A fisheries‑independent survey using longlines and drumlines, and an acoustic telemetry study, re... more A fisheries‑independent survey using longlines and drumlines, and an acoustic telemetry study, revealed that bull sharks Carcharhinus leucas and tiger sharks Galeocerdo cuvier occur throughout the year off the west coast of Reunion Island. The research, which commenced in 2011, was conducted in response to an increase in the rate of shark attacks. Juvenile, subadult and young adult tiger sharks of 111–390 cm total length (TL) were caught in all months except July; the sex ratio was 1:1 (M:F; n = 61). All bull sharks taken, except one (183 cm TL), were mature (205–329 cm TL), with a sex ratio of 0.7:1 (M:F; n = 54), and catches occurred throughout the year except in May and August, with the highest CPUE in September. Presence/absence of a total of 46 tagged tiger sharks and 36 tagged bull sharks was monitored by means of 42 acoustic receivers distributed along the west coast of the island. Tagged tiger sharks were detected in all months, with seasonal variation between sexes. Detections of males remained low throughout the year but with a peak in winter, whereas detections of females were low in May and August only and peaked in summer. Tagged bull sharks of both sexes were more abundant in winter than in summer, with females present year round. The reasons for the apparent differences in seasonality found in longline and drumline catches compared to acoustic monitoring are discussed.
The internal structure and behaviour of clupeid schools has been observed in situ during unstress... more The internal structure and behaviour of clupeid schools has been observed in situ during unstressed periods followed either by natural predation or by applying a controlled stress (a realistically painted model predator). Combined acoustic and visual (aerial and underwater) observation techniques were used. In situ observations are characterized by a high variability of fish reactions in space and time. The variability in space is obviously due to the large scale of observation which allows the simultaneous occurrence of different types of behaviour in different regions. The variability in time is probably due to a large number of physiological or environmental influences changing unexpectedly between repeated experiments. The variability of fish reaction depends on the characteristics of the source of the stimuli: some sources (e.g. predators) induce different behavioural patterns according to the number of stimuli; others (e.g. boats) induce a single pattern, varying only in intensity. Consequences for acoustic signal identification are discussed.
Journal of Experimental Marine Biology and Ecology, 2012
Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior... more Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior of fish in their environment. In this study, we characterized the spatial and temporal occupation of 37 acoustically tagged bigeye scads (Selar Crumenophthalmus) in an array of shallow Fish Aggregating Devices (FADs) to clarify the mechanism that leads fish to associate with FADs. A comparison of the number of visits and residence times exhibited by the fish at different FADs revealed a strong variability over the array of FADS, with the emergence of a leading FAD, which recorded the majority of visits and retained the fish for a longer period of time. We found diel variability in the residence times, with fish associated at daytime and exploring the array of FADs at nighttime. We demonstrated that this diel temporal pattern was amplified in the leading FAD. We identified a 24-hour periodicity for a subset of individuals aggregated to the leading FAD, thus suggesting that those fish were able to find this FAD after night excursions. The modeling of fish movements based on a Monte Carlo sampling of inter-FAD transitions revealed that the observed spatial heterogeneity in the number of visits could not be explained through simple arrayconnectivity arguments. Similarly, we demonstrated that the high residence times recorded at the leading FAD were not due to the spatial arrangement of individual fish having different associative characters. We discussed the relationships between these patterns of association with the FADs, the exploration of the FAD array and the possible effects of social interactions and environmental factors.
The study of the internal school structure and behaviour of pelagic fish provides interesting inf... more The study of the internal school structure and behaviour of pelagic fish provides interesting information in relation to acoustic surveys, especially when comparing the undisturbed structure with the structure observed beneath a vessel passing over a school. The methodological approach involves in situ observations and combines acoustic and visual (aerial and underwater) techniques. The internal school structure is heterogeneous, including vacuoles, and this structure changes when the school is overpassed by a vessel during the day. In this case compression of the upper layer of the school is observed. The influence of this school structure on the variability of the density estimation has been studied. For the subsurface schools, the usual rate of sampling may be too low for some heterogeneous schools. Other consequences of the school structure on acoustics are discussed.
