Ecology of freshwater shore zones

Freshwater shore zones are among the most ecologically valuable parts of the planet, but have been heavily damaged by human activities. Because the management and rehabilitation of freshwater shore zones could be improved by better use of ecological knowledge, we summarize here what is known about their ecological functioning. Shore zones are complexes of habitats that support high biodiversity, which is enhanced by high physical complexity and connectivity. Shore zones dissipate large amounts of physical energy, can receive and process extraordinarily high inputs of autochthonous and allochthonous organic matter, and are sites of intensive nutrient cycling. Interactions between organic matter inputs (including wood), physical energy, and the biota are especially important. In general, the ecological character of shore zone ecosystems is set by inputs of physical energy, geologic (or anthropogenic) structure, the hydrologic regime, nutrient inputs, the biota, and climate. Humans have affected freshwater shore zones by laterally compressing and stabilizing the shore zone, changing hydrologic regimes, shortening and simplifying shorelines, hardening shorelines, tidying shore zones, increasing inputs of physical energy that impinge on shore zones, pollution, recreational activities, resource extraction, introducing alien species, changing climate, and intensive development in the shore zone. Systems to guide management and restoration by quantifying ecological services provided by shore zones and balancing multiple (and sometimes conflicting) values are relatively recent and imperfect. We close by identifying leading challenges for shore zone ecology and management.     

European lake shores in danger — concepts for a sustainable development

In Europe, when small (> 0.01 km²) lakes are considered, approximately 500,000 natural lakes occur. This considerable number points to the fact that lakeshore habitats and ecosystems are of significant importance to the total biodiversity of European landscapes, not only because of their expanse, but also because they are ecotones between land and water, which attract many kinds of wildlife, economical, cultural and recreational uses and human settlement. These very considerable stretches of shore are not currently registered, mapped or evaluated anywhere.Apart from providing habitat, the littoral biocoenosis performs a string of additional functions in the ecosystem, of which several are also of great importance to people, such as self purification, buffer zone, erosion protection and recreation scenery. In particular, the recreational function provides a main economic base for many lake areas and even whole countries in Europe. However, human interests have resulted in lake shore deterioration, such that many European lakes are now bare natural shores or retain only relics of them. Apart from the loss of the ecological functions, this also leads to a substantial loss of economic benefits. This precipitates the need for a responsible management, which can be done only on the base of a sound assessment method and a continuous monitoring of the status of lake shore areas. On a European scale, the European Water Framework Directive (WFD, 2000/60/EC) and the Habitat Directive (92/43/EEC) provide the frame for the assessment and monitoring of the status of littoral habitats of lakes. The WFD focuses on entire surface water bodies, including their associated wetlands under influence of their natural water-level fluctuations. However, the provided set of quality elements has to be adapted for an approach specific to lake shores. Apart from quality elements indicating the natural spatial pre-requisites, biocoenotic diversity and integrity, and ecosystem function and dynamics, also quality elements representing the human pressure of land use and the social and economic value of the lake shore zone should be included in an integrated lakeshore assessment and monitoring concept.However, the application of such an integrated quality assessment scheme requires an integrated administrative counterpart, just as the WFD requires for water management aspects on a catchment scale. Only by overcoming the splitting of competence among a variety of authorities and planning corporations can an integrated approach to sustainable shore development be translated into action.     

The use of macroalgal species richness and composition on intertidal rocky seashores in the assessment of ecological quality under the European Water Framework Directive

The EC Water Framework Directive (WFD) suggests using abundance and species composition of intertidal seaweed communities for ecological quality classification of rocky seashores. There are two difficulties with this. According to WFD all sensitive species should be present on a shore. There is no accepted list of sensitive seaweed species and those which may be sensitive in one location may not be so in another. Second, natural successions can result in very large abundance changes of common species, e.g. from almost completely fucoid-dominated shores to almost totally barnacle-dominated shores, without any change in ecological quality. Studies have shown that numerical species richness, not the list of actual species present, is broadly constant in the absence of disturbance. The ephemeral species, possibly the sensitive members of the community, change regularly in such a way as to conserve species richness. It is proposed that species richness on a defined length of shore be used as a criterion of ecological quality. A database of species found on over 300 shores in the British Isles, under strictly controlled sampling conditions, has given ranges of values of species richness to be expected and has allowed for variations in these values due to sub-habitat variability, wave exposure and turbidity to be factored in. A major problem in applying such a tool is the lack of expertise of many workers in critical identification of seaweed species. A reduced species list has been extracted from the database using species commonly present and identifiable with reasonable certainty. A numerical index of ecological quality is proposed based on scores for various aspects of the physical nature of the habitat combined with a score for species richness which may be based on the reduced species list. The scoring system also uses further aspects of community structure, such as ecological status groups and the proportions of rhodophyta, chlorophyta and opportunist species. For this system to be effective there has to be close control of the way in which sampling is carried out to ensure a uniform level of thoroughness.     

