Tracing the origin of green macroalgal blooms based on the large scale spatio-temporal distribution of Ulva microscopic propagules and settled mature Ulva vegetative thalli in coastal regions of the Yellow Sea,China

From 2008 to 2016, massive floating green macroalgal blooms occurred annually during the summer months in the Yellow Sea. The original source of these blooms was traced based on the spatio-temporal distribution and species composition of Ulva microscopic propagules and settled Ulva vegetative thalli monthly from December 2012 to May 2013 in the Yellow Sea. High quantities of Ulva microscopic propagules in both the water column and sediments were found in the Pyropia aquaculture area along the Jiangsu coast before a green macroalgal bloom appeared in the Yellow Sea. The abundance of Ulva microscopic propagules was significantly lower in outer areas compared to in Pyropia aquaculture areas. A molecular phylogenetic analysis suggested that Ulva prolifera microscopic propagules were the dominant microscopic propagules present during the study period. The extremely low biomass of settled Ulva vegetative thalli along the coast indicated that somatic cells of settled Ulva vegetative thalli did not provide a propagule bank for the green macroalgal blooms in the Yellow Sea. The results of this study provide further supporting evidence that the floating green macroalgal blooms originate from green macroalgae attached to Pyropia aquaculture rafts along the Jiangsu coastline of the southern Yellow Sea.     

An integrated study of the temporal and spatial variation in the supply of propagules, recruitment and assemblages of intertidal macroalgae on a wave-exposed rocky coast, Victoria, Australia

Recruitment is known to influence distributions and abundances of benthic marine organisms. It is therefore important to document patterns of variability in recruitment and how these relate to patterns in established assemblages. This study provides an integrated assessment of the temporal and spatial variation in supply and recruitment of propagules and established populations of several macroalgae. Propagules in water samples from two stages of the incoming tide, recruitment to artificial substrata and percentage cover of species established on the shore were recorded every 2 months from December 1994 to October 1995, in two zones of an intertidal, wave-exposed rocky shore. Variability in recruitment was measured at three spatial scales: 10s cm, 100s cm and 100s m. Availability and recruitment of most taxa were greatest between April and August, although many species had available propagules and recruited throughout the year. Temporal variation in the established assemblages was, however, more species-specific. Differences in established assemblages between zones were reflected in differences in availability and recruitment of propagules between zones. Recruitment could not be predicted directly from supply of propagules, but the two processes were linked. For most species, the greatest variation in recruitment occurred at the smallest spatial scale of 10s cm, although there was also considerable large-scale (between site) variation in recruitment of several species. Results indicate that while pre-and post-settlement mortality are likely to influence macroalgal distribution and abundance, the temporal and spatial variability in supply and recruitment of propagules can explain much of the patchiness in macroalgal assemblages.     

The arrival of propagules of marine macroalgae in the intertidal zone

The composition and abundance of macroalgal propagules contained in sea water arriving at intertidal rocky shores was estimated monthly at Pelancura, central Chile, from June to December, 1984. Samples from surface water and from water running off rocky platforms with mixed algal vegetation were cultured in laboratory conditions and examined for development of sporelings. Thirty eight macroalgal entities grew in the cultures, 75% of them with opportunistic life-styles. The total number of sporelings was similar in the surface water of sites separated by 300 m. Marked variability in composition and number of sporelings was, however, observed between samples taken simultaneously at one site and between different months, suggesting patchiness in the dispersal of propagules. Marked differences were established in the ratio run-off water/surface water in the number of sporelings, which could be related to the dispersal shadows of the various taxa. Sporelings of late successional algae showed marked spatial and temporal variations in number. Some taxa of opportunistic algae were characterized by the development of several thousand sporelings per litre of sea water while in others the number of sporelings was two or three orders of magnitude less, suggesting differences in reproductive effort.     