Canadian Journal of Fisheries and Aquatic Sciences, 2016
Stomach contents and stable carbon and nitrogen isotope analyses (δ13C and δ15N) were used to inv... more Stomach contents and stable carbon and nitrogen isotope analyses (δ13C and δ15N) were used to investigate the trophic ecology of two apex predators, tiger sharks (Galeocerdo cuvier) and bull sharks (Carcharhinus leucas), from Reunion Island to describe their dietary habits at both the population and individual levels. In this oceanic island, the tiger and bull sharks were more piscivorous and teutophagous than noted in previous research from other localities. The δ13C values suggested that bull sharks depended on more neritic organic matter sources than tiger sharks, confirming a coastal habitat preference for bull sharks. Moreover, the total length of the bull shark influenced δ13C values, with smaller individuals being more coastal than larger individuals. All indicators suggest that there is a higher degree of similarity between individual tiger sharks compared with the more heterogeneous bull shark population, which is composed of individuals who specialize on different prey. Th...
Biases in the measurements of spatial distribution of fish schools and their consequences for sch... more Biases in the measurements of spatial distribution of fish schools and their consequences for school biomass estimates during conventional acoustic surveys are mainly due to vertical and lateral avoidance of the vessel. In this paper, we quantify school avoidance during an acoustic survey carried out from 13 to 29 May 1994 in the Catalan Sea. From a lateral multi-beam sonar the geometric characteristics (depth, length, width, height, surface, and volume) of 1268 schools were obtained. The 60 beams (1.5 15) of the sonar scanned a vertical plane from 0 to 90 , perpendicular to the vessel path within a range of 100 m. Within this plane, the projected area ensonified by the echo-sounder used aboard for acoustic evaluation was evaluated to simulate a comparison between the sonar and the echo-sounder. The results have enabled us to improve our knowledge on the vertical and lateral avoidance patterns of schools in relation to their size, external structure, and their position in the water column, and to quantify the vessel influence on biomass estimated by echo-sounder.
The size of pelagic fish schools depends on several parameters related to internal factors such a... more The size of pelagic fish schools depends on several parameters related to internal factors such as species, number of fish, fish swimming speed and physiological status and to external factors, such as hydrological factors and presence of predators. In order to better ...
Schooling fish species are conventionally subdivided into obligate and facultative species, depen... more Schooling fish species are conventionally subdivided into obligate and facultative species, depending on the proportion of the time spent in structured schools, but little is known about the factors affecting the group structure in the absence of external structuring stimuli such as food sources, risk of predators or water flow. Changes in attraction and repulsion indicators, depending on the species and the group size were explored under controlled conditions. Two species, displaying different schooling behaviours in the wild were observed: the bigeye scad Selar crumenophthalmus (Bloch) and the barred flagtail Kuhlia mugil (Forster). In the bigeye scad, the Polarity and Speed were high and stable, and the Nearest Neighbour Distance (NND) decreased when the group size increased. In contrast, for the barred flagtail, Polarity and Speed decreased according to the group size, inducing a loss of cohesion and leading to a disorganized school. The NND mean was stable whatever the group size and relatively high. This experiment indicated that the ability to polarize is first a species specific trait, rather than a property emerging from the group and led by the circumstances.
