Development of an epiphyte indicator of nutrient enrichment: Threshold values for seagrass epiphyte load

Metrics of epiphyte load on macrophytes were evaluated for use as quantitative biological indicators for nutrient impacts in estuarine waters, based on review and analysis of the literature on epiphytes and macrophytes, primarily seagrasses, but including some brackish and freshwater rooted macrophyte species. An approach is presented that empirically derives threshold epiphyte loads which are likely to cause specified levels of decrease in macrophyte response metrics such as biomass, shoot density, percent cover, production and growth. Data from 36 studies of 10 macrophyte species were pooled to derive relationships between epiphyte load and −25 and −50% seagrass response levels, which are proposed as the primary basis for establishment of critical threshold values. Given multiple sources of variability in the response data, threshold ranges based on the range of values falling between the median and the 75th quantiles of observations at a given seagrass response level are proposed rather than single, critical point values. Four epiphyte load threshold categories − low, moderate, high, very high, are proposed. Comparison of values of epiphyte loads associated with 25 and 50% reductions in light to macrophytes suggest that the threshold ranges are realistic both in terms of the principle mechanism of impact to macrophytes and in terms of the magnitude of resultant impacts expressed by the macrophytes. Some variability in response levels was observed among climate regions, and additional data collected with a standardized approach could help in the development of regionalized threshold ranges for the epiphyte load indicator.     

Relative effects of nutrient enrichment and grazing on epiphyte-macrophyte (Zostera marina L.) dynamics

The independent and interactive effects of nutrient concentration and epiphyte grazers on epiphyte biomass and macrophyte growth and production were examined in Zostera marina L. (eelgrass) microcosms. Experiments were conducted during early summer, late summer, fall, and spring in a greenhouse on the York River estuary of Chesapeake Bay. Nutrient treatments consisted of ambient or enriched (3× ambient) concentrations of inorganic nitrogen (ammonium nitrate) and phosphate. Grazer treatments consisted of the presence or absence of field densities of isopods, amphipods, and gastropods. epiphyte biomass increased with both grazer removal and nutrient enrichment during summer and spring experiments. The effect of grazers was stronger than that of nutrients. There was little epiphyte response to treatment during the fall, a result possibly of high ambient nutrient concentrations and low grazing pressure. Under low grazer densities of early summer, macrophyte production (g m^–2 d^–1) was reduced by grazer removal and nutrient enrichment independently. Under high grazer densities of late summer, macrophyte production was reduced by enrichment only with grazers absent. During spring and fall there were no macrophyte responses to treatment. The relative influence of epiphytes on macrophyte production may have been related to seasonally changing water temperature and macrophyte requirements for light and inorganic carbon.     

Benthic community responses to nutrient enrichment and predator exclusion: Influence of background nutrient concentrations and interactive effects

Potential community effects of nutrient enhancement are a topic of theoretical interest and increasing management concern in coastal marine systems. While increased nutrient levels may lead to increased microalgal production and biomass, studies have provided variable evidence regarding the existence of upward cascade effects on macrofauna. In benthic marine communities, limitation by predation or factors preventing recruitment response may contribute to weak coupling between resource availability and macrobenthos abundances. We conducted blocked nutrient addition and predator exclusion experiments in the intertidal of two estuaries that varied in background nutrient concentrations (Cape Fear and White Oak, southeastern North Carolina). Benthic community comparisons were also made among these and two other North Carolina estuaries to examine correlations in distribution patterns. Cape Fear, which had the highest background nitrogen and phosphorus concentrations, also had highest ambient benthic microalgal biomass. There was no significant response of microalgal biomass to local nutrient additions in Cape Fear and only one macrofaunal taxon during one season exhibited abundance responses to nutrient additions. White Oak, with lower background nutrient levels, was characterized by significant microalgal responses to nutrient additions and significant macrofauna abundance responses for 50% of the species examined during summer experiments. However, all of these macrofauna declined in abundance with nutrient enhancement while biomass remained constant or significantly increased with nutrient additions. This suggests a complex response of macrofauna to nutrient additions in this estuary with greater biomass per individual but a corresponding decline in abundances. Top-down/bottom-up interactive effects were observed for haustoriid amphipods, which were uncommon or absent when predators had access, but exhibited strong biomass responses to nutrient enhancement when predators were excluded. These results support a growing body of literature that indicates the importance of background conditions in regulating benthic community responses to nutrient enhancement. However, responses may be complex with biomass per individual rather than densities being the primary response variable for some taxa and predator moderation of responses occurring for some taxa but not others.     