Contrasting effects of cross-ecosystem subsidies and predation on benthic invertebrates in two Pacific coastal streams

Cross-ecosystem subsidies, such as terrestrial invertebrates and leaf litter falling into water as resources for aquatic communities, can vary across environmental gradients. We examined whether the effect of terrestrial subsidy inputs on benthic invertebrates was mediated by resident coastal cutthroat trout (Oncorhynchus clarki) in two representative streams. We experimentally manipulated the input rates (reduced, ambient) of terrestrial subsidies (terrestrial invertebrates and leaf litter) as well as the presence or absence of cutthroat trout in the two streams. The hypothesis that the reduction of terrestrial subsidies to the stream influences benthic invertebrate assemblages was supported by experimental results. The treatments of terrestrial subsidy reduction and cutthroat trout presence had a significant negative effect on benthic invertebrate community biomass and shredder biomass in East Creek with high natural terrestrial subsidy input and small amount of large wood in channel. In contrast, results from Spring Creek with low subsidy input and large amount of large wood in channel showed that only the terrestrial subsidy reduction significantly reduced the biomass of shredders. The effects of the terrestrial subsidy input and trout predation on benthic invertebrate communities varied between the two streams. Our results indicate that a subsidy effect on benthic communities can vary between nearby streams differing in canopy and habitats. This study, with the major finding of highly context-dependent effects of spatial subsidies, suggests that the interplay of resource subsidies and predators on invertebrate community assemblages can be site-specific and context-dependent on habitat features.     

Identifying and mitigating dam-induced declines in river health:Three case studies from the western United States

River health can be defined as the degree to which riverine energy source,water quality,flow regime, habitat and biota match the natural conditions.In a healthy river,physical process and form remain actively connected and able to mutually adjust,and biological communities have natural levels of diversity and are resilient to environmental stress.Both physical diversity and biodiversity influence river health.Physical diversity is governed by hydrology,hydraulics,and substrate,as reflected in the geometry of the river channel and adjacent floodplain,which create habitat for aquatic and riparian organisms.Biodiversity is governed by biological processes such as competition and predation,but biodiversity also reflects the diversity,abundance and stability of habitat,as well as connectivity. Connectivity within a river corridor includes longitudinal,lateral,and vertical dimensions.River health declines as any of these interacting components is compromised by human activities.The cumulative effect of dams and other human alterations of rivers has been primarily to directly reduce physical diversity and connectivity,which indirectly reduces biodiversity.Restoration and maintenance of physical diversity and biodiversity on rivers affected by dams requires quantifying relations between the driver variables of flow and sediment supply,and the response variables of habitat,connectivity,and biological communities.These relations can take the form of thresholds(e.g., entrainment of streambed sediment) or response curves(e.g.,fish biomass versus extent and duration of floodplain inundation).I use examples from Wyoming,Colorado,and Arizona in the western United States to illustrate how to quantify relations between driver and response variables on rivers affected by dams.     

Elemental and stable isotopic constraints on river influence and patterns of nitrogen cycling and biological productivity in Hudson Bay

