Primary Producer Assemblages Research Articles

Spatial and Ontogenetic Patterns in the Trophic Ecology of Juvenile Bull Sharks (Carcharhinus leucas) From the Northwest Gulf of Mexico

A combination of stomach content and stable isotope (δ13C, δ15N, and δ34S) analyses were used to characterize and examine spatiotemporal and ontogenetic trends in the feeding ecology of juvenile bull sharks (Carcharhinus leucas) captured in estuaries throughout the northwest Gulf of Mexico (GoM) between 2013 and 2016. Shark diets were dominated by fish prey taxa [>98% index of relative importance (%IRI)], and of those identified to the family level, two families comprised greater than 50% IRI, Mugilidae (mullets: ∼32%) and Sciaenidae (drums and croakers: 27%). Clupeidae (herrings: 14%) and Ariidae (sea catfishes: 15%) also contributed substantially to the diet of juvenile sharks, though consumption of Ariidae increased as consumption of Clupeidae decreased in juvenile sharks larger than 893 mm Fork Length (FL) (∼1 year old). Values of δ15N increased significantly with shark size, indicating a shift toward larger or higher trophic level prey with increasing shark size. Latitudinal and temporal trends in δ13C and δ34S suggest isotopic variation occurred in correspondence with shifts in primary producer assemblages and environmental drivers of sampled estuaries. These results highlight the importance of teleost prey resources along the freshwater-marine continuum in the diet of juvenile bull sharks, as well as the utility of natural tracers in tracking ontogenetic trends in feeding ecology.

Co-limitation by N and P Characterizes Phytoplankton Communities Across Nutrient Availability and Land Use

Historically, freshwater lakes have been widely assumed to be singly limited by phosphorus (P) because the dominant paradigm assumes that nitrogen fixation (N2 fixation) will compensate for any nitrogen (N) deficits. However, a growing body of evidence demonstrates that primary producer response to nutrient manipulation most frequently indicates co-limitation by N and P. Differences in N and P supply ratio have been shown to influence the identity and severity of nutrient limitation, but whether N and P concentration and the ratio of N to P concentrations can explain the frequency of co-limitation in aquatic primary producer assemblages remains unclear, especially in ecosystems subject to human perturbation that strongly increase nutrient availability. We determined how resource availability influences nutrient limitation by N and P of phytoplankton primary production across 12 lakes in Minnesota that vary in watershed land use and lake nutrient levels. We measured epilimnetic lake metabolism and indicators of N2 fixation to evaluate their influence on nutrient limitation status of planktonic algal assemblages. Despite large differences in land use (agricultural, urban, and suburban) and water column N and P availability, planktonic algal response to nutrient manipulation was consistently characterized by co-limitation by N and P across years and months. Neither P availability (as concentrations of total and inorganic forms) nor N2-flux rate predicted responses to nutrient additions. N availability significantly influenced responses of phytoplankton to nutrient additions across years, but this effect was small. The ratio of total N to total P significantly influenced the response to single additions of N and P (these effects were negative and positive, respectively) in summer 2013. Importantly, higher lake primary production and heterocyte count (number of nitrogen fixing cells) were also associated with a stronger, positive response to N + P addition. Overall, these data suggest that planktonic algal assemblages are predominantly characterized by co-limitation by N and P despite large and diverse human impacts on nutrient inputs. Additionally, higher rates of primary production increase the likelihood of co-limitation. Together, these results further support the paradigm shift toward dual management of N and P in aquatic ecosystems.

Holocene shifts in the primary producer community of large, shallow European Lake Peipsi, inferred from sediment pigment analysis

We used HPLC to identify and quantify pigments in a Holocene sediment record from large, shallow Lake Peipsi, Estonia. The aim of our study was to track the influence of long-term climate change (i.e. temperature fluctuations) on past dynamics of aquatic primary producers. Sedimentary pigments were separated and quantified in 182 samples that span the last ca. 10,000 years. There was an increasing trend in sedimentary pigment concentrations from basal to upper sediment layers, suggesting a gradual increase in lake trophic status through time. Using additive models, our results suggested that primary producer dynamics in Lake Peipsi were closely related to temperature fluctuations. We, however, identified two periods (early Holocene and after ca. 2.5 cal ka BP) when the relationship between primary producer composition and temperature was weak, suggesting the influence of additional drivers on the primary producer community. We postulate that: (a) the increase of primary producer biomass in the early Holocene could have been caused by input of allochthonous organic matter and nutrients from the flooded areas when water level in Lake Peipsi was increasing, and (b) changes in the abundance and structure of primary producer assemblages since ca. 2.5 cal ka BP was related to widespread agricultural activities in the Lake Peipsi catchment. These results suggest that human activities can disrupt the relationship between the primary producer community and temperature in large, shallow lakes.

