Jean-Marc Guarini

Dynamic behaviour of benthic microalgal biomass in intertidal mudflats

The dynamic behaviour of microphytobenthic biomass in a European intertidal mudflat has been analysed by comparing field and laboratory measurements. In an experimental mesocosm, where the effects of grazing by deposit-feeders and resuspension by tides had been significantly decreased, the benthic microalgal biomass followed a logistic-type growth curve, and thus converged towards a maximum value at which production is theoretically equal to zero. In the field, the kinetics of microalgal biomass—measured during 14 consecutive days at the beginning and at the end of every daytime exposures—exhibited a different pattern with biomass increases during daytime exposures and biomass decreases during the other periods, thus describing a series of oscillations. It is suggested that in the field, the effect of grazing and resuspension prevents biomass from holding steady at its maximum level. Moreover, it has been found a significant negative relationship which states that the higher the biomass level at the beginning of daytime exposures, the lower the net production during that daytime exposure. In other words, it turned out that in the field, the biomass also tends to converge towards a “field maximum” where net production is equal to zero. Taken together, these observations allow to hypothesize that the high productivity of the microphytobenthic community in intertidal mudflats is due to the tight coupling between physical and biological processes.

SEASONAL EFFECT ON THE RELATIONSHIP BETWEEN THE PHOTOSYNTHETIC CAPACITY OF INTERTIDAL MICROPHYTOBENTHOS AND TEMPERATURE1

The response of the photosynthetic capacity (Pmax) of microphytobenthos to short‐term variations of temperature (in the range 5–35° C) was assessed on a seasonal basis. The relationship is described mathematically, and relevant physiological parameters are identified: PMAX, the maximum value of Pmax achieved at Topl, the optimum temperature. Estimated values of Topt do not change significantly throughout the year and remain close to 25° C. It is thus concluded that Topt is not influenced by seasonal variations in the daily range of mud surface temperature. Identical conclusions hold for Tmax (ca. 38° C), the thermal threshold beyond which no photosynthesis occurs. Conversely, PMA estimates exhibit substantial variability: PMAX (mean ± root mean square error) is highest in April (11.18 ± 0.42 [μg C · [μg Chl a]−1· h−1) during the beginning of the annual increase in temperature, photoperiod, and maximum irradiance and is lowest in December (3.04 ± 0.16 μg C · [μg Chl a]−1· h−l). From an ecological point of view, the short‐term and seasonal variations of PMAX suggest that the microphytobenthic community takes advantage of the abiotic spring environmental conditions, allowing the onset of the bloom. Nevertheless, no “acclimation strategy” (i.e. shifts in Topt and Tmax that prevent temperature inhibition in summer or improve photosynthetic rates in winter) is apparent from our results.

Measurement of ingestion rate of Hydrobia ulvae (Pennant) on intertidal epipelic microalgae: the effect of mud snail density

The individual mean ingestion rate of Hydrobia ulvae was measured experimentally in controlled microcosms, in the dark to avoid primary production during measurement and at constant temperature. The experimental design was based on the addition of prelabelled epipelic microalgae to microcosms in a constant proportion with unlabelled diatoms, and in such a way that algal food availability was not a limiting factor within the range of tested densities (0.3 to 4.1 snails cm(-2)). Results show that the individual mean ingestion rate decreased significantly from 26.6+/-1.1 ng Chl a snail(-1) h(-1) to 22.4+/-1.0 ng Chl a snail(-1) h(-1) between 0.7 and 3 snails cm(-2). We hypothesize that this sharp decrease (the threshold density was between 1.4 and 2.5 snails cm(-2)) may account for a density-dependent effect. We have tested this hypothesis by using a simple random walk model including basic behavioural processes such as a break in feeding activity when two individuals contact each other. The model represents quantitatively well the threshold effect, suggesting that behavioural processes have to be taken into account for estimating a global feeding activity of H. ulvae populations.

