Assessing the effect of emergent vegetation in a surface-flow constructed wetland on eutrophication reversion and biodiversity enhancement

Abstract A free water-surface constructed wetland (F4-unit) was created in a Mediterranean protected site to treat eutrophic inflows to the Albufera de Valencia lagoon, and enhance biodiversity. In 2013, the F4-unit was divided into three subunits and planted with different compositions of emergent macrophytes (Phragmites australis, Iris pseudacorus, and a mixture of several macrophytes: F4-Reed, F4-Iris and F4-Mixed subunits). Two nine-month periods were monitored after planting, and water quality variables were measured in the inflows and outflows. Waterfowl occupation and plankton biomass were analyzed to evaluate their role in eutrophication reversion. The low removal efficiencies relat…

Foreseeable global warming will differentially affect Chara vulgaris populations from different altitudes

Abstract A 2–4 °C increase in temperature is foreseen for the Mediterranean region by the end of the 21st century due to climate change. This warming is expected to occur as heat waves, and will be particularly important for shallow water bodies which are the main freshwater ecosystems in the Mediterranean. We subjected four populations of Chara vulgaris to a common garden experiment. Each population came from locations at different altitudes, hence living at different mean temperatures. Individual charophyte cultures were initiated with specimens from four origins and acclimatized at 20 °C in a controlled environment for 3 weeks. After this, all the populations were incubated under 3 water…

Inferring the Relative Resilience of Alternative States

Ecological systems may occur in alternative states that differ in ecological structures, functions and processes. Resilience is the measure of disturbance an ecological system can absorb before changing states. However, how the intrinsic structures and processes of systems that characterize their states affects their resilience remains unclear. We analyzed time series of phytoplankton communities at three sites in a floodplain in central Spain to assess the dominant frequencies or "temporal scales" in community dynamics and compared the patterns between a wet and a dry alternative state. The identified frequencies and cross-scale structures are expected to arise from positive feedbacks that…

The antagonistic effect of UV radiation on warming or nitrate enrichment depends on ecotypes of freshwater macroalgae (Charophytes).

Increases in ultraviolet radiation (UVR), a negative global change factor, affect aquatic primary producers. This effect is expected to be modulated by other global change factors, and to be different for populations adapted to different environments. A common garden experimental approach using freshwater green macroalgae, the cosmopolitan charophyte species Chara hispida and C. vulgaris, allowed us to test whether the beneficial increases in water temperature (T) and nitrate concentration (N) mitigate negative UVR effects. Also, whether these interactions would be not only species-specific but also according to the origin of the population; therefore, two populations of each species were u…

Allelopathic effects of microcystin-LR on the germination, growth and metabolism of five charophyte species and a submerged angiosperm

Abstract Microcystins (MCs) are produced by cyanobacteria in aquatic environments and adversely affect macrophytes at very high concentrations. However, the effects of MC on macrophytes at concentrations of environmental relevance are largely unknown. The main objective of this study was to analyze the allelopathic effects of MC-LR at natural concentrations (1, 8 and 16 μgMC-LR/L) on five charophyte species ( Chara aspera , C. baltica , C. hispida , C. vulgaris and Nitella hyalina ) and the angiosperm Myriophyllum spicatum . Macrophyte specimens were obtained from a restored area located in Albufera de Valencia Natural Park, a protected coastal Mediterranean wetland. Two different experimen…

The main limnological characteristics of the biogenic meromictic Lake La Cruz, a small karstic lake located on a dolomitic substrate, are outlined below. The lake owes its chemical stratification mainly to high concentrations of bicarbonate and calcium in the permanent monimolimnion, located below a depth of 19 m. A massive precipitation of CaCO3 crystals ("whiting"), occurring in the euphotic zone every summer, leads to further sedimentation and redissolution in deep waters with enhanced CO2 concentrations. Other ions, like magnesium, ammonium, reduced iron, silicate, etc. also contribute to the chemically-induced stability of the monimolimnion. The patterns of stratification during two co…

Time and heat for sexual reproduction: comparing the phenology of Chara hispida of two populations at different latitudes