Regional versus local drivers of water quality in the Windermere catchment, Lake District, United Kingdom: The dominant influence of wastewater pollution over the past 200years.

Freshwater ecosystems are threatened by multiple anthropogenic stressors acting over different spatial and temporal scales, resulting in toxic algal blooms, reduced water quality and hypoxia. However, while catchment characteristics act as a 'filter' modifying lake response to disturbance, little is known of the relative importance of different drivers and possible differentiation in the response of upland remote lakes in comparison to lowland, impacted lakes. Moreover, many studies have focussed on single lakes rather than looking at responses across a set of individual, yet connected lake basins. Here we used sedimentary algal pigments as an index of changes in primary producer assemblages over the last ~200years in a northern temperate watershed consisting of 11 upland and lowland lakes within the Lake District, United Kingdom, to test our hypotheses about landscape drivers. Specifically, we expected that the magnitude of change in phototrophic assemblages would be greatest in lowland rather than upland lakes due to more intensive human activities in the watersheds of the former (agriculture, urbanization). Regional parameters, such as climate dynamics, would be the predominant factors regulating lake primary producers in remote upland lakes and thus, synchronize the dynamic of primary producer assemblages in these basins. We found broad support for the hypotheses pertaining to lowland sites as wastewater treatment was the main predictor of changes to primary producer assemblages in lowland lakes. In contrast, upland headwaters responded weakly to variation in atmospheric temperature, and dynamics in primary producers across upland lakes were asynchronous. Collectively, these findings show that nutrient inputs from point sources overwhelm climatic controls of algae and nuisance cyanobacteria, but highlights that large-scale stressors do not always initiate coherent regional lake response. Furthermore, a lake's position in its landscape, its connectivity and proximity to point nutrients are important determinants of changes in production and composition of phototrophic assemblages.

From phytoplankton to oil shale reservoirs: A 19-million-year record of the Late Cretaceous Tethyan upwelling regime in the Levant Basin

Abstract The complexity of sedimentary organic-matter formation, its source to burial, and influence on bottom-water communities are addressed in this study of the 19 million years-long Upper Cretaceous Tethyan upwelling regime in the Levant. A multi-proxy approach is amalgamated into a model through which the paleoceanographic complexity of an intense and long-lasting upwelling system is better understood. A shift in primary producer assemblages and reorganization of the bottom water bacterial consortium followed the Campanian-Maastrichtian boundary, enforcing conspicuous sedimentological and faunal changes. The siliceous and phosphatic-bearing sediments of the Campanian gave way to Maastrichtian uniform and regionally distributed organic-rich carbonates, accompanied by an increase in sedimentation rates from ∼1 to >6 cm/kyr. This has facilitated both the increase in organic-matter production and its excellent preservation, and through diagenetic pathways affected its elemental composition. These paleoceanographic modifications have also affected foraminiferal assemblages, from massive blooms of triserial (buliminid) benthic foraminifera during the Campanian which have sequestered diatom chloroplasts, to diverse trochospiral forms during the Maastrichtian using nitrate instead of oxygen for their respiratory pathways. We propose that shifts in the type of primary producers promote a change in the amount and type of preserved organic matter, in bottom water communities, on the lithological composition of the rock-mass, and ultimately on the hydrocarbon generation potential of the studied source rock. The continued rise of southern Tethyan sea level and global highstands during the Upper Maastrichtian, culminating into Arabian Plate-wide maximum flooding surfaces, cause the migration of upwelling activity away from the study area, terminating the high productivity sequence in the Levant.

Herbivory enhances the diversity of primary producers in pond ecosystems.

Diversity of primary producer is often surprisingly high, despite few limiting factors such as nutrients and light to facilitate species coexistence. In theory, the presence of herbivores could increase the diversity of primary producers, resolving this "paradox of the plankton." Little experimental evidence supports this natural enemies hypothesis, but previous tests suffer from several deficiencies. Previous experiments often did not allow for multigeneration effects; utilized low diversity assemblages of herbivores; and limited opportunities for new primary producer and herbivore species to colonize and undergo species sorting that favors some species over others. Using pond plankton, we designed a mesocosm experiment that overcame these problems by allowing more time for interactions over multiple generations, openness to allow new colonists, and manipulated higher diversity of primary producers and grazers than have previous studies. With this design, the presence of zooplankton grazers doubled phytoplankton richness. The additional phytoplankton species in grazed mesocosms were larger, and therefore likely more grazer resistant. Furthermore, phytoplankton richness in grazed mesocosms was similar to that observed in natural ponds whereas it was much lower in mesocosms without grazers. However, stoichiometric imbalance caused by variation in nitrogen:phosphorus ratios and light supply did not alter phytoplankton richness. Therefore, grazers enhanced primary producer richness more strongly than ratios of nutrient supply (even though both grazing and ratios of resource supply altered composition of primary producer assemblages). Taken together, these experimental and field data show that grazing from a diverse assemblage of herbivores greatly elevated richness of phytoplankton producers in pond ecosystems.

