Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs,with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an “individual” being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns.     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

Biomass, diversity and production of rocky shore macroalgae at two nutrient enrichment and wave action levels

The littoral zone of temperate rocky shores is normally dominated by perennial macroalgae (e.g. Fucus, Ascophyllum, Laminaria), but nutrient enrichment and/or permanently decreased wave action may lead to structural community changes from dominance of perennials to increased amounts of annual opportunistic species (mainly green algae). Macroalgal biomass, diversity and production as well as relationships between the two latter were studied using Solbergstrand’s rocky shore mesocosms in SE Norway in connection with a long-term experimental manipulation of nutrient addition and wave action (high and low levels of both factors applied in a crossed way to eight outdoor basins). After more than 2 years of experimental treatment, the total standing stock of macroalgae was larger in low nutrient than in high nutrient treatments as well as in high wave compared to low wave treatments (in autumn only). For macroalgal functional groups, bushy and filamentous brown and filamentous red algae were generally favoured by low nutrient concentrations, while annual filamentous and sheet-like green algae were stimulated by the nutrient enrichment. There was only one significant interaction between nutrient enrichment and wave action (for brown filamentous algae in autumn) and also only one significant main effect of the wave treatment (for bushy brown algae in autumn). Surprisingly, the high nutrient treatments supported a higher diversity of macroalgae, whereas the low nutrient treatments generally showed higher production rates. Moreover, significantly negative correlations were found between macroalgal diversity and primary productivity in both summer and autumn. This study shows that it is the biological components of the communities subjected to external forcing (nutrient addition or decreased wave action) that regulate production and this contradicts the common misperception that resource production in natural systems simply can be fast-forwarded by fertilization. The negative relationships between diversity and productivity, although a consequence of unexpected results for diversity and production, are also novel and hint towards species identities having more important functional consequences than general species dominance patterns and the amount of species per se. These results also emphasise the context dependency of findings within the field of biodiversity and ecosystem functioning.     

The role of sponge competition on coral reef alternative steady states

Sponges constitute an abundant and functionally important component of coral reef systems. Given their demonstrated resistance to environmental stress, it might be expected that the role of sponges in reef systems under modern regimes of frequent and severe disturbance may become even more substantial. Disturbances have recently reshaped the community structure of many Caribbean coral reefs shifting them towards a state of persistent low coral cover and often a dominance of macroalgae. Using competition and growth rates recorded from Glover's Atoll in Belize, we parameterise a mathematical model used to simulate the three-way competition between sponges, macroalgae and coral. We use the model to determine the range of parameters in which each of the three species might be expected to dominate. Emergent properties arise from our simple model of this complex system, and these include a special case in which heightened competitive ability of macroalgae versus coral may counter-intuitively prove to be advantageous to the persistence of corals. Importantly, we show that even under scenarios whereby sponges fail to invade the system, inclusion of this third antagonist can qualitatively affect the likelihood of alternative stable states - generally in favour of macroalgal dominance. The interplay between multi-species competition and predation is complex, but our efforts highlight a key process that has, until now, remained unexplored: the extent to which sponges dissipate algal grazing pressure by providing generalist fish with an alternative food source. We highlight the necessity of identifying the extent by which this process takes place in tropical systems in order to improve projections of alternative stable states for Caribbean coral reefs.     

Analysis of annual fluctuations of C. nodosa in the Venice lagoon: Modeling approach

A simple simulation model was developed to describe the growth trends of Cymodocea nodosa (Ucria) Ascherson based on data sets from the Venice lagoon. The model reproduces the seasonal fluctuations in the above and belowground biomass and in shoot density. The modeling results are in good agreement with data on net production, growth rates and chemical–physical parameters of water. It was assumed that light and temperature are the most important factors controlling C. nodosa development, and that the growth was not limited by nutrient availability. The aim was to simulate biomass production as a function of external forcing variables (light, water temperature) and internal control (plant density). A series of simulation experiments were performed with the basic model showing that among the most important phenomena affecting C. nodosa growth are: (1) inhibition of production and recruitment of new shoots by high temperature and (2) light attenuation due to seasonal fluctuation.     

Simulating the temporal pattern of waste production in farmed gilthead seabream (Sparus aurata), European seabass (Dicentrarchus labrax) and Atlantic bluefin tuna (Thunnus thynnus)

Most fish farming waste output models provide gross waste rates as a function of stocked or produced biomass for a year or total culture cycle, but without contemplating the temporality of the discharges. This work aims to ascertain the temporal pattern of waste loads by coupling available growth and waste production models and developing simulation under real production rearing conditions, considering the overlapping of batches and management of stocks for three widely cultured species in the Mediterranean Sea: gilthead seabream (Sparus aurata), European seabass (Dicentrarchus labrax) and Atlantic bluefin tuna (Thunnus thynnus). For a similar annual biomass production, the simulations showed that waste output and temporal dumping patterns differ between the three species as a result of the disparities in growth velocity, nutrient digestibility, maintenance metabolic budget and husbandry. The simulations allowed the temporal patterns including the periods of maximum discharge and the dissolved and particulate nitrogen and phosphorus content in the wastes released to be determined, both of which were seen to be species-specific.     

