Christophe Lett

Short communication: A Lagrangian tool for modelling ichthyoplankton dynamics

Ichthyop is a free Java tool designed to study the effects of physical and biological factors on ichthyoplankton dynamics. It incorporates the most important processes involved in fish early life: spawning, movement, growth, mortality and recruitment. The tool uses as input time series of velocity, temperature and salinity fields archived from ROMS or MARS oceanic models. It runs with a user-friendly graphic interface and generates output files that can be post-processed easily using graphic and statistical software.

Factors affecting information transfer from knowledgeable to naive individuals in groups

There is evidence that individuals in animal groups benefit from the presence of knowledgeable group members in different ways. Experiments and computer simulations have shown that a few individuals within a group can lead others, for a precise task and at a specific moment. As a group travels, different individuals possessing a particular knowledge may act as temporary leaders, so that the group will, as a whole, follow their behaviour. In this paper, we use a model to study different factors influencing group response to temporary leadership. The model is based on four individual behaviours. Three of those, attraction, repulsion, and alignment, are shared by all individuals. The last one, attraction toward the source of a stimulus, concerns only a fraction of the group members. We explore the influence of group size, proportion of stimulated individuals, number of influential neighbours, and intensity of the attraction to the source of the stimulus, on the proportion of the group reaching this source. Special attention is given to the simulation of large group size, close to those observed in nature. Groups of 100, 400 and 900 individuals are currently simulated, and up to 8,000 in one experiment. We show that more stimulated individuals and a larger group size both induce the arrival of a larger fraction of the group. The number of influential neighbours and the intensity of the stimulus have a non-linear influence on the proportion of the group arrival, displaying first a positive relationship and then, above a given threshold, a negative one. We conclude that an intermediate level of group cohesion provides optimal transfer information from knowledgeable to naive individuals.

Effects of successive predator attacks on prey aggregations

We study the cumulative effect of successive predator attacks on the disturbance of a prey aggregation using a modelling approach. Our model intends to represent fish schools attacked by both aerial and underwater predators. This individual-based model uses long-distance attraction and short-distance repulsion between prey, which leads to prey aggregation and swarming in the absence of predators. When intermediate-distance alignment is added to the model, the prey aggregation displays a cohesive displacement, i.e., schooling, instead of swarming. Including predators, i.e. with repulsion behaviour for prey to predators in the model, leads to flash expansion of the prey aggregation after a predator attack. When several predators attack successively, the prey aggregation dynamics is a succession of expanding-grouping-swarming/schooling phases. We quantify this dynamics by recording the changes in the simulated prey aggregation radius over time. This radius is computed as the longest distance of individual prey to the aggregation centroid, and it is assumed to increase along with prey disturbance. The prey aggregation radius generally increases during flash expansion, then decreases during grouping until reaching a constant lowest level during swarming/schooling. This general dynamics is modulated by several parameters: the frequency, direction (vertical vs. horizontal) and target (centroid of the prey aggregation vs. random prey) of predator attacks; the distance at which prey detect predators; the number of prey and predators. Our results suggest that both aerial and underwater predators are more efficient at disturbing fish schools by increasing their attack frequency at such level that the fish cannot return to swarming/schooling. We find that a mix between aerial and underwater predators is more efficient at disturbing a fish school than a single type of attack, suggesting that aerial and underwater foragers may gain mutual benefits in forming foraging groups.

Linking local retention, self-recruitment, and persistence in marine metapopulations.

Three indices of larval retention have been used in the literature to assess the tendency for self-maintenance of local marine populations: local retention (LR), self-recruitment (SR), and relative local retention (RLR). Only one of these, LR, defined as the ratio of locally produced settlement to local egg production, has a clear relationship to self-persistence of individual sites. However, SR, the ratio of locally produced settlement to settlement of all origins at a site, is generally easier to measure experimentally. We use theoretical, simulation, and empirical approaches to bridge the gap between these different indices, and demonstrate that there is a proportional relationship between SR and LR, for metapopulations close to a stable state and with lifetime egg production (LEP) approximately uniform over space. Similarly, for systems where larval mortality rates are a relatively uniform function of release site, RLR (defined as the ratio of locally produced settlement to all settlement of local origin) and LR will also be proportional. Therefore, SR and RLR provide information on relative rates of LR for systems satisfying these conditions. Furthermore, the ratio between LR and SR can be used to evaluate global persistence of metapopulations, and therefore provides valuable information not necessarily available if only LR is considered.

OPTIMAL NUMBER OF SITES IN MULTI-SITE FISHERIES WITH FISH STOCK DEPENDENT MIGRATIONS

We present a stock-effort dynamical model of a fishery subdivided into fishing zones. The stock corresponds to a fish population moving between these zones, on which they are harvested by fishing fleets. We consider a linear chain of identical fishing zones. Fish movements between the zones, as well as vessels displacements, are assumed to take place at a faster time scale than the variation of the stock and the change of the fleet size. The vessels movements between the fishing areas are assumed to be stock dependent, i.e. the larger the stock density is in a zone the more vessels tends to remain in it. We take advantage of these two time scales to derive a reduced model governing the dynamics of the total harvested stock and the total fishing effort. Under some assumption, we obtain either a stable equilibrium or a stable limit cycle which involves large cyclic variations of the total fish stock and fishing effort. We show that there exists an optimal number of fishing zones that maximizes the total catch at equilibrium. We discuss the results in relation to fish aggregating devices (FADs) fisheries.