Inferences for sources of oils from the Norwegian Barents Sea using statistical analysis of biomarkers

Conventional biomarker age indicators may not always give conclusive inferences for the age of an oil’s source rock, particularly with regard to condensates, in which most of the key biomarkers are often too sparse to determine. It is possible that other compounds, reflecting changes in primary producer assemblages or depositional environment, can provide age indications in basins that have experienced a progressive change in conditions. The Norwegian Barents Sea provides an opportunity to evaluate this approach, although the gradual changes are punctuated by intervals of stratification, OAEs and extinctions. Partial least squares analysis was used to identify a number of apparently suitable parameters, to establish an age calibration from source rock extracts and then to apply it to a variety of oils and condensates from the Barents Sea. The resulting source age inferences are consistent with conventional age parameters for the normal oils, providing a degree of confidence in their applicability to the problematic condensates. In the main part of the Hammerfest Basin, oils of Jurassic origin dominate, whereas contributions from Triassic sources are suggested towards the margins, particularly in pre-Jurassic reservoirs. To the east, on the Norsel High and in the Nordkapp Basin, the main oil sources are suggested to be Early Triassic, although a Palaeozoic, possibly Permian, source is likely on the margin of the Finnmark Platform in well 7128/4-1. Among the novel parameters, the relative abundances of phytanyl arenes and diadrimanes appear to be controlled by diagenetic conditions.

Hydrologic alteration affects aquatic plant assemblages in an arid-land river

Abstract We evaluated the effects of long-term flow alteration on primary-producer assemblages. In 1962, Flaming Gorge Dam was constructed on the Green River. The Yampa River has remained an unregulated hydrologically variable river that joins the Green River 100 km downstream from Flaming Gorge Dam. In the 1960s before dam construction only sparse occurrences of two macroalgae, Cladophora and Chara, and no submerged vascular plants were recorded in the Green and Yampa rivers. In 2009–2010, aquatic plants were abundant and widespread in the Green River from the dam downstream to the confluence with the Yampa River. The assemblage consisted of six vascular species, Elodea canadensis, Myriophyllum sibiricum, Nasturtium officinale, Potamogeton crispus, Potamogeton pectinatus, and Ranunculus aquatilis, the macroalgae Chara and Cladophora, and the bryophyte, Amblystegium riparium. In the Green River downstream from the Yampa River, and in the Yampa River, only sparse patches of Chara and Cladophora growing in ...

Eutrophication-Driven Shifts in Primary Producers in Shallow Coastal Systems: Implications for System Functional Change

Significant progress has been made recently towards a better understanding of the nature, causes, and consequences of anthropogenic eutrophication of shallow coastal systems. It is well established that, in pristine systems dominated by seagrasses, incipient to moderate eutrophication often leads to the replacement of seagrasses by phytoplankton and loose macroalgal mats as the dominant producers. However, less is known about the interactions between phytoplankton and loose macroalgae at intense eutrophication. Using a combination of original research and literature data, we provide support for the hypothesis that substantial macroalgal decline may occur at intense eutrophication due to severe water column shading. Our results suggest that such declines may be widespread. However, we also show that intense eutrophication is not always necessarily conducive to severe water column shading and large macroalgal declines, possibly due to short water residence time and/or elevated grazing on phytoplankton. Furthermore, we provide support to the hypothesis that the occurrence of hypoxic/anoxic conditions in eutrophication-driven shifts in dominant primary producer assemblages influences the nature and extent of functional change in the system. Focusing on the macroalgal blooms and seagrass decline that often occur at incipient/moderate eutrophication, we show the blooms have a positive effect on epifaunal abundance under well-oxygenated conditions, but a negative effect if pervasive anoxic/hypoxic conditions develop with the bloom. These findings provide support to prior suggestions that secondary productivity in shallow coastal systems may increase as seagrasses get replaced by loose macroalgal stands if the stands remain well oxygenated. In concert, our results contribute to an improvement of our current model of eutrophication of shallow coastal systems and suggest that further effort should be put on ascertaining the mechanisms that may prevent severe water column shading and large macroalgal decline at intense eutrophication, as well as thorough documentation of the impacts of anoxic/hypoxic conditions on system functionality at different stages of eutrophication.