Abstract Geographical location affects the main ecological factors driving the timing of plants’ life events. In addition, studying phenology is the simplest procedure to track current global warming and its effects on the success and survival of different populations of the same species. Little is known about the effect of water temperature and its corresponding accumulated heat on charophytes’ phenology. We compared differences in water temperature and sexual reproductive phenology of Chara hispida in two ponds of two countries located at different latitudes (Spain and Switzerland) over the same year. We estimated the accumulated heat required to develop from one phenophase to another (un…

The role of charophytes in a Mediterranean pond created forrestoration purposes

A small, shallow basin was created and flooded with groundwater in a Reserve Area in Albufera de València Natural Park (AVNP) under the scope of a restoration program intended to mimic typical environments such as freshwater springs, abundant in the past but currently suffering from deterioration (desiccation, pollution, etc.), with the ultimate goal of increasing local biodiversity of submerged macrophytes, particularly charophytes. In this study we have monitored, from April 2009 (a few months after its first flooding) to September 2012, the dynamics of charophyte growth and related physical, chemical and biotic variables to highlight the factors determining charophyte success. Just after…

We carried out enclosure experiments to assess the potential deleterious effects of the alien species common carp (Cyprinus carpio L.), mosquitofish (Gambusia holbrookii Gir.), and pumpkinseed sunfish (Lepomis gibbosus L.) on water quality and food web in a severely degraded floodplain wetland, the Spanish National Park Las Tablas de Daimiel. With addition of either carp or pumpkinseed sunfish, chlorophyll a and turbidity levels, as well as concentrations of total phosphorus and total nitrogen increased. The magnitude of this increase depended on fish species and was severest in the carp treatment. Mosquitofish did not significantly affect water quality. In the treatments with fish, cyclopo…

Carbon stable isotope composition of charophyte organic matter in a small and shallow Spanish water body as a baseline for future trophic studies

<p>Quantitative descriptions of foodweb structure based on isotope niche space require knowledge of producers’ isotopic signatures. In freshwater ecosystems charophytes are one of the main components of submerged vegetation and the feeding base for many herbivorous consumers, but knowledge about their organic carbon isotopic signatures is sparse. In this study, the δ<sup>13</sup>C organic values (and organic %C and %N) of the four species of submerged macrophytes (three charophytes - <em>Chara hispida</em>, <em>Nitella hyalina</em> and <em>Tolypella glomerata </em>- and one angiosperm, <em>Myriophyllum spicatum</em>) growing …

Aproximación a las comunidades de carófitos que existieron en la albufera de valencia a partir del estudio de las oósporas del sedimento

14 páginas, 8 figuras, 3 tablas.

Effects of overabundant nitrate and warmer temperatures on charophytes: The roles of plasticity and local adaptation

Global change effects, such as warming and increases in nitrogen loading, alter vulnerable Mediterranean aquatic systems, and charophytes can be one of the most affected groups. We addressed the possible interaction between these factors on two populations of the cosmopolitan charophytes Chara hispida and Chara vulgaris. Populations were taken from two different environments, a nitrate-poor mountain lake and a nitrate-rich Mediterranean coastal spring. The laboratory experiment had a 2 × 2 factorial design based on two nitrate levels (similar to and double the local conditions) and two temperatures. Increased temperatures favoured the growth of the four populations, but an increase in nitra…

Shifts in plankton assemblages promoted by free water surface constructed wetlands and their implications in eutrophication remediation

Abstract Two units of free water surface constructed wetlands (FWSCWs) were created in 2009 in a Mediterranean protected site, the Albufera de Valencia Natural Park, to treat eutrophic inflows to the Albufera de Valencia lagoon, the largest coastal lagoon in the Iberian Peninsula, affected by cultural eutrophication. Data of phytoplankton and zooplankton biomass and richness of the inflows and outflows corresponding to the first 3.5 years of operation of the system have been analyzed to evaluate the role of plankton in the eutrophication reversion. The FWSCWs significantly reduced the exported microalgal biomass, stimulated the potential mixotrophic groups of phytoplankton such as Euglenoph…