Elemental (carbon and nitrogen) ratios and stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) are examined in sediments and suspended particulate matter from Hudson Bay to study the influence of river inputs and autochthonous production on organic matter distribution. River-derived particulate organic matter (POM) is heterogeneous, nitrogen-poor and isotopically depleted, consistent with expectations for OM derived from terrestrial C3 vascular plant sources, and distinct from marine OM sources. Both δ¹³C and C/N source signatures seem to be transmitted to sediments with little or no modification, therefore making good tracers for terrigenous OM in Hudson Bay. They suggest progressively larger contributions from marine sources with distance from shore and secondarily from south to north, which broadly corresponds to the distribution of river inputs to Hudson Bay. Processes other than mixing of marine and terrigenous OM influence sedimentary δ¹⁵N values, including variability in the δ¹⁵N of phytoplankton in the Bay's surface waters due to differences in relative nitrate utilization, and post-production processes, which bring about an apparently constant ¹⁵N-enrichment between surface waters and underlying sediments. Variability in the δ¹⁵N of phytoplankton in the Bay's surface waters, in contrast, seems to be organized spatially with a pattern that suggests an inshore–offshore difference in surface water nitrogen conditions (open- vs. closed-system) and hence the δ¹⁵N value of phytoplankton. The δ¹⁵N patterns, supported by a simple nitrate box-model budget, suggest that in inshore regions of Hudson Bay, upwelling of deep, nutrient-rich waters replenishes surface nitrate, resulting in ‘open system’ conditions which tend to maintain nitrate δ¹⁵N at low and constant values, and these values are reflected in the sinking detritus. River inflow, which is constrained to inshore regions of Hudson Bay, appears to be a relatively minor source of nitrate compared to upwelling of deep waters. However, river inflow may contribute indirectly to enhanced inshore nutrient supply by supporting large-scale estuarine circulation and consequently entrainment and upwelling of deep water in this area. In contrast to previous proposals that Hudson Bay is oligotrophic because it receives too much fresh water (Dunbar, 1993), our results support most of the primary production being organized around the margin of the Bay, where river flow is constrained.     

Aquatic prey subsidies to riparian spiders in a stream with different land use types

Land use related habitat degradation in freshwater ecosystems has considerably increased over the past decades, resulting in effects on the aquatic and the riparian communities. Previous studies, mainly in undisturbed systems, have shown that aquatic emergent insects contribute substantially to the diet of riparian predators. To evaluate the effect of land use on aquatic prey subsidies of riparian spiders, we performed a longitudinal study from June to August 2012 along a first order stream (Rhineland-Palatinate, Germany) covering three land use types: forest, meadow and vineyard. We determined the contribution of aquatic and terrestrial resources to the diet of web-weaving (Tetragnathidae spp.) and ground-dwelling (Pardosa sp.) riparian spiders using stable isotope analyses of aquatic emergent insects and terrestrial arthropods. The contribution of aquatic and terrestrial sources differed between Tetragnathidae spp. and Pardosa sp. as well as among land use types. Tetragnathidae spp. consumed 80–100% of aquatic insects in the meadows and 45–65% in the forest and vineyards. Pardosa sp. consumed 5–15% of aquatic insects in the forest, whereas the proportions of aquatic and terrestrial sources were approximately 50% in the meadow and vineyard. Thus, aquatic emergent insects are an important subsidy to riparian spiders and land use is likely to affect the proportion of aquatic sources in the spider diet.     

淀山湖水生维管束植物群落研究

论述了淀山湖水生维管束植物的种类组成、分布、群落类型、群落的数量特征及演替。结果是:（1）湖中分布有26种水生维管束植物,隶属16科,21属;其优势种为苦草、菹草、马来眼子菜、芦苇等。（2）湖中水生植被呈不规则环带状分布,并可划分为挺水、浮叶、沉水三个植物带,12个主要水生维管束植物群落。（3）水生维管束植物的重要值以苦草最大,茨藻属植物最小。（4）淀山湖水生维管束植物群落有由沉水型向浮叶型、挺水型或漂浮型过渡的趋势。     

长江中下游湖泊大型底栖动物群落结构及多样性

2007年10月至2008年4月,对长江中下游地区四种不同类型(草型、天然养殖、施肥养殖以及城市湖泊)的10个湖泊的大型底栖动物群落结构和多样性进行研究,并分析其与水体营养状态之间的关系.研究结果表明,不同类型湖泊底栖动物的密度、生物量、多样性及特征种类均存在显著差异.草型湖泊具有最高的生物量和多样性,但密度最低,其特征种类为腹足纲动物.天然养殖湖泊生物量也较高,物种多样性处于中间水平,特征种类为河蚬、寡鳃齿吻沙蚕及苏氏尾鳃蚓.施肥养殖湖泊和城市湖泊底栖动物密度较高,并呈现出最低的物种多样性,特征种类主要是耐污能力较强的颤蚓类和摇蚊科幼虫.相关性分析表明湖泊营养状态指数与底栖动物密度呈显著正相关,而与生物量呈显著负相关,说明随着营养水平的增加,底栖动物群落逐渐被小个体的耐污种类所主导.Margalef丰富度指数及Pielou均匀度指数与营养状态指数亦呈显著负相关,反映目前长江中下游湖泊随营养水平增加底栖动物群落趋于简单化的演替趋势.     

