subtidal zone than in the lower intertidal zone. Although the photon flux density was significantly lower in transplant plots containing Z. marina shoots, the growth of Z. japonica transplants did not differ significantly between plots with and those without Z. marina shoots. Z. japonica transplants exhibited photoacclimatory responses such as increased shoot height and chlorophyll content under the lower-light conditions, offsetting the reduced light availability so that no significant differences in transplant growth occurred between plots with and those without Z. marina shoots. As the growth of Z. japonica transplants decreased significantly in the subtidal zone, the interactive effects of environmental stresses associated with tidal inundation and reduced light availability may restrict penetration of the intertidal seagrass Z. japonica into the subtidal zone. The persistence of high photosynthetic performance after air exposure and a regular arrangement of the densely overlapped leaves atop wet sediments may be desiccation tolerance mechanisms for Z. japonica in the intertidal zone.

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.

SMRT sequencing of full-length transcriptome of seagrasses Zostera japonica.

Seagrass meadows are among the four most productive marine ecosystems in the world. Zostera japonica (Z. japonica) is the most widely distributed species of seagrass in China. However, there is no reference genome or transcriptome available for Z. japonica, impeding progress in functional genomic and molecular ecology studies in this species. Temperature is the main factor that controls the distribution and growth of seagrass around the world, yet how seagrass responds to heat stress remains poorly understood due to the lack of genomic and transcriptomic data. In this study, we applied a combination of second- and third-generation sequencing technologies to sequence full-length transcriptomes of Z. japonica. In total, we obtained 58,134 uniform transcripts, which included 46,070 high-quality full-length transcript sequences. We identified 15,411 simple sequence repeats, 258 long non-coding RNAs and 28,038 open reading frames. Exposure to heat elicited a complex transcriptional response in genes involved in posttranslational modification, protein turnover and chaperones. Overall, our study provides the first large-scale full-length trascriptome in Zostera japonica, allowing for structural, functional and comparative genomics studies in this important seagrass species. Although previous studies have focused specifically on heat shock proteins, we found that examination of other heat stress related genes is important for studying response to heat stress in seagrass. This study provides a genetic resource for the discovery of genes related to heat stress tolerance in this species. Our transcriptome can be further utilized in future studies to understand the molecular adaptation to heat stress in Zostera japonica.

Estimation of a whole plant Q10 to assess seagrass productivity during temperature shifts.

Through respiration and photosynthesis, seagrass meadows contribute greatly to carbon and oxygen fluxes in shallow coastal waters. There is increasing concern about how shallow-water primary producers will react to a near-future climate scenario with increased temperature variation. When modelling primary productivity under high temperature variability, Q10 values are commonly used to predict rate changes depending on biophysical factors. Q10 values are often assumed to be constant and around 2.0 (i.e. a doubling of the rate with a temperature increase of 10 °C). We aimed to establish how the gas exchange of seagrass (Zostera marina) tissues at various maturity stages would respond over a broad range of temperatures. Seagrass shoot maturity stage clearly affected respiration and apparent photosynthesis, and the Q10 results indicated a skewed balance between the two processes, with a higher photosynthetic Q10 during periods of elevated temperatures. When estimating whole-plant Q10 in a realistic maximal temperature range, we found that the overall response of a seagrass plant's net O2 exchange balance can be as much as three to four times higher than under ambient temperatures. Our findings indicate that plant tissue age and temperature should be considered when assessing and modelling carbon and oxygen fluctuations in vegetated coastal areas.

The influence of hydrodynamics and ecosystem engineers on eelgrass seed trapping.

Propagule dispersal is an integral part of the life cycle of seagrasses; important for colonising unvegetated areas and increasing their spatial distribution. However, to understand recruitment success, seed dispersal and survival in habitats of different complexity remains to be quantified. We tested the single and synergistic effects of three commonly distributed ecosystem engineers-eelgrass (Zostera marina), oysters (Magellana gigas) and blue mussels (Mytilus edulis)-on trapping of Z. marina seeds in a hydraulic flume under currents. Our results suggest that seed retention increases with habitat complexity and further reveal insights into the underlying mechanisms. In eelgrass canopy, trapping occurred mostly through direct blocking of a seed's pathway, while trapping in bivalve patches was mainly related to altered hydrodynamics in the lee side, i.e. behind each specimen. With increasing flow velocity (24-30 cm s-1 in eelgrass canopy, 18-30 cm s-1 in bivalve patches), modifications of the sediment surface through increased turbulence and erosive processes became more important and resulted in high seed trapping rates. Furthermore, we show that while monospecific patches of seagrass and bivalves had different trapping optima depending on flow velocities, intermixing resulted in consistently high trapping rates throughout the investigated hydrodynamic gradient. Our results highlight the importance of positive interactions among ecosystem engineers for seed retention and patch emergence in eelgrass.