Wetland Restoration with Hydrophytes: A Review

Restoration cases with hydrophytes (those which develop all their vital functions inside the water or very close to the water surface, e.g., flowering) are less abundant compared to those using emergent plants. Here, I synthesize the latest knowledge in wetland restoration based on revegetation with hydrophytes and stress common challenges and potential solutions. The review mainly focusses on natural wetlands but also includes information about naturalized constructed wetlands, which nowadays are being used not only to improve water quality but also to increase biodiversity. Available publications, peer-reviewed and any public domain, from the last 20 years, were reviewed. Several countrie…

Selecting submerged macrophyte species for replanting in Mediterranean eutrophic wetlands

Abstract The criteria to select submerged plant species for restoration still need further evaluations for a better understanding of their suitability prior to undertaking large plantings which represent a large effort in terms of economical and human resources. In this study, previously to the restoration of a eutrophic area (Tancat de la Pipa) within the Albufera de Valencia Natural Park, the growth of three submerged plants (Ceratophyllum demersum, Myriophyllum spicatum and Stuckenia pectinata), an amphibious species (Ranunculus sceleratus), and three charophytes species (Chara vulgaris, Tolypella glomerata and Nitella hyalina) has been evaluated under controlled laboratory conditions in…

Effects of UVB radiation exposure from the molecular to the organism level in macrophytes from shallow Mediterranean habitats

Abstract The available data on the effects of UVB radiation (UVBR) are scarce for submerged macrophytes, particularly charophytes. We studied the effects of UVBR on Chara baltica, Chara hispida, Chara vulgaris, Nitella hyalina and Myriophyllum spicatum, collected from shallow Mediterranean waterbodies. In a short-term laboratory experiment, we subjected these species to three different UVBR treatments corresponding to daily biologically effective integrated doses of 0, 1 and 6 kJ m−2. The analysed response variables were DNA damage, UV-absorbing compounds (both the methanol-soluble–SUVACs- and, for the first time in charophytes, the methanol-insoluble cell wall-bound fraction–WUVACs-), the …

Plankton Ecology and Diversity

In this chapter we describe plankton responses induced by fluctuating hydrology and eutrophication process in the semi-arid TDNP wetland. We have followed the planktonic community since 1992, at seasonal and interannual scales, covering as well the spatial heterogeneity of the wetland. The studied planktonic components were bacterioplankton, autotrophic picoplankton, nano and microphytoplankton and zooplankton, including ciliates. Plankton has been studied in terms of species composition (diversity), functional groups, spatial heterogeneity, population dynamics. The present data from this wetland allow us to determine which factors (resources and conditions) are relevant for each group and …

Non‐trophic key players in aquatic ecosystems: a mesocosm experiment

Reconstruction of the former charophyte community out of the fructifications identified in Albufera de València lagoon sediments

Abstract Several decades ago, due to eutrophication, Albufera de Valencia lagoon lost the charophyte meadows that covered the bottom in the past but had never been studied in detail. An area close to the lagoon will be restored now to improve water quality and to recover submerged vegetation. To have information on the background of the charophytes from this area, four 80–87 cm long sediment cores were extracted from the lagoon. The subfossil charophyte fructifications (oospores and gyrogonites) were isolated and identified in two of the four cores. Moreover, sediment core fractions from the third and fourth cores were allowed to germinate under several conditions. The study of the lagoon s…

Plankton participation in the performance of three constructed wetlands within a Mediterranean natural park.