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

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 València 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 effect of macrophyte assemblages on environmental phytoplankton communities. Three experiments were carried out using spring communities: in Experiment 1, a diverse phytoplankton assemblage (29% of biomass Chlorophyceae, 26% Cryptophyceae, 19% diatoms and 9% cyanobacteria) was cultivated with exudates from monocultures of C. hispida, N. hyalina and M. spicatum. Experiment 2 proved the allelopathic effect of a macrophyte assemblage exudate (C. hispida, C. baltica, C. vulgaris, N. hyalina and M. spicatum in a mixed culture) on two different phytoplankton communities (one diverse: 53% biomass of diatoms, 27% cyanobacteria, 18% chlorophytes and another dominated by small chlorophytes). The response variables were Chl a concentration, phytoplankton biovolume and main algal groups’ biovolumes. When phytoplankton grew in water with exudates from monocultures, microalgal biomass was from 4 to 6 times lower than in the control after 5 days and Chl a concentration was up to 4 times lower. The inhibitory effect of C. hispida was greater than that of M. spicatum. Mixed macrophyte assemblages resulted in even stronger allelopathic effects than monocultured macrophytes; the biomass was reduced by 7 fold after 5 days using the mixed exudates and Chl a concentration was between 3 and 5 times lower. The experiments demonstrate that macrophytes are particularly effective in inhibiting the growth of both small diatoms and the least desirable phytoplankton component in these wetlands (filamentous cyanobacteria), but not chlorophytes (reduction by 37–69, 7–14 and 1–7 fold for diatoms, cyanobacteria and chlorophytes, respectively). The predictions are that spring macrophytes might enhance microalgae that are suitable for grazing (mostly small chlorophytes) and will decrease non-edible filamentous taxa. Thus, restoration managers should replant with the mixture of submerged native macrophytes that provide the most harmful allelopathic effects to promote benefits on aquatic communities in two synergistic ways: by directly reducing microalgal biomass and by indirectly enhancing grazing, which in turn, would promote a clear-water phase. We present replanted macrophytes as ecosystem engineers, i.e. organisms that directly or indirectly modulate the availability of resources for the wetland food webs, by causing changes in primary producer assemblages.

Regional‐scale effects of eutrophication on ecosystem structure and services of seagrass beds

Using large‐scale field surveys across 12 estuaries in two provinces in Atlantic Canada, we analyzed changes in phytoplankton and benthic macroalgal communities as well as the canopy structure of eelgrass beds and quantified their carbon and nitrogen storage with increasing eutrophication. As eutrophication increased, phytoplankton biomass increased on average 1.8 times and phaeopigments doubled. Among macroalgae, the epiphytic Ulothrix speciosa increased 40 times in New Brunswick, and benthic Ulva lactuca 670 times in Prince Edward Island covering 61% of the bottom. Eelgrass showed a significant increase in leaf length and declines in shoot density and aboveground and belowground biomass, consistent with increased shading by opportunistic algae. As eelgrass biomass declined, so did the carbon storage capacity of the habitat. Nitrogen storage only declined in belowground eelgrass beds due to increasing tissue nitrogen content above ground with eutrophication. Despite province‐ and species‐specific responses of primary producers to nutrient loading, principal component analysis revealed an overall shift from perennial eelgrass to opportunistic macroalgae and phytoplankton with eutrophication at the regional scale, indicating generalized eutrophication effects on primary producer assemblages.

Effects of temperature regime on primary producers in Icelandic geothermal streams

Eight adjacent sub-arctic streams with consistently different temperatures but broadly similar chemistry, other than some differences in conductivity and linked pH, were used to investigate effects of temperature regime on the structure of primary producer communities. Grazing by invertebrates was also taken into account to detect possible effects on the primary producers. The moss species Fontinalis antipyretica Hedw. (Bryophyta) was only observed in the warmer streams and its cover was positively linked with temperature. The liverwort Jungermannia exsertifolia (Steph.) was absent from the warmer streams and scarce in the colder streams. The biomass of F. antipyretica was inversely correlated with concentration of epilithic chlorophyll a, possibly due to shading from the bryophyte and/or competition between algae and bryophytes for nutrients. Epilithic algae and bryophyte biomass were not related to density of the main grazer, Radix peregra Müller (Gastropoda). A high biovolume of green algae (Chlorophyta) occurred in all streams. The biovolume Cyanobacteria was greatest in the coldest stream and in the two intermediately warm streams. Diatom biovolume was low compared with the biovolumes of Cyanobacteria and green algae in all streams, except in the two warmest streams, where the highest biovolume of diatom occurred. Regularized Canonical Correlation Analysis (RCCA) classification based on all primary producer assemblages and invertebrates indicated that warm streams (16.7–22.2 °C) were similar to each other. It also showed that the colder streams (6.6–13.2 °C) were different from the warmer streams and from each other because of differences in conductivity. Temperature and conductivity were the most influential variables in determining overall stream diversity.