Demonstrating and quantifying the respective roles of social interactions and external stimuli go... more Demonstrating and quantifying the respective roles of social interactions and external stimuli governing fish dynamics is key to understanding fish spatial distribution. If seminal studies have contributed to our understanding of fish spatial organization in schools, little experimental information is available on fish in their natural environment, where aggregations often occur in the presence of spatial heterogeneities. Here, we applied novel modeling approaches coupled to accurate acoustic tracking for studying the dynamics of a group of gregarious fish in a heterogeneous environment. To this purpose, we acoustically tracked with submeter resolution the positions of twelve small pelagic fish (Selar crumenophthalmus) in the presence of an anchored floating object, constituting a point of attraction for several fish species. We constructed a fieldbased model for aggregated-fish dynamics, deriving effective interactions for both social and external stimuli from experiments. We tuned the model parameters that best fit the experimental data and quantified the importance of social interactions in the aggregation, providing an explanation for the spatial structure of fish aggregations found around floating objects. Our results can be generalized to other gregarious species and contexts as long as it is possible to observe the fine-scale movements of a subset of individuals. Citation: Capello M, Soria M, Cotel P, Deneubourg J-L, Dagorn L (2011) Quantifying the Interplay between Environmental and Social Effects on Aggregated-Fish Dynamics. PLoS ONE 6(12): e28109.
Collective motion phenomena in large groups of social organisms have long fascinated the observer... more Collective motion phenomena in large groups of social organisms have long fascinated the observer, especially in cases, such as bird flocks or fish schools, where large-scale highly coordinated actions emerge in the absence of obvious leaders. However, the mechanisms involved in this self-organized behavior are still poorly understood, because the individual-level interactions underlying them remain elusive. Here, we demonstrate the power of a bottom-up methodology to build models for animal group motion from data gathered at the individual scale. Using video tracks of fish shoal in a tank, we show how a careful, incremental analysis at the local scale allows for the determination of the stimulus/response function governing an individual's moving decisions. We find in particular that both positional and orientational effects are present, act upon the fish turning speed, and depend on the swimming speed, yielding a novel schooling model whose parameters are all estimated from data. Our approach also leads to identify a density-dependent effect that results in a behavioral change for the largest groups considered. This suggests that, in confined environment, the behavioral state of fish and their reaction patterns change with group size. We debate the applicability, beyond the particular case studied here, of this novel framework for deciphering interactions in moving animal groups.
The trajectories of Kuhlia mugil fish swimming freely in a tank are analyzed in order to develop ... more The trajectories of Kuhlia mugil fish swimming freely in a tank are analyzed in order to develop a model of spontaneous fish movement. The data show that K. mugil displacement is best described by turning speed and its auto-correlation. The continuous-time process governing this new kind of displacement is modelled by a stochastic differential equation of Ornstein-Uhlenbeck family: the persistent turning walker. The associated diffusive dynamics are compared to the standard persistent random walker model and we show that the resulting diffusion coefficient scales non-linearly with linear swimming speed. In order to illustrate how interactions with other fish or the environment can be added to this spontaneous movement model we quantify the effect of tank walls on the turning speed and adequately reproduce the characteristics of the observed fish trajectories.
Journal of Experimental Marine Biology and Ecology, 2012
Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior... more Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior of fish in their environment. In this study, we characterized the spatial and temporal occupation of 37 acoustically tagged bigeye scads (Selar Crumenophthalmus) in an array of shallow Fish Aggregating Devices (FADs) to clarify the mechanism that leads fish to associate with FADs. A comparison of the number of visits and residence times exhibited by the fish at different FADs revealed a strong variability over the array of FADS, with the emergence of a leading FAD, which recorded the majority of visits and retained the fish for a longer period of time. We found diel variability in the residence times, with fish associated at daytime and exploring the array of FADs at nighttime. We demonstrated that this diel temporal pattern was amplified in the leading FAD. We identified a 24-hour periodicity for a subset of individuals aggregated to the leading FAD, thus suggesting that those fish were able to find this FAD after night excursions. The modeling of fish movements based on a Monte Carlo sampling of inter-FAD transitions revealed that the observed spatial heterogeneity in the number of visits could not be explained through simple arrayconnectivity arguments. Similarly, we demonstrated that the high residence times recorded at the leading FAD were not due to the spatial arrangement of individual fish having different associative characters. We discussed the relationships between these patterns of association with the FADs, the exploration of the FAD array and the possible effects of social interactions and environmental factors.