Structure and function of a warm monomictic lake

Construction and simulation of a model of Lake Conway, Florida, U.S.A., provided a framework for defining major characteristics of this ecosystem. The relationships that are formalized in this model comprise a set of hypotheses about the nature of a warm monomictic lake. The data that were used to parameterize the model came primarily from literature estimates, although approximations of biomass levels were available from associated research conducted on the lake.Submersed macrophytes are a major biomass component in Lake Conway; simulation suggested that their role in nutrient recycling overshadows their importance in the grazing food chain. Phytoplankton biomass and degree of fluctuation are considerably lower than are observed in most cool temperate lakes, although simulated respiration and herbivory rates are closer to temperate values than tropical values. Simulated epipelic algae biomass varies an order of magnitude during the year, and this group appears to be a significant part of the food chain. Simulated zooplankton consumption and turnover rates are very high, in part because of the relatively small biomass per individual. Simulation of the model suggests that slightly more carbon is processed through the grazing food chain in Lake Conway than through the detritus food chain.     

Modelling the dynamics of coral reef macroalgae using a Bayesian belief network approach

Macroalgae are a major benthic component of coral reefs and their dynamics influence the resilience of coral reefs to disturbance. However, the relative importance of physical and ecological processes in driving macroalgal dynamics is poorly understood. Here we develop a Bayesian belief network (BBN) model to integrate many of these processes and predict the growth of coral reef macroalgae. Bayesian belief networks use probabilistic relationships rather than deterministic rules to quantify the cause and effect assumptions. The model was developed using both new empirical data and quantified relationships elicited from previous studies. We demonstrate the efficacy of the BBN to predict the dynamics of a common Caribbean macroalgal genus Dictyota. Predictions of the model have an average accuracy of 55% (implying that 55% of the predicted categories of Dictyota cover were assigned to the correct class). Sensitivity analysis suggested that macroalgal dynamics were primarily driven by top–down processes of grazing rather than bottom–up nutrification. BBNs provide a useful framework for modelling complex systems, identifying gaps in our scientific understanding and communicating the complexities of the associated uncertainties in an explicit manner to stakeholders. We anticipate that accuracies will improve as new data are added to the model.     

Herbivory by the Caribbean king crab on coral patch reefs

Caribbean coral reefs are increasingly dominated by macroalgae instead of corals due to several factors, including the decline of herbivores. Yet, virtually unknown is the role of crustacean macrograzers on coral reef macroalgae. We examined the effect of grazing by the Caribbean king crab (Mithrax spinosissimus) on coral patch reef algal communities in the Florida Keys, Florida (USA), by: (1) measuring crab selectivity and consumption of macroalgae, (2) estimating crab density, and (3) comparing the effect of crab herbivory to that of fishes. Mithrax prefers fleshy macroalgae, but it also consumes relatively unpalatable calcareous algae. Per capita grazing rates by Mithrax exceed those of most herbivorous fish, but Mithrax often occurs at low densities on reefs and its foraging activities are reduced in predator-rich environments. Therefore, the effects of grazing by Mithrax tend to be localized and when at low density contribute primarily to spatial heterogeneity in coral reef macroalgal communities.     

Effects of local fisheries and ocean productivity on the northeastern Ionian Sea ecosystem

To better understand the effects of fisheries and ocean productivity on the northeastern Ionian Sea we constructed an Ecopath with Ecosim model with 22 functional groups. Data on biomass, production/biomass, consumption/biomass, and diet for each group were estimated or extrapolated from the literature. Fisheries landings and discards were also included. Temporal trajectories were simulated using Ecosim. The model was fitted with time-series data for the most important groups from 1964 to 2006. Simulations highlighted a decline of top predators and of most of the commercial species since the late 1970s. The model shows that the decline of fish resources was mainly caused by an intensive fishing pressure that occurred in the area until the end of the 1990s and also by changes in primary production that impacted the trajectories of the main functional groups. In particular, simulated changes through time in PP impacted the abundance trends of all the commercial species, showing a cascade-up effect through the ecosystem. The application of Ecopath with Ecosim was a useful tool for understanding the trends of the main functional groups of the northeastern Ionian Sea. The model underlined that management actions are needed to restore and protect target species including marine mammals, pelagic and demersal fishes. In particular, measures to reduce overfishing, illegal fishing activities and to respect existing legislations are in need. Moreover, the adoption of marine protected areas could be an effective management measure to guarantee prey survival and to sustain marine predators.     