Variations in the dispersal curves of macroalgal propagules from a source

The process of dispersal is critical to marine benthic species (i.e. invertebrates and algae) as a fundamental element of population ecology and a crucial ecological process that maintains the diversity in communities. We simultaneously sampled the abundance of spores inhabiting the water column at different distances from known parent sources at 3 sites along the coast of central Chile. From these data we constructed 258 dispersal curves for common rocky intertidal macroalgae. Only 43.8% of these curves could be predicted by the expected model, which describes the spatial distribution of propagules to be dominated by a larger concentration near the parent individual or “source site”, followed by a marked decrease in abundance with increasing distance. The curves that departed from the expected model (56.2%) were grouped into three curve types, according to the number of propagule abundance maxima observed in space. This work suggests that macroalgal propagule dispersal patterns are more variable than previously thought. The existence of several alternative curves to the expected model, as well as the presence of one to several abundance maxima associated with the differential distribution of propagule patches in the water column, suggests the idea that propagules are released in pulses which can be transported variable distances from the source site.     

Effect of Seasonality on Distribution of Macroalgae in a Stream System （Xin＇an Spring）in Shanxi Province, North China

A survey of the seasonal distribution of macroalgae in a stream system in Shanxi Province, north China, was undertaken from July 2004 to April 2005. The relative abundance and percentage cover of macroalgae, and several environmental factors were monitored along a 20-m stretch at each of four sites, at intervals of approximately three months （one sampling per season）. Several stream conditions were relatively constant over the sampling period （pH, maximum width and maximum depth）, whereas others exhibited a distinct seasonal pattern （water temperature and specific conductance）, and some fluctuated with no discernable seasonal pattern （current velocity and dissolved oxygen）. Forty-two species of macroalgae were found, with a predominance of Chlorophyta （26 species, 61.9%）. Rhodophyts and Charophyta represented the smallest proportion （1 species each, 2.38%）. Six macroalgae species were the most widespread, occurring in all four sampling sites. Twelve species were found at only one site each. In terms of seasonality, eight species occurred throughout the year, whereas 16 species were found in only one season each. The macroalgal community at Xin＇an Spring was species rich relative to other streams. Species richness per sampling site was negatively correlated with pH. Principal component analysis revealed that no single variable had much influence on the macroalgal seasonal dynamics. We calculated Sorensen similarity indices to compare our study with other continent-wide surveys of stream macroalgae, but the similarity indices were all very low. This study also shows that macroalgae in different locations have significant reproductive isolation.     

Effect of Seasonality on Distribution of Macroalgae in a Stream System (Xin'an Spring)in Shanxi Province, North China

A survey of the seasonal distribution of macroalgae in a stream system in Shanxi Province, north China,was undertaken from July 2004 to April 2005. The relative abundance and percentage cover of macroalgae,and several environmental factors were monitored along a 20-m stretch at each of four sites, at intervals of approximately three months (one sampling per season). Several stream conditions were relatively constant over the sampling period (pH, maximum width and maximum depth), whereas others exhibited a distinct seasonal pattern (water temperature and specific conductance), and some fluctuated with no discernable seasonal pattern (current velocity and dissolved oxygen). Forty-two species of macroalgae were found, with a predominance of Chlorophyta (26 species, 61.9%). Rhodophyta and Charophyta represented the smallest proportion (1 species each, 2.38%). Six macroalgae species were the most widespread,occurring in all four sampling sites. Twelve species were found at only one site each. In terms of seasonality,eight species occurred throughout the year, whereas 16 species were found in only one season each. The macroalgal community at Xin'an Spring was species rich relative to other streams. Species richness per sampling site was negatively correlated with pH. Principal component analysis revealed that no single variable had much influence on the macroalgal seasonal dynamics. We calculated Sorensen similarity indices to compare our study with other continent-wide surveys of stream macroalgae, but the similarity indices were all very low. This study also shows that macroalgae in different locations have significant reproductive isolation.     

Hydrodynamic control of filamentous macroalgae in a sub-tropical spring-fed river in Florida,USA