Effects of salinity and temperature on seed germination and seedling establishment in the endangered seagrass Zostera japonica Asch. & Graebn. in northern China.

Seagrass meadows are recognized as critical and among the most vulnerable habitats on the planet. As a worldwide concern, there is an urgent need to develop techniques to restore and preserve these vital coastal ecosystems due to their alarming decline rate. To effectively preserve and restore seagrasses, more research is required on the germination ecology of seeds. The seagrass Zostera japonica Asch. & Graebn is an endangered species in its native range, the Northwestern Pacific Coast. The present study investigated the germination and seedling establishment in Z. japonica seeds (collected from northern China) under different temperature and salinity conditions to explore suitable seed germination and establishment conditions, as well as the seedling formation process. Results showed that salinity had a more significant influence on seed germination rate. Germination rate decreased with an increase in salinity, and seeds did not germinate when salinity was higher than 40 psu. Temperature was more likely to influence germination speed, which increased with an increase in temperature, with high germination rates and the most rapid germination speed observed at 30 °C. Therefore, the optimal culture conditions were 10 psu salinity at 30 °C for germination and 10-20 psu salinity at 20 °C for seedling establishment, with a seed germination rate of 45.6% after 6 days of germination culture and a seedling establishment rate of 14.3% after 6 days of seedling culture, respectively. A new seedling raising method with low salinity (5 psu) germination and high salinity (20 psu) seedling establishment was proposed and a flow chart of seedling formation of Z. japonica was created. The results provide insight into the seed germination and seedling establishment in Z. japonica, and will facilitate future large-scale seedling culture and field restoration activities for this seagrass species.

Examination of the Effects of Curvature in Geometrical Space on Accuracy of Scaling Derived Projections of Plant Biomass Units: Applications to the Assessment of Average Leaf Biomass in Eelgrass Shoots.

Conservation of eelgrass relies on transplants and evaluation of success depends on nondestructive measurements of average leaf biomass in shoots among other variables. Allometric proxies offer a convenient way to assessments. Identifying surrogates via log transformation and linear regression can set biased results. Views conceive this approach to be meaningful, asserting that curvature in geometrical space explains bias. Inappropriateness of correction factor of retransformation bias could also explain inconsistencies. Accounting for nonlinearity of the log transformed response relied on a generalized allometric model. Scaling parameters depend continuously on the descriptor. Joining correction factor is conceived as the partial sum of series expansion of mean retransformed residuals leading to highest reproducibility strength. Fits of particular characterizations of the generalized curvature model conveyed outstanding reproducibility of average eelgrass leaf biomass in shoots. Although nonlinear heteroscedastic regression resulted also to be suitable, only log transformation approaches can unmask a size related differentiation in growth form of the leaf. Generally, whenever structure of regression error is undetermined, choosing a suitable form of retransformation correction factor becomes elusive. Compared to customary nonparametric characterizations of this correction factor, present form proved more efficient. We expect that offered generalized allometric model along with proposed correction factor form provides a suitable analytical arrangement for the general settings of allometric examination.

A unique meadow of the marine angiosperm Zostera japonica, covering a large area in the turbid intertidal Yellow River Delta, China.