Abstract The contribution of plankton in the performance of three constructed wetlands (CWs) within the Albufera de Valencia Natural Park has been analyzed, taxonomic group by group, over a two-year operation period in the different sectors of each CW: horizontal subsurface-flow –A–, free-water surface flow –B– and lagoons –C–. Tancat de la Pipa CW (TPCW) only contains B–C sectors, while the others have all three types. Treatment efficiency of each sector type on the taxonomic groups was evaluated by calculating frequency of phytoplankton reduction (or zooplankton production), mass removal (or production) efficiencies and rates, and accumulated removed phytoplankton mass (or produced zoopla…

Autotrophic and Heterotrophic Picoplankton in Wetlands: Differences with Lake Patterns

This study describes the occurrence, importance and seasonal patterns of picoplankton in two wetlands (TDNP and La Safor), and compares them to a system of fifteen interconnected lakes (Ruidera). In TDNP we performed a six-year monthly study in three sites of the wetland. Bacterial abundance increased throughout time and the autotrophic picoplankton (APP) range was wide (up to 33 x 10 6 cells/ml). The annual averaged APP contribution to total picoplankton and phytoplankton biovolumes was 0.5-22% and 0.03-6% respectively. There were large differences among sites in terms of APP absolute and relative abundance and seasonal patterns. In La Safor, the APP relative contribution to picoplankton a…

Controlling factors of phytoplankton assemblages in wetlands: an experimental approach

The aim of this work is to answer some questions like: what factors control the phytoplankton assemblage? What factor or factors are perturbing the assemblage? What factors are driving or maintaining the stability? Are the different responses to the control factors dependent on the hierarchy level? For that, we tested experimentally the influence of herbivory, planktivory, nutrients and sediment on phytoplankton assemblages and its stability from a hypertrophic wetland (Las Tablas de Daimiel National Park, Spain) in three microcosm experiments. The study of the steady state phytoplankton assemblages in this perturbed system could point out some underlying processes instead of competition. T…

Environmental Variables and Planktonic Communities in Two Ponds of El Hondo Wetland (SE Spain)

The annual cycle of physical and chemical variables and plankton dynamics was studied in two shallow ponds (East and West Ponds) of the El Hondo wetland, an ecosystem of international importance. Water conductivity increased up to 31-49 mS cm -1 as water level decreased due to high evaporation and minimal water inputs. Initially considered mesohaline, the waters were reclassified as polyhaline during the hot season. EP was subject to successive desiccation-flooding cycles, and flooding of the dried sediment caused the release of high concentrations of nitrogen and phosphorus compounds, which quickly depleted. The algal species composition was typical of eutrophic ecosystems, and the chlorop…

Revision of the Eduardo Reyes Prósper’s herbarium and typification of three species of charophytes (<em>Charophyceae : Characeae</em>) present within it

A study of the charophytes collected by Eduardo Reyes Prósper in the period 1904–1906 and kept in the herbarium of the Royal Botanical Garden of Madrid (MA-ALGAE) is carried out. Reyes Prósper’s herbarium consists of 112 sheets, which had not been revised until now, among which are the type materials of Lamprothamnus aragonensis Reyes y Prosper, Lamprothamnus toletanus Reyes y Prosper and Tolypella giennensis Reyes y Prosper, of which their lectotypes are here designated.

The role of eutrophication reduction of two small man-made Mediterranean lagoons in the context of a broader remediation system: Effects on water quality and plankton contribution

In order to meet the requirements of the European Union Water Framework Directive for the Albufera de Valencia (AV) Natural Park, in 2009, several areas of free water surface constructed wetlands (FWSCWs) planted with emergent vegetation and two small shallow lagoons planted with submerged macrophytes were created over a 40-ha area formerly occupied by rice fields. This area is currently a reserve known as Tancat de la Pipa. The dual goal of this programme was to improve the quality of the hyper-eutrophicated waters of the AV lagoon, the largest littoral lagoon in the Iberian Peninsula, and to restore former lost habitats to increase the biodiversity of the area. The lagoons were mainly fed…

Nitrogen sedimentation in a lake affected by massive nitrogen inputs: autochthonous versus allochthonous effects

12 pages, and tables statics, and figures

Effects of Changing Rainfall on the Limnology of a Mediterranean, Flowthrough-Seepage Chain of Lakes

Relationships between groundwater and lake ecology are often overlooked, but they may be strong, particularly in seepage lakes. As a result, the nature and degree of groundwater effects on lakes are usually neglected. In this study interactions among rainfall, groundwater and surface water and their limnological effects were traced seasonally for two years of changing rainfall in a Spanish flowthrough, seepage lake complex. Cumulative rainfall dictated recharge of groundwater with delays of nine months. Groundwater discharge, in turn, increased surface discharge downstream. Mediated by the geographical setting of lakes, both fluxes impinged on lake water renewal time, but effects of the lat…