鄱阳湖湿地中低滩典型植物群落的生物多样性及影响因子

周期性水文节律是影响洪泛湖泊洲滩湿地植物群落物种组成与生物多样性的重要驱动力。本研究以鄱阳湖中低滩典型植物群落为研究对象,通过实地样方调查与统计方法分析植物群落的生物多样性格局及其关键环境因子。结果表明：鄱阳湖中低滩代表性群落分别是灰化薹草群落和虉草群落;植物群落的Shannon-Wiener多样性指数均值为1.5,Pielou均匀度指数均值为0.55,生物多样性与物种均匀度偏低。虉草群落的均匀度、多样性指数高于灰化薹草群落,但二者物种丰富度差异不明显。土壤铵态氮、总磷及土壤含水量和高程是影响植物群落结构与生物多样性的关键因子,其中灰化薹草的物种多样性与土壤铵态氮高度相关,而虉草的均匀度则与土壤总磷关系更密切。此外,鄱阳湖中低滩环境下,灰化薹草群落的均匀度随土壤总磷含量的升高呈现U型响应曲线,虉草则是倒U型,这表明,鄱阳湖中低滩环境下,灰化薹草群落和虉草群落生物多样性对单个、同一的环境因子存在不同的适应机制。     

Impact of reservoirs and channelization on lowland river macroinvertebrates: A case study from Central Europe

In order to assess and compare the ecological impacts of channelization and shallow lowland reservoirs, macroinvertebrate communities of a lowland metapotamal river below reservoirs with epilimnial release were studied. The study was carried out in the Dyje River (Czech Republic) at five sites located from 1.5 to 22.5 km downstream of the reservoir outfall. The five sites differed in the degree of channel modification from natural muddy banks to riprap regulation. Seven samples were collected during the years 1998 and 1999 at each site using a semiquantitative method. The data were processed using multivariate analyses and methods for assessing the ecological and functional structure of communities. Altogether, 261 species of benthic macroinvertebrates were recorded including several rare and threatened taxa. Based on the results of principal component analysis (PCA), most of the variability within the species data (the first PCA axis) was explained by the degree of channel modification, from natural muddy banks with aquatic vegetation to a man-made riprap. The second axis was strongly correlated with current velocity. The sites differed in species richness, total abundances, proportion of individual functional feeding groups, pattern of the distribution of the current preference groups, and values of several biotic indexes, all of which also corresponded to the degree of channel modification. Thus, the morphological man-made modifications of the river channel were found to be the main factor affecting lowland river macroinvertebrates and their biodiversity. Our results suggest that the biggest threat to benthic macroinvertebrate diversity of lowland rivers comes from channelization. The impact of reservoirs can be completely overwhelmed by the impact of channelization, especially when muddy banks with aquatic vegetation present a substantial part of habitat diversity and significantly contribute to the total species pool.     

Vegetation succession in deglaciated landscapes: implications for sediment and landscape stability

Landscapes exposed by glacial retreat provide an ideal natural laboratory to study the processes involved in transforming a highly disturbed, glacially influenced landscape to a stable, diverse ecosystem which supports numerous species and communities. Large‐scale vegetation development and changes in sediment availability, used as a proxy for paraglacial adjustment following rapid deglaciation, were assessed using information from remote sensing. Delineation of broad successional vegetation cover types was undertaken using Landsat satellite imagery (covering a 22 year period) to document the rate and trajectory of terrestrial vegetation development. Use of a space‐for‐time substitution in Glacier Bay National Park, Alaska, allowed ‘back‐calculation’ of the age and stage of development of six catchments over 206 years. The high accuracy (89.2%) of the remotely sensed information used in monitoring successional change allowed detection of a high rate of change in vegetation classes in early successional stages (bare sediment and alder). In contrast, later successional stages (spruce and spruce–hemlock dominated forest) had high vegetation class retention, and low turnover. Modelled rates of vegetation change generally confirmed the estimated rates of successional turnover previously reported. These data, when combined with the known influence of terrestrial succession on soil development and sediment availability, suggest how physical and biological processes interact over time to influence paraglacial adjustment following deglaciation. This study highlights the application of remote sensing of successional chronosequence landscapes to assess the temporal dynamics of paraglacial adjustment following rapid deglaciation and shows the importance of incorporating bio‐physical interactions within landscape evolution models. © 2014 The Authors. Earth Surface Processes and Landforms published by John Wiley & Sons Ltd.     