A fisheries‑independent survey using longlines and drumlines, and an acoustic telemetry study, re... more A fisheries‑independent survey using longlines and drumlines, and an acoustic telemetry study, revealed that bull sharks Carcharhinus leucas and tiger sharks Galeocerdo cuvier occur throughout the year off the west coast of Reunion Island. The research, which commenced in 2011, was conducted in response to an increase in the rate of shark attacks. Juvenile, subadult and young adult tiger sharks of 111–390 cm total length (TL) were caught in all months except July; the sex ratio was 1:1 (M:F; n = 61). All bull sharks taken, except one (183 cm TL), were mature (205–329 cm TL), with a sex ratio of 0.7:1 (M:F; n = 54), and catches occurred throughout the year except in May and August, with the highest CPUE in September. Presence/absence of a total of 46 tagged tiger sharks and 36 tagged bull sharks was monitored by means of 42 acoustic receivers distributed along the west coast of the island. Tagged tiger sharks were detected in all months, with seasonal variation between sexes. Detections of males remained low throughout the year but with a peak in winter, whereas detections of females were low in May and August only and peaked in summer. Tagged bull sharks of both sexes were more abundant in winter than in summer, with females present year round. The reasons for the apparent differences in seasonality found in longline and drumline catches compared to acoustic monitoring are discussed.
The internal structure and behaviour of clupeid schools has been observed in situ during unstress... more The internal structure and behaviour of clupeid schools has been observed in situ during unstressed periods followed either by natural predation or by applying a controlled stress (a realistically painted model predator). Combined acoustic and visual (aerial and underwater) observation techniques were used. In situ observations are characterized by a high variability of fish reactions in space and time. The variability in space is obviously due to the large scale of observation which allows the simultaneous occurrence of different types of behaviour in different regions. The variability in time is probably due to a large number of physiological or environmental influences changing unexpectedly between repeated experiments. The variability of fish reaction depends on the characteristics of the source of the stimuli: some sources (e.g. predators) induce different behavioural patterns according to the number of stimuli; others (e.g. boats) induce a single pattern, varying only in intensity. Consequences for acoustic signal identification are discussed.
Journal of Experimental Marine Biology and Ecology, 2012
Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior... more Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior of fish in their environment. In this study, we characterized the spatial and temporal occupation of 37 acoustically tagged bigeye scads (Selar Crumenophthalmus) in an array of shallow Fish Aggregating Devices (FADs) to clarify the mechanism that leads fish to associate with FADs. A comparison of the number of visits and residence times exhibited by the fish at different FADs revealed a strong variability over the array of FADS, with the emergence of a leading FAD, which recorded the majority of visits and retained the fish for a longer period of time. We found diel variability in the residence times, with fish associated at daytime and exploring the array of FADs at nighttime. We demonstrated that this diel temporal pattern was amplified in the leading FAD. We identified a 24-hour periodicity for a subset of individuals aggregated to the leading FAD, thus suggesting that those fish were able to find this FAD after night excursions. The modeling of fish movements based on a Monte Carlo sampling of inter-FAD transitions revealed that the observed spatial heterogeneity in the number of visits could not be explained through simple arrayconnectivity arguments. Similarly, we demonstrated that the high residence times recorded at the leading FAD were not due to the spatial arrangement of individual fish having different associative characters. We discussed the relationships between these patterns of association with the FADs, the exploration of the FAD array and the possible effects of social interactions and environmental factors.
The study of the internal school structure and behaviour of pelagic fish provides interesting inf... more The study of the internal school structure and behaviour of pelagic fish provides interesting information in relation to acoustic surveys, especially when comparing the undisturbed structure with the structure observed beneath a vessel passing over a school. The methodological approach involves in situ observations and combines acoustic and visual (aerial and underwater) techniques. The internal school structure is heterogeneous, including vacuoles, and this structure changes when the school is overpassed by a vessel during the day. In this case compression of the upper layer of the school is observed. The influence of this school structure on the variability of the density estimation has been studied. For the subsurface schools, the usual rate of sampling may be too low for some heterogeneous schools. Other consequences of the school structure on acoustics are discussed.