Effects of micro- and mesograzers on intertidal macroalgal recruitment

This study examined the effects of a guild of micrograzing harpacticoid copepods (dominated by two species of Paradactylopodia sp. nov. and one species of Scutellidium sp. nov.) and a mesograzing periwinkle, Afrolittorina praetermissa, on the early recruitment of intertidal macroalgae on a wave-exposed, rocky shore. This is the first study, to our knowledge, to examine the effects of micrograzers (<500 μm) on intertidal macroalgal recruitment. Data showed that microscopic harpacticoid copepods altered the assemblages and reduced the densities of several macroalgal taxa, while A. praetermissa changed the assemblages and reduced both the density and number of macroalgal taxa. Recruitment of encrusting coralline algae was actually higher in copepod inclusions than exclusions, suggesting that copepods may be beneficial to the recruitment of this algal group. These results contribute to the understanding of grazing as a factor causing high mortality of algal recruits, but also highlight the need for more studies that examine the effects of micro- and mesograzers on the distribution and abundance of macroalgae.     

Influences of canopy structure and physiological traits on flux partitioning between understory and overstory in an eastern Siberian boreal larch forest

Boreal forests play an important role in the global balance of energy and CO₂. Our previous study of elaborate eddy covariance observations in a Siberian boreal larch forest, conducted both above the forest canopy and at the forest floor, revealed a significant contribution of latent heat flux (LE) from the cowberry understory to the whole ecosystem LE. Thus, in the present study, we examined what factors control the partitioning of whole ecosystem LE and CO₂ flux into the understory and overstory vegetation, using detailed leaf-level physiology (for both understory and overstory vegetation) and soil respiration property measurements as well as a multilayer soil-vegetation-atmosphere transfer (SVAT) model. The modeling results showed that the larch overstory's leaf area index (LAI) and vertical profile of leaf photosynthetic capacity were major factors determining the flux partitioning in this boreal forest ecosystem. This is unlike other forest ecosystems that tend to have dense LAI. We concluded that control of the larch overstory's LAI had a relationship with both the coexistence of the larch with the cowberry understory and with the water resources available to the total forest ecosystem.     

Individual-based modeling of flooding and salinity effects on a coastal swamp forest

Coastal swamps are among the rapidly vanishing wetland habitats in Louisiana. Increased flooding, nutrient and sediment deprivation, and salt-water intrusion have been implicated as probable causes of the decline of coastal swamps. We developed a two-species individual-based forest succession model to compare the growth and composition of a cypress-tupelo swamp under various combinations of flooding intensity and salinity levels, using historical time-series of stage and salinity data as inputs. Our model simulates forest succession over 500 years by representing the growth, mortality, and reproduction of individual Taxodium distichum (baldcypress) and Nyssa aquatica (water tupelo) trees in a 1-km² spatial grid of 10 m × 10 m cells that vary in water levels and salinity through differences in elevation. We independently adjusted the elevations of each cell to obtain different grid-wide mean elevations and standard deviations of elevation; this affected the temporal and spatial pattern of flooding. We calibrated the model by adjusting selected parameters until averaged basal area, stem density and wood production rates under two different mean elevations (partially versus highly flooded) were qualitatively similar to comparable values reported for swamps in the literature. Corroboration involved comparing model predictions to four well-monitored contrasting habitat sites within the Maurepas Basin, Louisiana, USA. Model predictions of both species combined showed the same patterns among sites as the data, but the model overestimated wood production and the dominance of T. distichum. Exploratory simulations predicted that increased flooding leads to swamps with reduced basal areas and stem densities, while increased salinity resulted in lower basal areas at low salinity concentration (∼1-3 psu) and complete tree mortality at higher salinity concentrations (∼2-6 psu). Our model can provide insight into the succession dynamics of coastal swamps and information for the effective design of restoration actions.     