Distinguishing the effects of oil pollution from natural cyclical phenomena on the biota of Bantry Bay,Ireland

Marked declines in the numbers/percentage cover of four common species of littoral organism occurred between July 1978 and July 1979 on mainland shores in Bantry Bay, Ireland. Regular monthly monitoring has enabled most of these declines to be related to natural phenomena rather than to pollution of the Bay by oil from the Tanker Betelgeuse which exploded in January 1979. Only Pelvetia canaliculata and recently settled spat of Balanus balanoides show any indication of having been affected by oil/dispersants. By contrast, the North shore of Whiddy Island was severely affected.     

Bacterial communities are sensitive indicators of contaminant stress

With many environments worldwide experiencing at least some degree of anthropogenic modification, there is great urgency to identify sensitive indicators of ecosystem stress. Estuarine organisms are particularly vulnerable to anthropogenic contaminants. This study presents bacterial communities as sensitive indicators of contaminant stress. Sediments were collected from multiple sites within inner and outer zones of three heavily modified and three relatively unmodified estuaries. Bacterial communities were censused using Automated Ribosomal Intergenic Spacer Analysis and analysed for a suite of metal and PAH contaminants. Shifts in both bacterial community composition and diversity showed strong associations with sediment contaminant concentrations, particularly with metals. Importantly, these changes are discernable from environmental variation inherent to highly complex estuarine environments. Moreover, variation in bacterial communities within sites was limited. This allowed for differences between sites, zones and estuaries to be explained by variables of interest such as contaminants that vary between, but not within individual sites.     

Effects of the 'Prestige' oil spill on macroalgal assemblages: large-scale comparison

An assessment of the effects of the 'Prestige' oil spill on intertidal, macroalgal assemblages was carried out comparing abundance data obtained before and after the spill. Four zones in the North and Northwest coast of Spain were sampled, one of them located at the immediate vicinity of the spill, the zone most heavily oiled. Macroalgal assemblages had similar structure between years. Neither critical decrease in abundance of the dominant macroalgae, nor increase in opportunistic species were found. Some differences in abundance were observed, but they did not show any pattern, being more likely the result of the natural variability of the assemblage. Extensive, but not intense fuel deposition on the shores and a limited use of aggressive cleanup methods are suggested as possible causes for the lack of the effects in these assemblages after the 'Prestige' oil spill.     

Distribution,characteristics and potential impacts of chromophoric dissolved organic matter (CDOM) in Hudson Strait and Hudson Bay,Canada

The characteristics of chromophoric dissolved organic matter (CDOM) were studied in Hudson Bay and Hudson Strait in the Canadian Arctic. Hudson Bay receives a disproportionately large influx of river runoff. With high dissolved organic matter (DOM) concentrations in Arctic rivers the influence of CDOM on coastal and ocean systems can be significant, yet the distribution, characteristics and potential consequences of CDOM in these waters remain unknown. We collected 470 discrete water samples in offshore, coastal, estuarine and river waters in the region during September and October 2005. Mixing of CDOM appeared conservative with salinity, although regional differences exist due to variable DOM composition in the rivers discharging to the Bay and the presence of sea-ice melt, which has low CDOM concentrations and low salinity. There were higher concentrations of CDOM in Hudson Bay, especially in coastal waters with salinities <28$<28$, due to river runoff. Using CDOM composition of water masses as a tracer for the freshwater components revealed that river runoff is largely constrained to nearshore waters in Hudson Bay, while sea-ice melt is distributed more evenly in the Bay. Strong inshore–offshore gradients in the bio-optical properties of the surface waters in the Hudson Bay cause large variation in penetration of ultraviolet radiation and the photic depth within the bay, potentially controlling the vertical distribution of biomass and occurrence of deep chlorophyll maxima which are prevalent only in the more transparent offshore waters of the bay. The CDOM distribution and associated photoprocesses may influence the thermodynamics and stratification of the coastal waters, through trapping of radiant heating within the top few meters of the water column. Photoproduction of biologically labile substrates from CDOM could potentially stimulate the growth of biomass in Hudson Bay coastal waters. Further studies are needed to investigate the importance of terrestrial DOM in the Hudson Bay region, and the impact of hydroelectric development and climate change on these processes.     