Different responses of coexisting Chara species to foreseeable Mediterranean temperature and salinity increases

Abstract An increase in temperature of approximately 4 °C is expected to occur in the Mediterranean by the end of the century. Concomitant to this warming, a foreseeable rise in salinity will affect aquatic species. We addressed the effects of warming and salinity, and their interaction on three coexisting characean species (Chara aspera, C. hispida and C. vulgaris) from a Spanish Mediterranean interdunal pond (spring water temperature 20–23 °C, when charophytes re-grew; salinity 1.3–1.8 PSU). A laboratory experiment was designed with two levels of water temperature treatment (23 and 27 °C), plus two levels of salinity treatment (0.4 and 4.0 PSU). The variables considered were total length,…

Persistence of submerged macrophytes in a drying world: Unravelling the timing and the environmental drivers to produce drought-resistant propagules

Plankton biodiversity in a landscape of shallow water bodies (Mediterranean coast, Spain)

A large spatial heterogeneity was detected in La Safor, a coastal area with different kinds of small and shallow water bodies. The area exhibits a sharp gradient in eutrophication (0.004–20 mgP l−1; 0.6–457 μg Chl a l−1) and varied water body features (morphology: size, depth; hydrology; vegetation, etc.). These factors result in a high diversity of aquatic habitats within the area. One hundred and twenty eight species of microalgae were identified in the 32 sampling stations studied (in the wet and dry periods). The phytoplankton richness varied between 1 and 21 species, the Shannon-Wiener index range was 0–3 bits, the index of floral originality (IFO) range was 0.1 – 0.6. A total number o…

Structure and vulnerability of the multi‐interaction network in macrophyte‐dominated lakes

The network approach is crucial to understand how ecosystems are structured and how they will respond to the disturbances (e.g. the current global change). We have recreated the multi‐interaction network of a shallow freshwater lake dominated by submerged macrophytes (Charophytes), a known system very vulnerable to environmental changes, considering both trophic and non‐trophic relationships among its elements. To minimize the environmental variability, we established it in an experimental mesocosm, including three habitats: the pelagic, the habitat around the meadow and the periphytic community living on macrophytes. We aimed to study the structure of this network and the roles of its elem…

Population dynamics of Cyclotella ocellata (Bacillariophyceae): endogenous and exogenous factors

Population dynamics of Cyclotella ocellata PANTOCSEK were examined from 1991 to 1993 in an oligo-mesotrophic, gravel-pit lake near Madrid (Spain). Monthly and daily sampling was carried out to track seasonal- and auxosporulation dynamics, respectively. This species was the most abundant planktonic diatom in the lake. The population increased steadily in spring until the seasonal peak (3.8.10 6 ind/ L in April) and then abundance declined slowly through late winter. Time series techniques showed that the seasonal dynamics were driven by exogenous factors: there was a relationship between Cyclotella abundance and the chemical environment, both TP (synchronic) and SRSi (with delay). Auxosporul…

Restoration of two small Mediterranean lagoons: The dynamics of submerged macrophytes and factors that affect the success of revegetation

Abstract Two small, shallow lagoons newly created from former rice fields in the Mediterranean coast (Albufera de Valencia Natural Park, Spain) were restored by planting several species of submerged macrophytes (Myriophyllum spicatum, Ceratophyllum demersum, Potamogeton spp., Zannichellia peltata and Ranunculus peltatus) in 2008. Charophytes also appeared spontaneously. M. spicatum was the species that dominated both lagoons and almost completely covered their surface. M. spicatum reached a high biomass and displayed a seasonal pattern, declining during the cold season and reappearing in the spring. No submerged vegetation was observed in the lagoons from 2011 onward. The major purpose of t…

Unravelling the bacterial diversity found in the semi-arid Tablas de Daimiel National Park wetland (central Spain)