The response of primary producer assemblages to mitigation measures to reduce eutrophication in a temperate estuary

The Mondego estuary is a well-described system located on the North Atlantic Ocean, where cultural eutrophication progressed over the last decades of the 20th century. Consequently, and due to a large productivity of Ulva spp., Zostera noltii meadows were severely reduced with a concomitant decrease in environmental quality. In 1998, experimental mitigation measures were implemented, via changes in hydrology to increase circulation and diversion of nutrient-rich freshwater inflow, to reverse the process in the most affected area of the estuary – its South arm. The objective of this study was to assess the differences in response of primary producer assemblages to the implemented measures to reduce eutrophication. Results show that the mean concentrations of DIN suffered a notorious decrease due to a significant reduction in the ammonium concentration in the water column, while DIP increased significantly. Primary producer assemblages showed different responses to these changes: phytoplankton, measured as concentration of chlorophyll a, did not show any significant changes; green macroalgae, mostly Ulva spp., suffered a large reduction in biomass, whereas Gracilaria gracilis and the macrophyte Zostera noltii biomasses increased greatly. Results show that phytoplankton biomass has remained constant and suggest that the reduction in ammonium could have been responsible for the changes in the green macroalgal biomass. Light was the most likely factor in the response of seagrass whereas red macroalgal reaction seemed to be dependent on both light and ammonium.

Molecular indicators for palaeoenvironmental change in a Messinian evaporitic sequence (Vena del Gesso, Italy). II: High-resolution variations in abundances and 13C contents of free and sulphur-bound carbon skeletons in a single marl bed

The extractable organic matter of 10 immature samples from a marl bed of one evaporitic cycle of the Vena del Gesso sediments (Gessoso-solfifera Fm., Messinian, Italy) was analyzed quantitatively for free hydrocarbons and organic sulphur compounds. Nickel boride was used as a desulphurizing agent to recover sulphur-bound lipids from the polar and asphaltene fractions. Carbon isotopic compositions (δ vs PDB) of free hydrocarbons and of S-bound hydrocarbons were also measured. Relationships between these carbon skeletons, precursor biolipids, and the organisms producing them could then be examined. Concentrations of S-bound lipids and free hydrocarbons and their δ values were plotted vs depth in the marl bed and the profiles were interpreted in terms of variations in source organisms, 13C contents of the carbon source, and environmentally induced changes in isotopic fractionation. The overall range of δ values measured was 24.7‰, from −11.6‰ for a component derived from green sulphur bacteria (Chlorobiaceae) to −36.3‰ for a lipid derived from purple sulphur bacteria (Chromatiaceae). Deconvolution of mixtures of components deriving from multiple sources (green and purple sulphur bacteria, coccolithophorids, microalgae and higher plants) was sometimes possible because both quantitative and isotopic data were available and because either the free or S-bound pool sometimes appeared to contain material from a single source. Several free n-alkanes and S-bound lipids appeared to be specific products of upper-water-column primary producers (i.e. algae and cyanobacteria). Others derived from anaerobic photoautotrophs and from heterotrophic protozoa (ciliates), which apparently fed partly on Chlorobiaceae. Four groups of n-alkanes produced by algae or cyanobacteria were also recognized based on systematic variations of abundance and isotopic composition with depth. For hydrocarbons probably derived from microalgae, isotopic variations are well correlated with those of total organic carbon. A resistant aliphatic biomacromolecule produced by microalgae is, therefore, probably an important component of the kerogen. These variations reflect changes in the depositional environment and early diagenetic transformations. Changes in the concentrations of S-bound lipids induced by variations in conditions favourable for sulphurization were discriminated from those related to variations in primary producer assemblages. The water column of the lagoonal basin was stratified and photic zone anoxia occurred during the early and middle stages of marl deposition. During the last stage of the marl deposition the stratification collapsed due to a significant shallowing of the water column. Contributions from anaerobic photoautotrophs were apparently associated with variations in depth of the chemocline.