Biodiversity-productivity relations: an experimental evaluation of mechanisms

To examine the mechanisms underlying productivity-diversity relationships, we manipulated nutrient levels in replicate small-scale artificial habitat units in the marine subtidal zone and followed the process of community assembly. In contrast to most enrichment studies, algal diversity increased in enriched habitats relative to controls along with biomass – a result that may be explained by the low nutrient status of the region. Both the total number of faunal species and the total number of individuals were also significantly greater in enriched habitats, but the relationship between algal resources and faunal diversity did not support the resource heterogeneity hypothesis. Received: 14 June 1999 / Accepted: 29 October 1999     

Assessing the sustainability of water companies: A synthetic indicator approach

Performance indicators (PIs) are essential in the benchmarking process used to rate and rank water companies. However, a set of individual PIs does not provide a holistic assessment of company performance from multiple perspectives. A multidimensional evaluation of the performance of water companies can be achieved by aggregating the PIs into a synthetic indicator. Although the concept of sustainability involves economic, environmental and social criteria, most of the previous studies have not considered these three dimensions simultaneously. This paper discusses a process of indicator aggregation using two approaches based on multi-criteria decision analysis to evaluate and compare the sustainability of water companies from a holistic perspective. A synthetic indicator embracing economic, environmental and social PIs was computed for a sample of 154 Portuguese water companies. Both methods yielded similar rankings of water company sustainability. The techniques and results presented in this paper may be utilized as a means of improving the benchmarking process in regulated water industries, as well as providing valuable contributions to decision-makers on the most efficient steps for improving the sustainability of urban water services.     

The effects of light and nutrients on Caulerpa taxifolia and growth

Caulerpa taxifolia, an invasive species elsewhere in the world, is native to Moreton Bay where its distribution has been increasing in recent years. In Australia, dense beds of C. taxifolia are predominantly found in areas of low light and high nutrients (low water quality). Monitoring data from Moreton Bay suggests that native C. taxifolia is not directly replacing seagrass, but that there is a successional trend of seagrass loss and subsequent C. taxifolia colonization. The current study examined responses of C. taxifolia in relation to changes in environmental conditions using ambient water quality and a light/nutrient manipulative experiment. In the ambient water quality experiment we found that C. taxifolia grew significantly faster in areas with higher light (lower turbidity). The manipulative experiment demonstrated that nutrients stimulate C. taxifolia growth, however, light availability and seasonality appear to influence the response of C. taxifolia growth to nutrients in Moreton Bay. These findings suggest that C. taxifolia is unlikely to colonize seagrass beds in areas with high light and low nutrients; however, in areas with moderate light and moderate to high nutrients C. taxifolia and seagrass are likely to coexist.     

Effect of different Mg/Ca ratios and electrolyte concentrations in irrigation water on the nutrient content of wheat crop

Summary A study conducted in pots to evaluate the effect of different Mg/Ca ratios (2, 4, 8 and 16) and electrolyte concentrations (20 and 80 meq/l) at SAR 10 in irrigation water on the nutrient uptake and yield of wheat crop in two soils revealed that the average grain and dry matter yields of wheat decreased significantly with an increase in Mg/Ca ratio in irrigation water, but the magnitude of decrease was greater at higher electrolyte concentration than at lower electrolyte concentration. The concentration of Na in both straw and grain of wheat increased and that of K decreased with an increase in Mg/Ca ratio and electrolyte concentration of irrigation water, which led to higher Na/Ca and Na/K ratios in the plant. Further, the concentration of Ca and Mg both in straw as well as in grain increased with increasing electrolyte concentration of the irrigation water. An increasing proportion of Mg in saline irrigation water resulted in decreased concentration of Ca and increased concentration of Mg in both straw and grain of wheat crop. It was also noticed that the increasing proportion of Mg over Ca in the poor quality irrigation water increased the P content of both straw and grain of wheat crop.     