Our knowledge of microbial diversity in the environment is still limited, and there are many species as yet unidentified in both soil and water. Studies of the microbial diversity of wetland ecosystems have been neglected for years, as is the case of Tablas de Daimiel National Park (TDNP), a Spanish semi-arid wetland system of international importance in terms of waterfowl. We report the bacterial diversity of water column, sediment (upper and lower layers) and biofilm samples from the TDNP system using a 16S rRNA gene library approach. A sequence comparison of the 703 clones obtained revealed a number of bacterial phylogroups unreported to date. Bacterial diversity was high (Shannon values…

On the tolerance of charophytes to high-nitrate concentrations

Currently a debate exists about whether the reduced growth of macrophytes with increased nitrogen loading in shallow ecosystems is determined by ecological or physiological factors. To discover whe...

Chara hispida beds as a sink of nitrogen: Evidence from growth, nitrogen uptake and decomposition

8 pages, and tables stadistics, and figures.

Reproductive phenology of submerged macrophytes: A tracker of year‐to‐year environmental variations

AIMS: Phenological shifts are useful trackers of climate change effects on species and communities, but few datasets focus on the climate‐driven phenological responses of aquatic ecosystem vegetation. We aimed to analyse the reproductive seasonality of a submerged macrophyte community, the synchrony among coexisting species, and their responses to several environmental factors. LOCATION: A Mediterranean coastal pond in Valencia, Spain. METHODS: We analysed three years of data on a submerged macrophyte community. Plants were collected biweekly or monthly (57 collection dates) to quantify the intensity of flowering/fruiting per species. The reproductive seasonality of the community was tested…

Germination capability of four charophyte populations (Characeae) from Mediterranean brackish ponds under warm experimental conditions

ABSTRACTThe diaspore banks from two brackish ponds within the Albufera de Valencia Natural Park (Spain) containing oospores of Chara hispida, C. aspera, C. canescens and Nitella hyalina were analys...

The allelopathic capacity of submerged macrophytes shapes the microalgal assemblages from a recently restored coastal wetland

Abstract We have tested the efficiency of isolated and combined submerged macrophyte cultures to inhibit, through allelopathy, the natural phytoplankton growth. Both plants and microalgae come from the same wetland, a recently restored area in Albufera de Valencia Natural Park (Spain). The need to replant the area under restoration with submerged macrophytes makes this information essential for wetland management. The selection and culture of the submerged macrophytes used in that restoration (four charophytes: Chara hispida , Chara vulgaris , Chara baltica , Nitella hyalina , and one angiosperm: Myriophyllum spicatum ) provided a good opportunity to test in the laboratory the allelopathic …

Assessing the potential of Albufera de València Lagoon sediments for the restoration of charophyte meadows

Abstract Albufera de Valencia Lagoon, the largest littoral lagoon on the Iberian Peninsula, has been affected by eutrophication processes since the 1970s due to the direct dumping of sewage waters of industrial, agricultural and urban origin. Consequently, the submerged vegetation that covered the bottom of the lagoon has been lost (charophytes and Potamogeton spp., Myriophyllum spp. and Ceratophyllum spp.). Despite efforts to improve water quality, this vegetation has not recovered. No information about the potential of the recent sediments to host charophyte development is available. In this study, we analysed several sediment cores taken from different areas of the lagoon to determine th…

Nitrate uptake rates in freshwater plankton: the effect of food web structure

9 pages, 3 figure, 1 table

On the tolerance of charophytes to high-nitrate concentrations

Currently a debate exists about whether the reduced growth of macrophytes with increased nitrogen loading in shallow ecosystems is determined by ecological or physiological factors. To discover whether nitrate in the water is detrimental per se to charophytes, we subjected Chara hispida and Chara vulgaris specimens, collected from two habitats greatly differing in nitrate concentrations (1.5 and 10 mg NO3-N/L, annual means), to a wide nitrate range (0.5–50 mg NO3-N/L) in two experiments (with free-floating specimens using nitrate as the sole N source, and with planted specimens, with other N sources in sediment). Charophytes grew both unplanted and planted in all treatments, and growth redu…