Marine submerged aquatic angiosperms (seagrasses) are declining globally. The species Zostera japonica Asch. & Graebn. is endangered in its native range in Asia, but has been successfully introduced to North America. A large area (1031.8 ha) of Z. japonica meadow has recently been discovered in the intertidal zone of Yellow River Delta, China. This seagrass occurs along both sides of the river mouth, forming dense meadows in turbid water conditions. Seasonal investigations over two years were conducted to examine the distribution, biomass, seed reproduction, seed bank, and population recruitment of the seagrass meadows at three sites in the intertidal zone. The meadows generally showed relatively high coverage, biomass, reproductive effort, and seed production in August. The seed bank was found to be large and contributed to population recruitment. There were significant inter-annual variations overall, and at individual sites. These variations are likely due to winter temperatures, which determine the abundance of overwintering shoots and seedling success. Differences in micro-topography may also play a role in producing variations in seedling success between sites. Microsatellite analysis revealed a high genetic exchange between the two sides of the river mouth. The results indicate that the seagrass bed in the Yellow River Delta shallow waters is in good condition, which can be attributed to its location within a national nature reserve. Establishment of protected areas might act as an effective way to mitigate the anthropogenic disturbance, conserve the seagrass meadows, and then enhance critical ecosystem functions.

Local patchiness of macrobenthic faunal abundance displays homogeneity across the disparate seagrass systems of an estuarine bay.

Spatial variation in the degree of local patchiness of macrobenthic assemblage abundance was assessed across the 16 km2 warm-temperate Knysna estuarine bay (South Africa) where the seagrass Zostera (Zosterella) capensis grows under a broad spectrum of environmental conditions and supports invertebrate assemblages at a wide range of local density (<2000->320000 ind. m-2). Macrobenthic assemblage abundance at all 27 representative sites examined displayed low-level but highly-significant spatial patchiness (mean Lloyd's index, IP = 1.148). Except at high tidal levels, however, the magnitude of this local patchiness did not vary statistically across the system (CV 4.3%) regardless of assemblage abundance, location or species composition. Patchinesses well within ±1 standard deviation of Knysna's value also characterise an equivalent Z. (Zosterella) capricorni assemblage in subtropical Queensland (IP 1.169) and another, Z. (Zosterella) noltei, assemblage in cool-temperate England (IP 1.135), suggesting that at local scales intertidal dwarf-eelgrass macrobenthic abundance displays a characteristic level of patchiness.

Seed size and burial depth influence Zostera marina L. (eelgrass) seed survival, seedling emergence and initial seedling biomass development.

Seed burial in the sediment is critical for successful seedling establishment in seagrasses because it protects from predation and dispersal into unsuitable sites, and it may enhance germination by exposing the seeds to suitable germination stimuli. However, relatively little is known about the fate of buried seeds and their ability to emerge from greater depths. The goal of this study was to determine seed survival in the sediment, seedling emergence success and initial seedling biomass of Zostera marina in relation to burial depth and to evaluate if large seeds, having larger energy reserves, are more tolerant to burial than small seeds. Seeds from a perennial Z. marina population were buried at 7 different sediment depths (0.1-8 cm), and seeds sorted by size (large and small) were buried at depths of 2, 4 and 6 cm in outdoor mesocosms. Total seedling emergence after 2 months was significantly affected by seed burial depth, with maximum values in the top 2 cm of the sediment (48.1-56.7% of planted seeds), and a marked decline below 4 cm depth to only 5% seedling emergence at the deepest burial depth of 8 cm. Moreover, seeds had shorter time to emergence from shallow compared to deep burial depths. At all burial depths, a small fraction of seeds (<10%) died after germination but before emerging, and 15-30% remained viable after 6 months. Seed mortality was the major limitation to seedling recruitment from the deeper burial depths. The effect of seed size on seedling emergence success and time was not clear, but heavier seeds displayed greater longevity and gave rise to seedlings of significantly higher biomass, indicating that the mobilization of metabolic reserves may be important during initial seedling development.

The distribution of large floating seagrass (Zostera marina) aggregations in northern temperate zones of Bohai Bay in the Bohai Sea, China.

Seagrass meadows (Zostera marina) are important coastal ecosystems with high levels of productivity and biodiversity. They often have high biomass turnover and are susceptible to dislodgment, leading to export of biomass from seagrass beds. In September 2016, two cruises covering a total area of 52 km x 15 km (38°57'1.14"-39° 0'41.28" N, 118°45'23.22"-118°47'6.96" E) found floating Z. marina aggregations along 13 km of the transect in the northernmost area of Bohai Bay, in the Bohai Sea, China. This floating seagrass was 6.3-13.4 km northeast (offshore) of the Caofeidian seagrass bed, which is a large (10 km2) seagrass bed discovered in 2015 in the Bohai Sea, China. The modal length of floating intact shoots (from meristem to longest leaf tip) matched samples from the Caofeidian seagrass bed. The dominant individuals lengths were 40-50 cm, with less than 5% of the total number of individuals found in larger size categories (80-90 and 90-100 cm). We concluded that they originated from the nearby Caofeidian seagrass meadows.