Canadian Journal of Fisheries and Aquatic Sciences, 2016
Stomach contents and stable carbon and nitrogen isotope analyses (δ13C and δ15N) were used to inv... more Stomach contents and stable carbon and nitrogen isotope analyses (δ13C and δ15N) were used to investigate the trophic ecology of two apex predators, tiger sharks (Galeocerdo cuvier) and bull sharks (Carcharhinus leucas), from Reunion Island to describe their dietary habits at both the population and individual levels. In this oceanic island, the tiger and bull sharks were more piscivorous and teutophagous than noted in previous research from other localities. The δ13C values suggested that bull sharks depended on more neritic organic matter sources than tiger sharks, confirming a coastal habitat preference for bull sharks. Moreover, the total length of the bull shark influenced δ13C values, with smaller individuals being more coastal than larger individuals. All indicators suggest that there is a higher degree of similarity between individual tiger sharks compared with the more heterogeneous bull shark population, which is composed of individuals who specialize on different prey. Th...
Biases in the measurements of spatial distribution of fish schools and their consequences for sch... more Biases in the measurements of spatial distribution of fish schools and their consequences for school biomass estimates during conventional acoustic surveys are mainly due to vertical and lateral avoidance of the vessel. In this paper, we quantify school avoidance during an acoustic survey carried out from 13 to 29 May 1994 in the Catalan Sea. From a lateral multi-beam sonar the geometric characteristics (depth, length, width, height, surface, and volume) of 1268 schools were obtained. The 60 beams (1.5 15) of the sonar scanned a vertical plane from 0 to 90 , perpendicular to the vessel path within a range of 100 m. Within this plane, the projected area ensonified by the echo-sounder used aboard for acoustic evaluation was evaluated to simulate a comparison between the sonar and the echo-sounder. The results have enabled us to improve our knowledge on the vertical and lateral avoidance patterns of schools in relation to their size, external structure, and their position in the water column, and to quantify the vessel influence on biomass estimated by echo-sounder.
The size of pelagic fish schools depends on several parameters related to internal factors such a... more The size of pelagic fish schools depends on several parameters related to internal factors such as species, number of fish, fish swimming speed and physiological status and to external factors, such as hydrological factors and presence of predators. In order to better ...
Schooling fish species are conventionally subdivided into obligate and facultative species, depen... more Schooling fish species are conventionally subdivided into obligate and facultative species, depending on the proportion of the time spent in structured schools, but little is known about the factors affecting the group structure in the absence of external structuring stimuli such as food sources, risk of predators or water flow. Changes in attraction and repulsion indicators, depending on the species and the group size were explored under controlled conditions. Two species, displaying different schooling behaviours in the wild were observed: the bigeye scad Selar crumenophthalmus (Bloch) and the barred flagtail Kuhlia mugil (Forster). In the bigeye scad, the Polarity and Speed were high and stable, and the Nearest Neighbour Distance (NND) decreased when the group size increased. In contrast, for the barred flagtail, Polarity and Speed decreased according to the group size, inducing a loss of cohesion and leading to a disorganized school. The NND mean was stable whatever the group size and relatively high. This experiment indicated that the ability to polarize is first a species specific trait, rather than a property emerging from the group and led by the circumstances.