Hydrodynamic properties of rivers and streams are primary controls on the abundance and structure of algal communities. In Florida, USA, filamentous macroalgal proliferation in spring-fed rivers has largely been attributed to nitrogen enrichment; however, changes in hydrodynamics and other potential mechanisms have not been thoroughly evaluated. This study tested whether macroalgal abundance (primarily Lyngbya wollei) was related to hydrodynamics in Gum Slough, a sub-tropical Florida spring-fed river. Flow velocities and macroalgal cover were measured at three transects bimonthly between June 2010 and January 2013, a period during which river discharge fluctuated in response to a drought and a tropical storm. Temporal variation in macroalgal cover was significantly inversely related to flow velocity, excluding fall/winter sampling events when light limitation was likely. The results indicate a velocity threshold below which macroalgal cover increased substantially, and a lower threshold below which complete cover frequently occurred. Local differences in channel morphology and substrate type (i.e., aquatic plant beds) seemed to affect the relationship between macroalgal cover and velocity at each transect. Nitrogen and phosphorus concentrations and other water chemistry parameters were not associated with variation in macroalgal cover. Overall, this study suggests that hydrodynamics can strongly control filamentous macroalgae in Florida spring-fed rivers.     

Environmental and spatial controls of biotic assemblages in a discrete semi‐terrestrial habitat: comparison of organisms with different dispersal abilities sampled in the same plots

Aim The development of metacommunity theory inspired a series of studies exploring the importance of environmental and spatial effects on the composition of biotic assemblages. However, the comparison of different groups of organisms has been hampered by differences in sampling design, spatial scales or the environmental variables involved. Our aim was to test how dispersal ability affects metacommunity structure and associated species distributions by sampling different species groups in the same plots to avoid these problems. Location Western Carpathian Mountains (Europe). Methods In 191 fens we sampled the composition of diatom, bryophyte, vascular plant and mollusc assemblages, water chemistry, and macroclimatic data. We then generated spatial variables covering all relevant spatial scales using analysis of principal coordinates of neighbour matrices (PCNM). We applied the adjusted variation partitioning algorithm to quantify the effects of environment and space. Results Pure effects of water chemistry and space were highly significant for all groups of organisms. Spatial effects were stronger for groups with larger propagules (vascular plants, molluscs) than for those with smaller propagules (diatoms, bryophytes). Assemblages of macroscopic bryophytes were structured slightly less by geography and much more by environment than were those of microscopic diatoms. Vascular plant and mollusc assemblages turned out to be more spatially structured (as compared to diatom and bryophyte assemblages), with small differences between the two groups. Coarse‐scale spatial effects dominated in the bryophyte metacommunity, while in the other groups, including diatoms, finer‐scale effects were also important. Main conclusions Given that our analyses are based on a standardized sampling and analytical framework, our findings provide strong support for the hypothesis that both environmental and spatial variables structure metacommunities of organisms with very different dispersal abilities, including microscopic diatoms. In addition, we show for the first time that the strengths of these effects and their scale dependence may be predicted using important trait differences between organisms, for example differences in propagule size.     

A system for coupled multiple-column separation of proteins

Purification of proteins is commonly a multiple-step process involving size exclusion, ion exchange, affinity, hydrophobic, and other modes of chromatography. In an effort to circumvent the laborious process of collecting the solutes from each column and reintroducing them onto a second column, a valving system is described that directs the samples eluted from a high-performance liquid chromatographic column through a detector with a high-pressure cell into either a second column or into storage loops of a multiloop value. This multiloop value is referred to as a high-pressure fraction collector. After development of the first column is complete, a second solvent can be directed to the second column or high-pressure fraction collector to elute the solutes back through the detector and onto any other column in the system. The process of eluting a sample from a column through a single detector and directing it to the high-pressure fraction collector or any other column in the system may be repeated a number of times. Such valving systems make it possible to chromatograph a single protein component on two or three columns in a short time.     

Differential survival of macroalgae to digestion by intertidal herbivore molluscs

This study evaluates the capacity of algal propagules to survive digestion by seven intertidal herbivore molluscs. Thirty individuals of Littorina peruviana, 36 of Siphonaria lessoni, 35 of Collisella ceciliana, and 25 each of C. zebrina, Chiton granosus, Fissurella crassa and F. limbata were collected at different dates between October, 1983 and April, 1984. About half of the individuals of each species were examined for gut contents and the other half was used to provide faecal pellets for culture. Diet of these grazers ranged from 8 macroalgal species in Collisella zebrina to 17 in Fissurella limbata. In total, macroalgal propagules of 56% of the 27 algal species found in these gut contents survive digestion. There was no positive correlation between diet and number of algal taxa growing in the respective faecal cultures. Survival through Littorina peruviana and Chiton granosus was low (20–30% of the algal species consumed) while it was high (75–83%) through the two species of Collisella. No significant correlation was found between frequency of algae in the gut contents and its frequency in the faecal cultures. Algal capacity to survive digestion was almost exclusively restricted to opportunistic species, especially in the Chlorophyta and Phaeophyta. It seems that survival of algal propagules through the digestive tract of generalist grazers is a rather random phenomenon.     