CHARACTERIZING AND QUANTIFYING PHOTOINHIBITION IN INTERTIDAL MICROPHYTOBENTHOS1

This study characterizes the short‐term influence of the sustained saturating irradiance encountered by the microphytobenthos inhabiting intertidal mudflats. The kinetics of photoinhibition in epipelic microalgae from intertidal mudflats were investigated in the laboratory. Previously isolated benthic microalgae were exposed to a saturating photon flux density (PFD) for periods ranging from 0 to 180 min; every 30 min, a photosynthesis‐irradiance curve was established to quantify the effect of the saturating PFD on both parameters αB, the photosynthetic efficiency, and PmB, the photosynthetic capacity. The αB decreased from the beginning of light exposure until the end, whereas PmB first slightly increased and then diminished from 90 min exposure onward. It turned out that epipelic microphytobenthos undergoes photoinhibition after about 90 min of saturating PFD. The possible ecological consequences of these ecophysiological results are discussed.

Microphytobenthic Potential Productivity Estimated in Three Tidal Embayments of the San Francisco Bay: A Comparative Study

In this paper we describe a three-step procedure to infer the spatial heterogeneity in microphytobenthos primary productivity at the scale of tidal estuaries and embayments. The first step involves local measurement of the carbon assimilation rate of benthic microalgae to determine the parameters of the photosynthesis-irradiance (P-E) curves (using non-linear optimization methods). In the next step, a resampling technique is used to rebuild pseudo-sampling distributions of the local productivity estimates; these provide error estimates for determining the significance level of differences between sites. The third step combines the previous results with deterministic models of tidal elevation and solar irradiance to compute mean and variance of the daily areal primary productivity over an entire intertidal mudflat area within each embayment. This scheme was applied on three different intertidal mudflat regions of the San Francisco Bay estuary during autumn 1998. Microphytobenthos productivity exhibits strong (ca. 3-fold) significant differences among the major sub-basins of San Francisco Bay. This spatial heterogeneity is attributed to two main causes: significant differences in the photosynthetic competence (P-E parameters) of the microphytobenthos in the different sub-basins, and spatial differences in the phase shifts between the tidal and solar cycles controlling the exposure of intertidal areas to sunlight. The procedure is general and can be used in other estuaries to assess the magnitude and patterns of spatial variability of microphytobenthos productivity at the level of the ecosystems.

Contrôle de la dynamique à court terme du microphytobenthos intertidal par le cycle exondation-submersion

Abstract We test herein the hypothesis that the succession of low tides — during which biomass increases — and high tides — during which biomass decreases — controls the dynamic behaviour of the microphytobenthic compartment. Results indeed show that during a complete lunar cycle (14 d) biomass exhibits a series of oscillations due to biomass increases during low tides and decreases during high tides. Moreover, the datasets show that the production level is in the range of previously recorded values and they indicate that the major part of the photosynthetically produced biomass during low tide can be exported into the water column during high tide. This clearly suggests that microphytobenthos is the major autochtonous source of organic matter production in the littoral zone and that it directly supplies the associated pelagic trophic web.

Spatio-temporal differentiation in the population structure of Hydrobia ulvae on an intertidal mudflat (Marennes-Oleron Bay, France)

distributed in the upper half part of the mud£at and no individuals were found in the lower part.The breeding cycle extended from March to December and showed two annual peaks, in spring and in autumn.Three cohorts were recruited during the year and showed high growth rates during summer; the parameters of the von Bertalaniy model describing the growth curves were equal to kmax¼0.47 � 0.5 mm month 71 and L1¼5.4 � 0.2 mm. The snail population had similar size^ frequency structure along the transect at the beginning of the survey but summer recruitment initiated spatial diierentiation.Reproduction occurred in the middle part of the mud£at but recruits mainly settled down at the upper level of the mud£at; new cohorts appeared with increasing individual densities.The middle part of the mud£at was rather dominated by adult individuals which showed large density £uctuations.