Spatial characterisation of the Benguela ecosystem for ecosystem-based management

The three countries of the Benguela Current Large Marine Ecosystem (BCLME), namely Angola, Namibia and South Africa, have committed to implementing ecosystem-based management (EBM) including an ecosystem approach to fisheries (EAF) in the region, to put in practice the principles of sustainable development in ocean-related matters. There is also recognition of the need for marine spatial planning (MSP) as a process for informing EBM with regard to the allocation and siting of ocean uses so that ecosystem health is ensured and trade-offs between ecosystem services are appropriately dealt with. Marine spatial planning is both an integrated and an area-based process, and this paper produces a spatial characterisation of the BCLME for achieving a common basis for MSP in the region, focusing on the oceanography, biology and fisheries. Recognising spatial variation in physical driving forces, primary and secondary production, trophic structures and species richness, four different subsystems are characterised: (1) north of the Angola–Benguela Front, (2) from the Angola–Benguela Front to Lüderitz, (3) from Lüderitz to Cape Agulhas, and (4) from Cape Agulhas to Port Alfred on the south-east coast of South Africa. Research and monitoring requirements of relevance for MSP and EBM in the region are identified, focusing on understanding variability and change, including with regard to the boundary areas identified for the system. To this end, 14 cross-shelf monitoring transects are proposed (including seven that are already being monitored) to estimate fluxes of biota, energy and materials within and between the subsystems. The usefulness of models for understanding ecosystem variability and changes is recognised and the need for fine-scale resolution of both sampling and modelling for adequate MSP as input to EBM for the often-conflicting interests of conserving biodiversity, and managing fisheries, recreation, offshore oil and gas exploration and exploitation, offshore mining and shipping routes, is emphasised.

Modelling the effect of food availability on recruitment success of Cape anchovy ichthyoplankton in the southern Benguela upwelling system

Cape anchovy Engraulis encrasicolus adapted its reproductive strategies to the southern Benguela system by spawning over the Agulhas Bank, an area of low productivity that is located upstream of the predominant upwelling system. Frontal jet currents transport eggs and larvae toward the west coast of South Africa, where recruitment takes place. To characterise the recruitment dynamics of Cape anchovy ichthyoplankton, we used an individual-based model forced by a coupled hydrodynamic–biogeochemical model. The results show the importance of food (especially diatoms and copepods) dynamics on the spatial and temporal patterns of recruitment success, and also confirm the importance of the spawning area, timing and water depth on the recruitment success of Cape anchovy larvae.

Influence of Biological Factors on Connectivity Patterns for Concholepas concholepas (loco) in Chile

In marine benthic ecosystems, larval connectivity is a major process influencing the maintenance and distribution of invertebrate populations. Larval connectivity is a complex process to study as it is determined by several interacting factors. Here we use an individual-based, biophysical model, to disentangle the effects of such factors, namely larval vertical migration, larval growth, larval mortality, adults fecundity, and habitat availability, for the marine gastropod Concholepas concholepas (loco) in Chile. Lower transport success and higher dispersal distances are observed including larval vertical migration in the model. We find an overall decrease in larval transport success to settlement areas from northern to southern Chile. This spatial gradient results from the combination of current direction and intensity, seawater temperature, and available habitat. From our simulated connectivity patterns we then identify subpopulations of loco along the Chilean coast, which could serve as a basis for spatial management of this resource in the future.

Use of Lagrangian simulations to hindcast the geographical position of propagule release zones in a Mediterranean coastal fish

The study of organism dispersal is fundamental for elucidating patterns of connectivity between populations, thus crucial for the design of effective protection and management strategies. This is especially challenging in the case of coastal fish, for which information on egg release zones (i.e. spawning grounds) is often lacking. Here we assessed the putative location of egg release zones of the saddled sea bream (Oblada melanura) along the south-eastern coast of Spain in 2013. To this aim, we hindcasted propagule (egg and larva) dispersal using Lagrangian simulations, fed with species-specific information on early life history traits (ELTs), with two approaches: 1) back-tracking and 2) comparing settler distribution obtained from simulations to the analogous distribution resulting from otolith chemical analysis. Simulations were also used to assess which factors contributed the most to dispersal distances. Back-tracking simulations indicated that both the northern sector of the Murcia region and some traits of the North-African coast were hydrodynamically suitable to generate and drive the supply of larvae recorded along the coast of Murcia in 2013. With the second approach, based on the correlation between simulation outputs and field results (otolith chemical analysis), we found that the oceanographic characteristics of the study area could have determined the pattern of settler distribution recorded with otolith analysis in 2013 and inferred the geographical position of main O. melanura spawning grounds along the coast. Dispersal distance was found to be significantly affected by the geographical position of propagule release zones. The combination of methods used was the first attempt to assess the geographical position of propagule release zones in the Mediterranean Sea for O. melanura, and can represent a valuable approach for elucidating dispersal and connectivity patterns in other coastal species.

Biophysical models of larval dispersal in the Benguela Current ecosystem

We synthesise and update results from the suite of biophysical, larval-dispersal models developed in the Benguela Current ecosystem. Biophysical models of larval dispersal use outputs of physical hydrodynamic models as inputs to individual-based models in which biological processes acting during the larval life are included. In the Benguela, such models were first applied to simulate the dispersal of anchovy Engraulis encrasicolus and sardine Sardinops sagax ichthyoplankton, and more recently of the early life stages of chokka-squid Loligo reynaudii and Cape hakes Merluccius spp. We identify how the models have helped advance understanding of key processes for these species. We then discuss which aspects of the early life of marine species in the Benguela Current ecosystem are still not well understood and could benefit from new modelling studies.