Long-term effects of nutrient enrichment on the condition and size-structure of an alpine brown trout population

Long-term effects of nutrient enrichment on a population of brown trout inhabiting a small, alpine lake in north-central Sweden have been studied for nearly 20 years. The study took place between 1981 and 1999 starting up between 1982 and 1987 with full nutrient enrichment of phosphorus and nitrogen in a ratio by weight of 1:8, followed by a period of reduction by half between 1988 and 1994, and thereafter no enrichment at all. Growth of the brown trout population was low before the application of nutrients. Fertilization promoted the development of zooplankton in great abundance, which gave rise to abundant food for the trout. Already during the first year of nutrient addition the average 4+ and 5+ fish had increased in weight by nearly 50% and in length by about 30%. Maximum growth was reached 5–6 years later—weight by about 175% and length by about 50% higher than before application. The slope of the growth curves for fish of ages 2+ to 5+ increased significantly from 1981 to 1987, and so did the size-variation with a high proportion of the fish reaching larger size. After each change in nutrient treatment the mean weights of 5–6-year-old trout were maintained for about 3 years. Five years after termination of fertilization growth was nearly back to the original state. Although badly needed, long-term studies of fish populations like this are few in the literature.     

Resurvey studies of terricolous bryophytes and lichens indicate a widespread nutrient enrichment in German forests

Questions

Vegetation resurveys, both single studies and meta-analyses, have predominantly been based on vascular plant data while bryophytes and lichens have largely been neglected. Our study aims to fill this gap and addresses the following research questions: has the overall species richness of terricolous bryophytes and lichens in forests changed over time? Which are the winners and losers among single species and ecological species groups? Do the results give a signal of the impact of nutrient enrichment, of changes in the light regime and of climate change?

Location

Deciduous and coniferous forests in Germany.

Methods

We compiled 35 single resurvey data sets, including 1096 plots in total (each sampled twice). The time interval between initial surveys and resurveys ranged from 10 to 65 years. The differences between old and new plots were analysed with respect to the frequency of single species, total species richness, and the absolute and relative numbers of taxa in the species groups. Trend scores of species were related to ecological indicator values to identify the main environmental drivers behind the observed changes.

Results

Total species richness did not systematically change, while pleurocarpous mosses had increased at the expense of acrocarpous mosses and, in coniferous forests, of lichens. Weak changes were generally observed in deciduous forests on base-rich soils. In coniferous forests and in deciduous forests on acid soils, species with high nitrogen demand and high shade tolerance had increased, whereas those being typical for more infertile and open forest sites had decreased. There were trends towards a larger share of taxa with a more oceanic distribution.

Conclusions

The changes in the vegetation of terricolous bryophytes and lichens in the studied forests indicate nutrient enrichment and increasingly shady conditions in forests on acid soils, likely caused by nitrogen deposition and shrub layer closure.     

Anthropogenic nutrient enrichment of seagrass and coral reef communities in the Lower Florida Keys: discrimination of local versus regional nitrogen sources