Physiological responses of eelgrass (Zostera marina) to ambient stresses such as herbicide, insufficient light, and high water temperature.

This study aimed to elucidate the biological responses of eelgrass (Zostera marina) to artificially induced stresses such as herbicide (Irgarol 1051, Irg) exposure, insufficient light, and high water temperature (27 ± 1.0 °C) by evaluating growth inhibition, photosynthetic activity, and metabolomic profiles. After 14 days, all treatments inhibited growth, but photosynthetic activity was only reduced in the Irg-exposed group. In the Irg-exposed and insufficient light groups, the metabolomic profiles were characterized by decreased levels of sugar (sucrose) and increased levels of amino acids such as glutamine, glycine, and leucine. Biochemical and ultrastructural analyses revealed that the loss of sugar-derived metabolic energy was compensated for by energy generated during autophagic protein degradation. Furthermore, the level of myo-inositol, which has various biological roles and participates in several cellular processes such as cell wall synthesis, stress response, and mineral nutrient storage, was markedly increased in the Irg-exposed and insufficient light groups. A combination of metabolomic analysis with other analyses such as measurement of photosynthetic activity might further elucidate the response of eelgrass to ambient stresses in the natural environment.

Evidence for 'critical slowing down' in seagrass: a stress gradient experiment at the southern limit of its range.

The theory of critical slowing down, i.e. the increasing recovery times of complex systems close to tipping points, has been proposed as an early warning signal for collapse. Empirical evidence for the reality of such warning signals is still rare in ecology. We studied this on Zostera noltii intertidal seagrass meadows at their southern range limit, the Banc d'Arguin, Mauritania. We analyse the environmental covariates of recovery rates using structural equation modelling (SEM), based on an experiment in which we assessed whether recovery after disturbances (i.e. seagrass & infauna removal) depends on stress intensity (increasing with elevation) and disturbance patch size (1 m2 vs. 9 m2). The SEM analyses revealed that higher biofilm density and sediment accretion best explained seagrass recovery rates. Experimental disturbances were followed by slow rates of recovery, regrowth occurring mainly in the coolest months of the year. Macrofauna recolonisation lagged behind seagrass recovery. Overall, the recovery rate was six times slower in the high intertidal zone than in the low zone. The large disturbances in the low zone recovered faster than the small ones in the high zone. This provides empirical evidence for critical slowing down with increasing desiccation stress in an intertidal seagrass system.

Expanding Greenland seagrass meadows contribute new sediment carbon sinks.

The loss of natural carbon sinks, such as seagrass meadows, contributes to grenhouse gas emissions and, thus, global warming. Whereas seagrass meadows are declining in temperate and tropical regions, they are expected to expand into the Arctic with future warming. Using paleoreconstruction of carbon burial and sources of organic carbon to shallow coastal sediments of three Greenland seagrass (Zostera marina) meadows of contrasting density and age, we test the hypothesis that Arctic seagrass meadows are expanding along with the associated sediment carbon sinks. We show that sediments accreted before 1900 were highly 13C depleted, indicative of low inputs of seagrass carbon, whereas from 1940's to present carbon burial rates increased greatly and sediment carbon stocks were largely enriched with seagrass material. Currently, the increase of seagrass carbon inputs to sediments of lush and dense meadows (Kapisillit and Ameralik) was 2.6 fold larger than that of sparse meadows with low biomass (Kobbefjord). Our results demonstrate an increasing important role of Arctic seagrass meadows in supporting sediment carbon sinks, likely to be enhanced with future Arctic warming.

Differentiation in fitness-related traits in response to elevated temperatures between leading and trailing edge populations of marine macrophytes.