Variability in nitrogen stable isotope ratios of macroalgae: consequences for the identification of nitrogen sources

In our research, we collected and analyzed numerous macroalgal specimens (738) for isotopic analysis sampled over a year at monthly intervals across 20 sites within the Urías lagoon complex, a typical subtropical coastal ecosystem located in the Gulf of California. We quantified and characterized (chemically and isotopically) the N loads received by Urías throughout a year. We studied the spatial‐temporal variation of the chemical forms and isotopic signals of the available N in the water column, and we monitored in situ different environmental variables and other hydrodynamic parameters. Multiple N sources (e.g., atmospheric, sewage, seafood processing, agriculture and aquaculture effluents) and biogeochemical reactions related to the N cycle (e.g., ammonia volatilization, nitrification and denitrification) co‐occurring across the ecosystem, result in a mixture of chemical species and isotopic compositions of available N in the water column. Increased variability was observed in the δ¹⁵N values of macroalgae (0.41‰–22.67‰). Based on our results, the variation in δ¹⁵N was best explained by spatio‐temporal changes in available N and not necessarily related to the N sources. The variability was also explained by the differences in macroalgal biology among functional groups, species and/or individuals. Although the δ¹⁵N‐macroalgae technique was a useful tool to identify N sources, its application in coastal ecosystems receiving multiple N sources, with changing environmental conditions influencing biogeochemical processes, and high diversity of ephemeral macroalgal species, could be less sensitive and have less predictive power.     

Ecological,Dynamic and Taxonomic Problems Due to <Emphasis Type="Italic">Ludwigia</Emphasis> (Onagraceae) in France

We analyzed meta-community structure of bdelloid rotifers colonizing mosses along an 80 meter section of Rio Valnava in NW Italy. Bdelloid rotifers are small animals living associated with a substratum; colonization in bdelloids can be produced by active animals moving along the riverbed, or by passive dormant propagules, moved by wind. To detect which kind of colonization might be stronger at different spatial scales, we designed a spatially nested sampling experiment at three hierarchical levels: (1) single sample, (2) 10 communities inside each pool, (3) complete section of 10 pools. Assessing species richness and species similarity of communities, and coherence and nestedness of bdelloid meta-communities, we found that different forces may drive species composition at different spatial scales: at the largest scale, colonization of propagules may over-ride direct dispersal between pools, while at the scale of the single pool, differential movements of species give a nested structure to the meta-communities. The number of species increased as the level of analysis increased, even though this study was carried out along only a small stream section.     

Understorey benthic microalgae and their consumers depend on habitat complexity and light in a microtidal coastal ecosystem

Presence of habitat-forming macroalgae is supposed to mitigate effects of altered resources on benthic microalgae and their consumers. In a field experiment in a microtidal area of the Western Baltic Sea, we tested the interactive effects of nutrient enrichment, artificial shading, and habitat complexity on microalgal biomass and diversity as well as invertebrate abundance and richness. Habitat complexity comprised three levels, the presence of macroalgal canopy of Fucus vesiculosus, the presence of macroalgal propagules, and the absence of both (=control). Microalgal biomass (and richness) was significantly reduced by canopy presence (−88%, compared to control) and shading (−42%), with the highest biomass in the absence of both canopy and macroalgal propagules at ambient light. Within the microalgal assemblage, higher biomass was related to lower evenness (higher dominance). Density of two main invertebrate groups (snails and amphipods) strongly increased with canopy presence (on average from 53 to 154 individuals m⁻² stone area for snails, and from 234 to 1203 individuals m⁻² for amphipods) and so did invertebrate richness (from 4.3 to 10.3). Additionally, snail density doubled with increasing light availability. The snail responses to light and canopy were independent, the former relating to higher availability of microalgal prey, the latter to more structure. Microalgal taxon richness and biomass decreased with increasing invertebrate richness and with density of snails and amphipods. Our experiment thus showed that the presence of habitat-forming macroalgae alters biomass and diversity across trophic levels in benthic coastal communities as well as their response to resource manipulations.     