Land-based nutrient pollution represents a significant human threat to coral reefs globally. We examined this phenomenon in shallow seagrass and coral reef communities between the Content Keys (southern Florida Bay) and Looe Key (south of Big Pine Key) in the Lower Florida Keys by quantifying the role of physical forcing (rainfall, wind, tides) and water management on mainland South Florida to nutrient enrichment and blooms of phytoplankton, macroalgae, and seagrass epiphytes. Initial studies (Phase I) in 1996 involved daily water quality sampling (prior to, during, and following physical forcing events) at three stations (AJ, an inshore area directly impacted by sewage discharges; PR, a nearshore patch reef located inshore of Hawk Channel; and LK, an offshore bank reef at Looe Key) to assess the spatial and temporal patterns in advection of land-based nutrients to the offshore reefs. Concentrations of dissolved inorganic nitrogen (DIN=NH₄⁺+NO₃⁻+NO₂⁻), soluble reactive phosphorus (SRP), and chlorophyll a increased at PR and LK following a wind event (∼15 knots, northeast) in mid-February. The highest DIN (mostly NH₄⁺) and SRP concentrations of the entire study occurred at the inshore AJ during an extreme low tide in March. Following the onset of the wet season in May, mean NH₄⁺ and chlorophyll a concentrations increased significantly to maximum seasonal values at PR and LK during summer; relatively low concentrations of NO₃⁻ and a low f-ratio (NO₃⁻/NH₄⁺+NO₃⁻) at all stations during summer do not support the hypothesis that the seasonal phytoplankton blooms resulted from upwelling of NO₃⁻. A bloom of the seagrass epiphyte Cladosiphon occidentalis (phaeophyta) followed the onset of the rainy season and increased NH₄⁺ concentrations at LK, resulting in very high epiphyte:blade ratios (∼3:1) on Thalassia testudinum. Biomass of macroalgae increased at all three stations from relatively low values (<50 g dry wt m⁻²) in winter and early spring to higher values (∼100-300 g dry wt m⁻²) typical of eutrophic seagrass meadows and coral reefs following the onset of the rainy season. The mean δ¹⁵N value of Laurencia intricata (rhodophyta) during 1996 at AJ (+4.7‰) was within the range reported for macroalgae growing on sewage nitrogen; lower values at the more offshore PR (+3.1‰) and LK (+2.9‰) were at the low end of the sewage range, indicating an offshore dilution of the sewage signal during the 1996 study. However, transient increases in δ¹⁵N of Cladophora catanata (chlorophtyta) from ~+2% to +5% at LK concurrent with elevated NH₄⁺ concentrations following rain and/or wind events in May and July suggest episodic advection of sewage nitrogen to the offshore LK station. The Phase II study involved sampling of macroalgae for δ¹⁵N along a gradient from the Content Keys through Big Pine Key and offshore to LK in the summer wet season of 2000 and again in the drought of spring 2001. During the July 2000 sampling, macroalgae in nearshore waters around Big Pine Key had elevated δ¹⁵N values (~+4‰) characteristic of sewage enrichment; lower values (~+2‰) at LK were similar to values reported for macroalgae in upstream waters of western Florida Bay influenced by nitrogen-rich Everglades runoff. That pattern contrasted with the drought sampling in March 2001, when δ¹⁵N values of macroalgae were elevated (+6‰) to levels characteristic of sewage enrichment over a broad spatial scale from the Content Keys to LK. These results suggest that regional-scale agricultural runoff from the mainland Everglades watersheds as well as local sewage discharges from the Florida Keys are both significant nitrogen sources supporting eutrophication and algal blooms in seagrass and coral reef communities in the Lower Florida Keys. Hydrological and physical forcing mechanisms, including rainfall, water management on the South Florida mainland, wind, and tides, regulate the relative importance and variability of these anthropogenic nitrogen inputs over gradients extending to the offshore waters of the Florida Reef Tract.     

Effects of irradiance, temperature, and nutrients on growth dynamics of seagrasses: A review

Productivity of seagrasses can be controlled by physiological processes, as well as various biotic and abiotic factors that influence plant metabolism. Light, temperature, and inorganic nutrients affect biochemical processes of organisms, and are considered as major factors controlling seagrass growth. Minimum light requirements for seagrass growth vary among species due to unique physiological and morphological adaptations of each species, and within species due to photo-acclimation to local light regimes. Seagrasses can enhance light harvesting efficiencies through photo-acclimation during low light conditions, and thus plants growing near their depth limit may have higher photosynthetic efficiencies. Annual temperatures, which are highly predictable in aquatic systems, play an important role in controlling site specific seasonal seagrass growth. Furthermore, both thermal adaptation and thermal tolerance contribute greatly to seagrass global distributions. The optimal growth temperature for temperate species range between 11.5 °C and 26 °C, whereas the optimal growth temperature for tropical/subtropical species is between 23 °C and 32 °C. However, productivity in persistent seagrasses is likely controlled by nutrient availability, including both water column and sediment nutrients. It has been demonstrated that seagrasses can assimilate nutrients through both leaf and root tissues, often with equal uptake contributions from water column and sediment nutrients. Seagrasses use HCO₃⁻ inefficiently as a carbon source, thus photosynthesis is not always saturated with respect to DIC at natural seawater concentrations leading to carbon limitation for seagrass growth. Our understanding of growth dynamics in seagrasses, as it relates to main environmental factors such as light, temperature, and nutrient availability, is critical for effective conservation and management of seagrass habitats.     