Vegetation succession prevents dry lake beds from becoming dust sources in the semi‐arid steppe region of China

East Asian dust storms have become increasingly intense over the last two decades, and the arid inland regions of northern China have been recognized as the main dust source areas. Numerous lakes in this region have recently become desiccated, leaving large areas of bare ground prone to becoming potential dust sources. Vegetation cover characteristics and vegetation succession following lake desiccation remain unclear. Here we chose eight inland dry lakes, one outflow lake and one river on the southeast edge of the Inner Mongolian Plateau to investigate vegetation patterns along transects from lake bed to lake shore, and determine the relationships between vegetation patterns and environmental factors. The results show that dry lake bed soils do indeed have high contents of fine particles. Also, soil salt content is the most critical control on vegetation succession on desiccated lake beds, and vegetation is unlikely to colonize areas with soil salt content ≥5%. Soil texture additionally influenced vegetation patterns by affecting soil salt content. The likely vegetation succession on dry like beds is Nitraria tangutorum community > Suaeda corniculata and Suaeda glauca communities > Achnatherum splendens and Elymus sibiricus communities, and finally Carex duriuscula community as the probable climax. When vegetation is at the later stages of succession, for example with Achnatherum splendens communities, Elymus sibiricus communities and Carex duriuscula communities, soil may be protected from wind erosion because of their high vegetation cover and high proportion of perennials. We suggest grazing should be avoided around lake shores, especially in Achnatherum splendens communities, because high vegetation cover and biomass not only protect soil from erosion, but also promote the deposition of fine particles blown from upwind regions. Copyright © 2010 John Wiley & Sons, Ltd.     

Mangrove microbial diversity and the impact of trophic contamination

Mangroves are threatened ecosystems that provide numerous ecosystem services, especially through their wide biodiversity, and their bioremediation capacity is a challenging question in tropical areas. In a mangrove in Mayotte, we studied the potential role of microbial biofilm communities in removing nutrient loads from pre-treated wastewater. Microbial community samples were collected from tree roots, sediments, water, and from a colonization device, and their structure and dynamics were compared in two areas: one exposed to sewage and the other not. The samples from the colonization devices accurately reflected the natural communities in terms of diversity. Communities in the zone exposed to sewage were characterized by more green algae and diatoms, higher bacteria densities, as well as different compositions. In the area exposed to sewage, the higher cell densities associated with specific diversity patterns highlighted adapted communities that may play a significant role in the fate of nutrients.     

Impact of the Phaeocystis globosa spring bloom on the intertidal benthic compartment in the eastern English Channel: a synthesis

From 1999 to 2005, studies carried out in the frame of regional and national French programs aimed to determine whether the Phaeocystisglobosa bloom affected the intertidal benthic communities of the French coast of the eastern English Channel in terms of composition and/or functioning. Study sites were chosen to cover most of the typical shore types encountered on this coast (a rocky shore, an exposed sandy beach and a small estuary). Both the presence of active Phaeocystis cells and their degradation product (foam) did have a significant impact on the studied shores. The primary production and growth rates of the kelp Saccharina latissima decreased during the bloom because of a shortage of light and nutrient for the macroalgae. On sandy sediments, the benthic metabolism (community respiration and community primary production), as well as the nitrification rate, were enhanced during foam deposits, in relation with the presence of bacteria and active pelagic cells within the decaying colonies. In estuarine sediments, the most impressive impact was the formation of a crust at the sediment surface due to drying foam. This led to anoxic conditions in the surface sediment and resulted in a high mortality among the benthic community. Some organisms also tended to migrate upward and were then directly accessible to the higher trophic level represented by birds. Phaeocystis then created a shortcut in the estuarine trophic network. Most of these modifications lasted shortly and all the systems considered came back to their regular properties and activities a few weeks after the end of the bloom, except for the most impacted estuarine area.