A roadmap for a quantitative ecosystem-based environmental impact assessment

A new roadmap for quantitative methodologies of Environmental Impact Assessment (EIA) is proposed, using an ecosystem-based approach. EIA recommendations are currently based on case-by-case rankings, distant from statistical methodologies, and ecological ideas that lack proof of generality or predictive capacities. These qualitative approaches ignore process dynamics, scales of variations and interdependencies and are unable to address societal demands to link socio-economic and ecological processes (e.g. population dynamics). We propose to refocus EIA around the systemic formulation of interactions between organisms (organized in populations and communities) and their environments but inserted within a strict statistical framework. A systemic formulation allows scenarios to be built that simulate impacts on chosen receptors. To illustrate the approach, we design a minimum ecosystem model that demonstrates nontrivial effects and complex responses to environmental changes and validated with case study. We suggest that an Ecosystem-Based EIA-in which the socio-economic system is an evolving driver of the ecological one-is more promising than a socio-economic-ecological system where all variables are treated as equal. This refocuses the debate on cause-and-effect, processes, identification of essential portable variables, and allows for quantitative comparisons between projects, which is critical in cumulative effects determinations.

Estimating Muricid abundances from trapping methods used in Mediterranean Tyrian Purple industry

A new statistical method based on a stochastic dynamic model is proposed to assess population abundances of murcid species at scales relevant to both Ancient and Modern artisanal, coastal fisheries. Motivated by the long-term goal of reconstructing the dynamics of exploited murex populations during Antiquity, the objective was to quantify the population density of the banded-dye murex, Hexaplex trunculus (Linnaeus, 1758) from successive captures with baited traps, using a method similar to the technique employed in the Mediterranean purple dye industry. A stochastic model simulating cumulative captures while taking into account high variability was developed and calibrated with data acquired during a field experiment conducted on Crete Island, near Heraklion. Sampling devices were deployed in two shallow water habitats. The traps’ catchability and the Effective Area of Attraction were estimated using the individual speed and behavioural response toward the bait observed during independent laboratory experiments. The average density of H. trunculus was estimated at 2.2 ± 1.4 SE individuals per square meter, with no significant differences between seagrass and rocky habitats, respectively. The clearing time (the time to catch all individuals within reach of the trap) of the successive experiments was 84 ± 6 SE hours, on average. This means that clearing ca. 0.4 ha of subtidal area would be necessary to produce ca. 1.0 g of pure dye pigment. While the method is discussed here with respect to a particular historical context, it is generalizable to making population abundance estimates for other species such as whelks, in modern fisheries.

An inference procedure for behavioural studies combining numerical simulations, statistics and experimental results

The technical difficulties of performing underwater observation mean that marine ecologists have long relied on behavioural experiments to study reactions of marine organisms. In this article, we examine the underlying complexity of assumptions made in raceway experiments and we propose a statistical inference procedure tailored to this type of experimental protocol. As an example, experiments were performed to test if light of two different intensities affects the proximal behaviour (i.e. direct, local and immediate) of two species of crustaceans, the hermit crab (Pagurus bernhardus), and the green crab (Carcinus maenas). Individuals were collected in the vicinity of the Sven Loven Marine Center in Tjarno (Sweden). Their movements in raceways were recorded and the statistical distance between the resulting experimental distribution and a simulated null distribution was used to compare their behaviour in two situations: dim (when they were expected to feed) and bright light (when they were expected to shelter). Initial tests indicated no differences of behaviour between dim and bright light for the two species. However, when compared with the reference state (here, a null distribution) the behaviour in dim light deviates significantly from the null distribution suggesting non-random behaviour. Our results suggest that efforts should be made to understand the behaviours of the individuals of these two species to establish a comprehensive reference state as a basis for comparison. This fundamental information should be a prerequisite before implementing experiments testing how potential disturbances affect individual organisms in behavioural ecology.