Demonstrating and quantifying the respective roles of social interactions and external stimuli go... more Demonstrating and quantifying the respective roles of social interactions and external stimuli governing fish dynamics is key to understanding fish spatial distribution. If seminal studies have contributed to our understanding of fish spatial organization in schools, little experimental information is available on fish in their natural environment, where aggregations often occur in the presence of spatial heterogeneities. Here, we applied novel modeling approaches coupled to accurate acoustic tracking for studying the dynamics of a group of gregarious fish in a heterogeneous environment. To this purpose, we acoustically tracked with submeter resolution the positions of twelve small pelagic fish (Selar crumenophthalmus) in the presence of an anchored floating object, constituting a point of attraction for several fish species. We constructed a fieldbased model for aggregated-fish dynamics, deriving effective interactions for both social and external stimuli from experiments. We tuned the model parameters that best fit the experimental data and quantified the importance of social interactions in the aggregation, providing an explanation for the spatial structure of fish aggregations found around floating objects. Our results can be generalized to other gregarious species and contexts as long as it is possible to observe the fine-scale movements of a subset of individuals. Citation: Capello M, Soria M, Cotel P, Deneubourg J-L, Dagorn L (2011) Quantifying the Interplay between Environmental and Social Effects on Aggregated-Fish Dynamics. PLoS ONE 6(12): e28109.
Collective motion phenomena in large groups of social organisms have long fascinated the observer... more Collective motion phenomena in large groups of social organisms have long fascinated the observer, especially in cases, such as bird flocks or fish schools, where large-scale highly coordinated actions emerge in the absence of obvious leaders. However, the mechanisms involved in this self-organized behavior are still poorly understood, because the individual-level interactions underlying them remain elusive. Here, we demonstrate the power of a bottom-up methodology to build models for animal group motion from data gathered at the individual scale. Using video tracks of fish shoal in a tank, we show how a careful, incremental analysis at the local scale allows for the determination of the stimulus/response function governing an individual's moving decisions. We find in particular that both positional and orientational effects are present, act upon the fish turning speed, and depend on the swimming speed, yielding a novel schooling model whose parameters are all estimated from data. Our approach also leads to identify a density-dependent effect that results in a behavioral change for the largest groups considered. This suggests that, in confined environment, the behavioral state of fish and their reaction patterns change with group size. We debate the applicability, beyond the particular case studied here, of this novel framework for deciphering interactions in moving animal groups.
The trajectories of Kuhlia mugil fish swimming freely in a tank are analyzed in order to develop ... more The trajectories of Kuhlia mugil fish swimming freely in a tank are analyzed in order to develop a model of spontaneous fish movement. The data show that K. mugil displacement is best described by turning speed and its auto-correlation. The continuous-time process governing this new kind of displacement is modelled by a stochastic differential equation of Ornstein-Uhlenbeck family: the persistent turning walker. The associated diffusive dynamics are compared to the standard persistent random walker model and we show that the resulting diffusion coefficient scales non-linearly with linear swimming speed. In order to illustrate how interactions with other fish or the environment can be added to this spontaneous movement model we quantify the effect of tank walls on the turning speed and adequately reproduce the characteristics of the observed fish trajectories.
Journal of Experimental Marine Biology and Ecology, 2012
Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior... more Identifying spatial and temporal patterns can reveal the driving factors that govern the behavior of fish in their environment. In this study, we characterized the spatial and temporal occupation of 37 acoustically tagged bigeye scads (Selar Crumenophthalmus) in an array of shallow Fish Aggregating Devices (FADs) to clarify the mechanism that leads fish to associate with FADs. A comparison of the number of visits and residence times exhibited by the fish at different FADs revealed a strong variability over the array of FADS, with the emergence of a leading FAD, which recorded the majority of visits and retained the fish for a longer period of time. We found diel variability in the residence times, with fish associated at daytime and exploring the array of FADs at nighttime. We demonstrated that this diel temporal pattern was amplified in the leading FAD. We identified a 24-hour periodicity for a subset of individuals aggregated to the leading FAD, thus suggesting that those fish were able to find this FAD after night excursions. The modeling of fish movements based on a Monte Carlo sampling of inter-FAD transitions revealed that the observed spatial heterogeneity in the number of visits could not be explained through simple arrayconnectivity arguments. Similarly, we demonstrated that the high residence times recorded at the leading FAD were not due to the spatial arrangement of individual fish having different associative characters. We discussed the relationships between these patterns of association with the FADs, the exploration of the FAD array and the possible effects of social interactions and environmental factors.
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Papers by M. Soria