Relationships between macroalgal biomass and microbiological quality of water in a phytotreatment pond

The microbiological quality of wastewater in phytotreatment ponds with foliose macroalgae can be influenced by biofilm formation on thallus surface. This hypothesis was tested with an in situ experiment which was carried out in a pond with Ulva spp. receiving wastewater from a land based fish farm at Piombino (Italy). The total bacterial load (TBL) was determined in the inflowing and outflowing waters and a multifactorial design was employed to investigate the effect of different macroalgal biomass. Microbiological analysis revealed a high TBL in the water column (18.4 ± 7.4 × 10⁸ cells ml⁻¹). TBL of inlet water was significantly correlated with quantity and quality of particulate organic matter (POM) of inflowing water, whereas no correlation was found between TBL and POM in the outlet water. A significant decrease in the POM concentration was detected within macroalgal ponds, due to the mechanical action of thalli which favoured POM sedimentation. Nevertheless, great TBLs were found in the outlet water. These findings suggest that TBL probably depended upon macroalgae. Indeed high bacterial density was found on macroalgal thallus surface (~10⁸ cells cm²). Furthermore, high plate counts of faecal bacteria (faecal enterococci) were determined on thallus surface (~ 40 CFU cm²) and outlet water (11 886 ± 3984 CFU 100 ml⁻¹) supporting the evidence that macroalgae negatively affect the microbiological quality of treated water. Bacterial activities in terms of exoenzymatic rates and secondary production were two folds higher in the water within macroalgal beds, than in the open water. These preliminary results suggest that high macroalgal biomass represents a ‚hot spot’ of bacterial density and activity that may affect microbiological quality of the treated water. Bacterial control of inlet water and management of macroalgal biomass through periodic removal are essential for a more efficient treatment of wastewater in phytotreatment ponds.     

Automatic Monitoring of primary amines in preparative column effluents with fluorescamine.

A system is presented for automatically monitoring primary amines in preparative column effluents by fluorescamine assay. A small portion of column effluent, containing nanograms of protein or picomoles of peptide, is diverted via a sampling valve into the detection system, while the remainder is recovered in a fraction collector.     

Possible influences of a macroalgal bloom in eelgrass beds on fish assemblages in the lower Knysna Estuary,South Africa

The occurrence of a macroalgal bloom at eelgrass (Zostera capensis) sampling sites in the summer of 2014/2015 provided an opportunity to use underwater video cameras to monitor the possible effects of environmental change on fish diversity and abundance in the lower reaches of the Knysna Estuary. A General Linear Model (GLM) showed that there was a significant difference in the abundance of fish before and after an invasion of the sampling sites by the macroalga Ulva lactuca. The eelgrass bed fish abundance was more affected by the macroalgal bloom than the bare substratum fish abundance, with the Ulva impacting negatively on the relative abundance of Mugilidae in the former habitat. The hypothesis that macroalgal invasions have a negative effect on fish diversity and abundance is therefore supported by this preliminary study.     

Propagule banks, herbivory and nutrient supply control population development and dominance patterns in macroalgal blooms

Destructive macroalgal mass blooms threaten estuarine and coastal ecosystems worldwide. We asked which factors regulate macroalgal bloom intensity, distribution and species composition. In field experiments in the Baltic Sea, we analyzed the relative effects of nutrients, herbivores and algal propagule banks on population development and dominance patterns in two co-occurring bloom-forming macroalgae, Enteromorpha intestinalis and Pilayella littoralis . Both species were highly affected by the combined effects of a propagule bank, herbivory and nutrients. The magnitude of effects varied with season. The propagule bank was an important overwintering mechanism for both algae, and allowed for recruitment two months earlier than recruitment via freshly dispersed propagules. This provided a seasonal escape from intense herbivory and nutrient limitation later in the year. Favored by massive recruitment from the propagule bank, Enteromorpha was the superior space occupier in early spring, thereby reducing recruitment of Pilayella . Elimination of the propagule bank and recruitment via freshly dispersed propagules favored Pilayella . Strong and selective herbivory on Enteromorpha supported Pilayella in the presence, but not in the absence of the propagule bank. Nutrient enrichment in summer counteracted herbivore pressure on Enteromorpha , thereby negatively affecting Pilayella . Herbivore and nutrient effects were more pronounced for early life stages than adult algae. These results show that recruitment processes and forces affecting early life stages at the beginning of the vegetation period determine development and dominance patterns of macroalgal blooms. Herbivores naturally suppress blooms but increasing nutrient enrichment can override this important control mechanism. The propagule bank plays a previously unrecognized role for population and community dynamics.     