A 2-year experimental study on nutrient and predator influences on food web constituents in a shallow lake of north-west Spain

1. A 2‐year study was carried out on the roles of nutrients and fish in determining the plankton communities of a shallow lake in north‐west Spain. Outcomes were different each year depending on the initial conditions, especially of macrophyte biomass. In 1998 estimated initial ‘per cent water volume inhabited’ (PVI) by submerged macrophytes was about 35%. Phytoplankton biomass estimated as chlorophyll a was strongly controlled by fish, whereas effects of nutrient enrichment were not significant. In 1999 estimated PVI was 80%, no fish effect was observed on phytoplankton biomass, but nutrients had significant effects. Water temperatures were higher in 1998 than in 1999. 2. In the 1998 experiment, cladoceran populations were controlled by fish and cyanobacteria were the dominant phytoplankton group. There were no differences between effects of low (4 g fresh mass m^?2) and high (20 g fresh mass m^?2) fish density on total zooplankton biomass, but zooplankton biomass was higher in the absence of fish. With the high plant density in 1999, fish failed to control any group of the zooplankton community. 3. Total biovolume of phytoplankton strongly decreased with increased nutrient concentrations in 1998, although chlorophyll a concentrations did not significantly change. At higher nutrient concentrations, flagellate algae became more abundant with likely growth rates that could have overcompensated cladoceran feeding rates. This change in phytoplankton community composition may have been because of increases in the DIN : SRP ratio. Both chlorophyll a concentration and total phytoplankton biovolume increased significantly with nutrients in the 1999 experiment. 4. A strong decline of submerged macrophytes was observed in both years as nutrients increased, resulting in shading by periphyton. This shading effect could account for the plant decline despite lower water turbidity at the very high nutrient levels in 1998.     

Difference characteristics of phosphorus in Chara and two submerged angiosperm species: implications for phosphorus nutrient cycling in an aquatic ecosystem

Phosphorus speciation in three submerged macrophytes species, Chara fibrosa Agardh ex Bruzelius, Najas marina Linnaeus and Vallisneria gigantea Graebner, and the implications for phosphorus nutrient cycling in an aquatic ecosystem were studied, using sequential phosphorus fractionations. The results showed that C.␣fibrosa had a far higher residual ash and calcium content compared with the two angiosperm species, but lower total phosphorus content. Two different fractionation methods for phosphorus showed that the bioavailable water-soluble phosphorus (H₂O-P) and ammonium chloride extractable phosphorus (NH₄Cl-P) of the extractions used represented the major part of total plant phosphorus in the two angiosperm species, while organic phosphorus (NaOH-P) represented a relatively large fraction in C. fibrosa. In this species, about 12–15% of total plant phosphorus was calcium-bound phosphorus (HCl-P), occurring as co-precipitation with calcite encrustation, but this fraction was negligible in the two angiosperm species, i.e. less than 1%. The redox-insensitive forms of HCl-P are considered less bioavailable and not affected by anoxic conditions of bottom sediment, thus have potential as a phosphorus nutrient sink in aquatic ecosystems.     

Non-electrolyte permeability of plant cuticles: A critical evaluation of experimental methods

A method for measurement of non-electrolyte transport through enzymatically isolated cuticles of Ficus elastica Roxb. var. decora is presented. Permeance coefficients and hold-up times are determined under steady-state conditions. The transport system consists of two small reservoirs separated by the cuticle. The compounds tested (4-nitrophenol and pentachlorophenol) differ widely in lipid solubility and enable a critical evaluation of the apparatus used. The sorption capacity of the cuticle for lipophilic compounds is related to their cuticle/water partition coefficients. The problems of unstirred layers and of membrane integrity are discussed. The permeances measured ranged from 10⁻¹¹ to 10⁻⁵ m s⁻¹, the hold-up times from 50 s to 22 h.     

Water chemistry and nutrient budgets in an undisturbed evergreen rainforest of southern Chile