Simulation modeling to understand how selective foraging by beaver can drive the structure and function of a willow community

Beaver–willow (Castor-Salix) communities are a unique and vital component of healthy wetlands throughout the Holarctic region. Beaver selectively forage willow to provide fresh food, stored winter food, and construction material. The effects of this complex foraging behavior on the structure and function of willow communities is poorly understood. Simulation modeling may help ecologists understand these complex interactions. In this study, a modified version of the SAVANNA ecosystem model was developed to better understand how beaver foraging affects the structure and function of a willow community in a simulated riparian ecosystem in Rocky Mountain National Park, Colorado (RMNP). The model represents willow in terms of plant and stem dynamics and beaver foraging in terms of the quantity and quality of stems cut to meet the energetic and life history requirements of beaver. Given a site where all stems were equally available, the model suggested a simulated beaver family of 2 adults, 2 yearlings, and 2 kits required a minimum of 4 ha of willow (containing about10 stems m⁻²) to persist in a steady-state condition. Beaver created a willow community where the annual net primary productivity (ANPP) was 2 times higher and plant architecture was more diverse than the willow community without beaver. Beaver foraging created a plant architecture dominated by medium size willow plants, which likely explains how beaver can increase ANPP. Long-term simulations suggested that woody biomass stabilized at similar values even though availability differed greatly at initial condition. Simulations also suggested that willow ANPP increased across a range of beaver densities until beaver became food limited. Thus, selective foraging by beaver increased productivity, decreased biomass, and increased structural heterogeneity in a simulated willow community.     

Mathematical analysis of change in forest carbon use efficiency with stand development: A case study on Abies veitchii Lindl.

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were analyzed for Abies veitchii Lindl. forests with respect to stand development by developing a simple mathematical model incorporating data on physiological variables and leaf mass ratio. A decrease in CUE with stand development was successfully expressed as a function of stand biomass (y) based on the following three assumptions: (1) a power-law relationship between mean respiration and mean individual tree mass, (2) a power-functional relationship between mean gross primary production and mean individual tree mass, and (3) self-thinning relationship between stand biomass and density. Based on this model, a parameter of CUE–y relationship was defined, and it was clarified that CUE decrease with stand development is caused not by the ratio of specific respiration rate to specific gross photosynthetic rate, but by leaf mass ratio. Since CUE is high in young forests, helpful information on selecting woody species when planting seedlings was provided from the viewpoints of reducing CO₂ in the atmosphere and global warming.     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

Stability of sea urchin dominated barren grounds following destructive grazing of kelp in St. Margaret's Bay,Eastern Canada

Kelp regeneration was observed for the first time in St. Margaret's Bay, Nova Scotia, Canada in an area known to have been devoid of macroalgae for several years. The regeneration was destroyed by sea urchins within 10 months. Experimentally induced kelp regeneration met a similar fate under normal grazing pressure. At the lowest sea urchin biomass and density encountered re-establishment of mature kelp stands seems highly unlikely. The sea urchin population required to suppress kelp regeneration is fed by benthic microalgae. Diatoms and other pioneer algal community species were found in the guts of sea urchins. The mean standing crop of benthic microalgae was found to be, 2.2 g C m^-2 and production estimated as ca 15 g C m^-2yr^-1 at 8m depth. Most of the primary production of St. Margaret's Bay has been lost with the disappearing kelp populations.     

Modeling approach to regime shifts of primary production in shallow coastal ecosystems

Pristine coastal shallow systems are usually dominated by extensive meadows of seagrass species, which are assumed to take advantage of nutrient supply from sediment. An increasing nutrient input is thought to favour phytoplankton, epiphytic microalgae, as well as opportunistic ephemeral macroalgae that coexist with seagrasses. The primary cause of shifts and succession in the macrophyte community is the increase of nutrient load to water; however, temperature plays also an important role. A competition model between rooted seagrass (Zostera marina), macroalgae (Ulva sp.), and phytoplankton has been developed to analyse the succession of primary producer communities in these systems. Successions of dominance states, with different resilience characteristics, are found when modifying the input of nutrients and the seasonal temperature and light intensity forcing.     

A spatial model of phytoplankton patchiness

The one-dimensional theory of critical-length scales of phytoplankton patchiness is developed to include phytoplankton growth and herbivore grazing as functions of time and space. The critical-length scale L _c for the pathch is then determined by the initial spatial distribution and concentration of the limiting nutrient and herbivores in addition to the daily averaged values of the growth and loss processes. The response of an initial phytoplankton patch to the stresses of turbulent diffusion, nutrient depletion, light periodicity, and nocturnal or continuous herbivore grazing is investigated numerically for several oceanic conditions. Nocturnal grazing, while less stressful on primary production than continous grazing, results in lower phytoplankton standing stocks. Increase in biomass of vertically migrating zooplankton results in a net loss of nutrient which might otherwise be egested, recycled, and utilized in the euphotic zone under continuous grazing conditions. The Ivlev constant is shown via sensitivity analysis to be a significant parameter ultimately influencing phytoplankton production. It is demonstrated numerically that diffusion of phytoplankton cells from areas of high concentration to low concentration prevents the local extinction of the standing stock, thereby rendering a positive herbivore grazing-threshold unnecessary for ecosystem stability.