The nature of species distribution boundaries is a key subject in ecology and evolution. Edge populations are potentially more exposed to climate-related environmental pressures. Despite research efforts, little is known about variability in fitness-related traits in leading (i.e., colder, high latitude) versus trailing (i.e., warmer, low latitude) edge populations. We tested whether the resilience, i.e. the resistance and recovery, of key traits differs between a distributional cold (Greenland) and warm (Portugal) range edge population of two foundation marine macrophytes, the intertidal macroalga Fucus vesiculosus and the subtidal seagrass Zostera marina. The resistance and recovery of edge populations to elevated seawater temperatures was compared under common experimental conditions using photosynthetic efficiency and expression of heat shock proteins (HSP). Cold and warm edge populations differed in their response, but this was species specific. The warm edge population of F. vesiculosus showed higher thermal resistance and recovery whereas the cold leading edge was less tolerant. The opposite was observed in Z. marina, with reduced recovery at the warm edge, while the cold edge was not markedly affected by warming. Our results confirm that differentiation of thermal stress responses can occur between leading and trailing edges, but such responses depend on local population traits and are thus not predictable just based on thermal pressures.

Changes in benthic community structure and sediment characteristics after natural recolonisation of the seagrass Zostera muelleri.

Macrofauna are important contributors to estuarine ecosystem services within and outside of seagrass beds. Here we documented the natural recolonisation of a temperate seagrass (Zostera muelleri) community over 15 years in an urban estuary (Waitemata Harbour, North Island, New Zealand). We also investigated the change in macrofaunal communities in relation to seagrass cover over time, from transition from bare sandflat to seagrass. Colonisation by seagrass was associated with an increase in macrofaunal species diversity (from an average of 32 species per core in 2001 to 46 species per core in 2015) and abundance (from 482 to 2273 individuals per core), as well as an increase in sediment mud (from 4.09% to 12.37%) and organic matter content (from 0.90% to 1.41%). The most abundant species within both seagrass and adjacent unvegetated sandflat were similar, the polychaetes Heteromastus filiformis, Aricidea sp., and Prionospio aucklandica, and the amphipod Paracalliope novizealandiae. The difference in macrofaunal community structure between seagrass and unvegetated sandflat was primarily associated with higher abundance of P. novizealandiae and lower abundance of Pseudopolydora sp. in seagrass. A successional effect was observed in macrofaunal communities over the 15 years following seagrass expansion, primarily associated with an increase in the abundance of Aricidea sp., H. filiformis, and P. novizealandiae, and a reduction in the abundance of the bivalve Linucula hartvigiana. This study is the first to document long-term changes in seagrass and their associated communities during a natural recolonisation event, providing insight into timeframes required both for the regrowth of a seagrass meadow from initial colonisation of individual patches, as well as the trajectories and timeframes of change from a sandflat to a seagrass-associated macrofaunal community. This research enhances our understanding of how changes in seagrass distributions due to seagrass loss or restoration may affect macrofaunal community composition and ultimately ecosystem function.

Individual-based genetic analyses support asexual hydrochory dispersal in Zostera noltei.

Dispersal beyond the local patch in clonal plants was typically thought to result from sexual reproduction via seed dispersal. However, evidence for the separation, transport by water, and re-establishment of asexual propagules (asexual hydrochory) is mounting suggesting other important means of dispersal in aquatic plants. Using an unprecedented sampling size and microsatellite genetic identification, we describe the distribution of seagrass clones along tens of km within a coastal lagoon in Southern Portugal. Our spatially explicit individual-based sampling design covered 84 km2 and collected 3 185 Zostera noltei ramets from 803 sites. We estimated clone age, assuming rhizome elongation as the only mechanism of clone spread, and contrasted it with paleo-oceanographic sea level change. We also studied the association between a source of disturbance and the location of large clones. A total of 16 clones were sampled more than 10 times and the most abundant one was sampled 59 times. The largest distance between two samples from the same clone was 26.4 km and a total of 58 and 10 clones were sampled across more than 2 and 10 km, respectively. The number of extremely large clone sizes, and their old ages when assuming the rhizome elongation as the single causal mechanism, suggests other processes are behind the span of these clones. We discuss how the dispersal of vegetative fragments in a stepping-stone manner might have produced this pattern. We found higher probabilities to sample large clones away from the lagoon inlet, considered a source of disturbance. This study corroborates previous experiments on the success of transport and re-establishment of asexual fragments and supports the hypothesis that asexual hydrochory is responsible for the extent of these clones.