In a pristine evergreen rainforest of Nothofagus betuloides, located at the Cordillera de los Andes in southern Chile (41 °S), concentrations and fluxes of nutrients in bulk precipitation, cloud water, throughfall water, stemflow water, soil infiltration and percolation water and runoff water were measured. The main objectives of this study were to investigate canopy-soil-atmosphere interactions and to calculate input-output budgets. From May 1999 till April 2000, the experimental watershed received 8121 mm water (86% incident precipitation, 14% cloud water), of which the canopy intercepted 16%. Runoff water volume amounted 9527 mm. Bulk deposition of inorganic (DIN) and organic (DON) nitrogen amounted 3.6 kg ha^–1 year^–1 and 8.2 kg ha^–1 year^–1 respectively. Occult deposition (clouds + fog) contributes for 40% to the atmospheric nitrogen input (bulk + occult deposition) of the forest. An important part of the atmospheric ammonium deposition is retained within the canopy or converted to nitrate or organic nitrogen by epiphytic bacteria or lichens. Also the export of inorganic (0.9 kg ha^–1 year^–1) and organic (5.2 kg ha^–1 year^–1) nitrogen via runoff is lower than the input to the forest floor via throughfall and stemflow water (3.2 kg DIN ha–1 year^–1 and 5.6 kg DON ha^–1 year^–1). The low concentrations of NO ₃ ^– and NH ₄ ⁺ under the rooting depth suggest an effective biological immobilization by vegetation and soil microflora. Dry deposition and foliar leaching of base cations (K⁺, Ca²⁺, Mg²⁺) was estimated using a canopy budget model. Bulk deposition accounted for about 50% of the total atmospheric input. Calculated dry and occult deposition are both of equal value (about 25%). Foliar leaching of K⁺, Ca²⁺, and Mg²⁺ accounted for 45%, 38% and 6% of throughfall deposition respectively. On an annual basis, the experimental watershed was a net source for Na⁺, Ca²⁺ and Mg²⁺.     

The use of nutrient reduction and food-web management to improve water quality in the deep stratifying Wupper Reservoir, Germany

Only a combination of nutrient load abatement and food-web management proved efficient for the management of water quality in the deep stratifying Wupper Reservoir. Reduction of nutrient loading, was completed in winter 1992/1993, but resulted only in reduced winter/spring mixing of phosphorus concentrations. Since the capacity of the diatom spring bloom to remove nutrients from the trophogenic layer of this slightly eutrophic water-body was never exhausted, the surplus of total phosphorus available to support summer algal growth remained unchanged. Thus, nutrient reduction alone did not improve the water quality, as expected. Subsequent replacement of the smaller Daphnia cucullata by the larger Daphnia galeata-hyalina complex that was attributable to successful food-web management did, however, result in a shift from a turbid to a clear water regime in 1999. Clearly, the zooplankton community, and therefore food-web structure, played an integral role in nutrient recycling and in the repartitioning of the phosphorus pool. As diatom settling and grazing became much more tightly linked with the appearance of the larger-bodied Daphnia galeata-hyalina complex, which exploits lower-level food resources as early as May, daphnids increasingly acted as a sink for phosphorus. This increased export fluxes out of the pelagic zone and leaves a smaller surplus of total phosphorus to support the accumulation of summer algae. Consequently, water transparency and total chlorophyll concentrations in summer improved with food-web restructuring, indicating real oligotrophication of Wupper Reservoir driven by internal feedbacks. Handling editor: S. I. Dodson     

Effects of experimental warming and nitrogen enrichment on leaf and litter chemistry of a wetland grass,Phragmites australis

As global climate is warming and the nitrogen cycle accelerates, plants are likely to respond not only by shifting community composition, but also by adjusting traits such as tissue chemistry. We subjected a widespread wetland plant, Phragmites australis, to increased nitrate supply and elevated temperature in enclosures that were established in a littoral permanently submerged freshwater marsh. The nitrogen (N) and phosphorus (P) concentrations in green leaves ranged from 11.4 to 13.8 mg N and from 1.5 to 2.0 mg P g⁻¹ dry mass. While the N concentration changed little in brown litter, the P concentration decreased to 0.53–0.65 mg P g⁻¹ litter dry mass. Neither experimental warming of the water and sediment surface, nor nitrate enrichment during the growing season affected nitrogen or phosphorus concentrations in green leaves. Concentrations of the two major structural carbon compounds in plant litter, cellulose and lignin, were also unaffected, ranging from 32.1 to 34.2% of dry mass for cellulose and from 16.3 to 17.7% of dry mass for lignin. Warming, however, significantly increased the nitrogen concentration of fully brown leaf litter. Thus, temperature appears to be more important than the supply of dissolved N in the water, especially in affecting leaf litter N concentrations in P. australis, even when only water but not air temperature is increased. This result may have implications for decomposition processes and decomposer food webs, which both depend on the quality of plant litter.