Implementation of a new index to assess intertidal seaweed communities as bioindicators for the European Water Framework Directory

An index CCO (cover, characteristic species, opportunistic species) has been developed for the implementation of the European Water Framework Directory (WFD) in coastal waters, using intertidal macroalgal communities as bio-indicator (Biological Quality Element). CCO is based on the calculation of three metrics corresponding to the global cover of macroalgal communities (metric 1), the number of characteristic species per topographic level/seaweed community (metric 2) and the cover of opportunistic species (metric 3). The final rating is obtained by pooling the scores of the three metrics. Results are given for 32 sites in 29 water bodies, grouped into four biogeographic regions along the Channel–Atlantic coasts of France. Over the six-year study, most of sites were sampled twice each (every three years). CCO index revealed that 25 coastal water bodies of both the Channel and the Bay of Biscay were in good or high ecological quality status (EQS), whereas only 4 of them were moderate and none in poor to bad status. However, significant differences have been found between sites and between geographic regions, water bodies located in Brittany obtaining the best EQS. No significant change occurred between the three-year sampling sets. A significant correlation has been established between a three-component anthropogenic pressure index and CCO ratings, showing the accuracy of CCO to evaluate the impact of anthropic activities on the structure and development of macroalgal communities as indicator of the ecological quality of coastal water bodies.     

The movement ecology of seagrasses

A movement ecology framework is applied to enhance our understanding of the causes, mechanisms and consequences of movement in seagrasses: marine, clonal, flowering plants. Four life-history stages of seagrasses can move: pollen, sexual propagules, vegetative fragments and the spread of individuals through clonal growth. Movement occurs on the water surface, in the water column, on or in the sediment, via animal vectors and through spreading clones. A capacity for long-distance dispersal and demographic connectivity over multiple timeframes is the novel feature of the movement ecology of seagrasses with significant evolutionary and ecological consequences. The space–time movement footprint of different life-history stages varies. For example, the distance moved by reproductive propagules and vegetative expansion via clonal growth is similar, but the timescales range exponentially, from hours to months or centuries to millennia, respectively. Consequently, environmental factors and key traits that interact to influence movement also operate on vastly different spatial and temporal scales. Six key future research areas have been identified.     

Small-scale spatial structure within patterns of seed dispersal

A large proportion of dispersing propagules land near their maternal plant, even in species that have evolved structures which enhance dispersal. For these propagules, their post-dispersal spatial pattern is likely to reflect the overall shape and scale of the parental plant canopy and, especially in poorly dispersing species, aggregation of propagules on the plant prior to dispersal. Localised patterns within seed shadows are also likely to be affected by secondary movement after dispersal, leading to either more or less small-scale aggregation, depending on the mechanism. Our general aim was to study the small-scale spatial structure within patterns of seed dispersal of Raphanus raphanistrum L. to generate hypotheses about the sequence of processes and events leading to the spatial pattern of dispersal in this species. More specifically, we determined the sizes of small-scale structures within the seed shadows on the ground after dispersal and the extent to which these match the sizes of pre-dispersal aggregations within the parental canopy. Variation in plant size and shape was provided by four levels of inter-specific competition resulting from differing wheat crop densities. Positions of propagules were determined using a three-dimensional digitizer, and the data for each plant were analysed using point pattern analysis. Not surprisingly, larger plants, growing at lower plant density, had larger seed shadows, showing an overall influence of maternal plant size. The pattern of propagules exhibited significant small-scale aggregates, with similar sizes on the plant and on the ground. There was no evidence that aggregation size was greater on the ground or increased with time, but the strength of the aggregation increased with length of time on the ground.