ORCID Profile
0000-0003-0243-4518
Current Organisation
University of Cambridge
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Biological Mathematics | Simulation and Modelling | Ecology | Behavioural Ecology |
Expanding Knowledge in the Biological Sciences | Communication Networks and Services not elsewhere classified | Expanding Knowledge in Technology
Publisher: The Royal Society
Date: 03-2021
Abstract: The emergent patterns of collective motion are thought to arise from application of in idual-level rules that govern how in iduals adjust their velocity as a function of the relative position and behaviours of their neighbours. Empirical studies have sought to determine such rules of interaction applied by ‘average’ in iduals by aggregating data from multiple in iduals across multiple trajectory sets. In reality, some in iduals within a group may interact differently from others, and such in idual differences can have an effect on overall group movement. However, comparisons of rules of interaction used by in iduals in different contexts have been largely qualitative. Here we introduce a set of randomization methods designed to determine statistical differences in the rules of interaction between in iduals. We apply these methods to a case study of leaders and followers in pairs of freely exploring eastern mosquitofish ( Gambusia holbrooki ). We find that each of the randomization methods is reliable in terms of: repeatability of p -values, consistency in identification of significant differences and similarity between distributions of randomization-based test statistics. We observe convergence of the distributions of randomization-based test statistics across repeat calculations, and resolution of any ambiguities regarding significant differences as the number of randomization iterations increases.
Publisher: Springer Science and Business Media LLC
Date: 05-03-2010
Publisher: The Royal Society
Date: 14-10-2020
Abstract: Grouping is a widespread form of predator defence, with in iduals in groups often performing evasive collective movements in response to attack by predators. In iduals in these groups use behavioural rules to coordinate their movements, with visual cues about neighbours’ positions and orientations often informing movement decisions. Although the exact visual cues in iduals use to coordinate their movements with neighbours have not yet been decoded, some studies have suggested that stripes, lines, or other body patterns may act as conspicuous conveyors of movement information that could promote coordinated group movement, or promote dazzle camouflage, thereby confusing predators. We used phylogenetic logistic regressions to test whether the contrasting achromatic stripes present in four different taxa vulnerable to predation, including species within two orders of birds (Anseriformes and Charadriiformes), a suborder of Artiodactyla (the ruminants), and several orders of marine fishes (predominantly Perciformes) were associated with group living. Contrasting patterns were significantly more prevalent in social species, and tended to be absent in solitary species or species less vulnerable to predation. We suggest that stripes taking the form of light-coloured lines on dark backgrounds, or vice vers a , provide a widespread mechanism across taxa that either serves to inform conspecifics of neighbours' movements, or to confuse predators, when moving in groups. Because detection and processing of patterns and of motion in the visual channel is essentially colour-blind, erse animal taxa with widely different vision systems (including mono-, di-, tri-, and tetrachromats) appear to have converged on a similar use of achromatic patterns, as would be expected given signal-detection theory. This hypothesis would explain the convergent evolution of conspicuous achromatic patterns as an antipredator mechanism in numerous vertebrate species.
Publisher: The Royal Society
Date: 06-01-2014
Abstract: Animals make use a range of social information to inform their movement decisions. One common movement rule, found across many different species, is that the probability that an in idual moves to an area increases with the number of conspecifics there. However, in many cases, it remains unclear what social cues produce this and other similar movement rules. Here, we investigate what cues are used by damselfish ( Dascyllus aruanus ) when repeatedly crossing back and forth between two coral patches in an experimental arena. We find that an in idual's decision to move is best predicted by the recent movements of conspecifics either to or from that in idual's current habitat. Rather than actively seeking attachment to a larger group, in iduals are instead prioritizing highly local and dynamic information with very limited spatial and temporal ranges. By reanalysing data in which the same species crossed for the first time to a new coral patch, we show that the in iduals use static cues in this case. This suggests that these fish alter their information usage according to the structure and familiarity of their environment by using stable information when moving to a novel area and localized dynamic information when moving between familiar areas.
Publisher: Cold Spring Harbor Laboratory
Date: 09-09-2020
DOI: 10.1101/2020.09.07.279711
Abstract: The environment contains different forms of ecological noise that can reduce the ability of animals to detect information. Here we ask whether animals can adapt their behaviour to either exploit or avoid areas of their environment with increased dynamic visual noise. By immersing three-spined sticklebacks ( Gasterosteus aculeatus ) into environments with a simulated form of naturally occurring visual noise – light bands created by the refraction of light from surface waves termed caustic networks – we tested how such visual noise affected the movements, habitat use, and perceptual abilities of these fish. Fish avoided areas of higher visual noise, and achieved this by increasing their activity as a function of the locally perceived noise level, resulting in in iduals moving away from noisier areas. By projecting virtual prey into the environment with different levels of visual noise, we found that the fish’s ability to visually detect prey decreased as visual noise increased. We found no evidence that fish increased their exploration (and decreased their refuge use) in environments with increased visual noise, which would have been predicted if they were exploiting increased visual noise to reduce their own likelihood of being detected. Our results indicate that animals can use simple behavioural strategies to mitigate the impacts of dynamic visual noise on their perceptual abilities, thereby improving their likelihood of gathering information in dynamically changing and noisy environments.
Publisher: The Royal Society
Date: 07-06-2014
Abstract: The istiophorid family of billfishes is characterized by an extended rostrum or ‘bill’. While various functions (e.g. foraging and hydrodynamic benefits) have been proposed for this structure, until now no study has directly investigated the mechanisms by which billfishes use their rostrum to feed on prey. Here, we present the first unequivocal evidence of how the bill is used by Atlantic sailfish ( Istiophorus albicans ) to attack schooling sardines in the open ocean. Using high-speed video-analysis, we show that (i) sailfish manage to insert their bill into sardine schools without eliciting an evasive response and (ii) subsequently use their bill to either tap on in idual prey targets or to slash through the school with powerful lateral motions characterized by one of the highest accelerations ever recorded in an aquatic vertebrate. Our results demonstrate that the combination of stealth and rapid motion make the sailfish bill an extremely effective feeding adaptation for capturing schooling prey.
Publisher: The Royal Society
Date: 07-2017
DOI: 10.1098/RSOS.170043
Abstract: Collective motion describes the global properties of moving groups of animals and the self-organized, coordinated patterns of in idual behaviour that produce them. We examined the group-level patterns and local interactions between in iduals in wild, free-ranging shoals of three-spine sticklebacks, Gasterosteus aculeatus . Our data reveal that the highest frequencies of near-neighbour encounters occur at between one and two body lengths from a focal fish, with the peak frequency alongside a focal in idual. Fish also show the highest alignment with these laterally placed in iduals, and generally with animals in front of themselves. Furthermore, fish are more closely matched in size, speed and orientation to their near neighbours than to more distant neighbours, indicating local organization within groups. Among the group-level properties reported here, we find that polarization is strongly influenced by group speed, but also the variation in speed among in iduals and the nearest neighbour distances of group members. While we find no relationship between group order and group size, we do find that larger groups tend to have lower nearest neighbour distances, which in turn may be important in maintaining group order.
Publisher: American Association for the Advancement of Science (AAAS)
Date: 09-09-2022
Abstract: In shallow water, downwelling light is refracted from surface waves onto the substrate creating bands of light that fluctuate in both time and space, known as caustics. This dynamic illumination can be a visual hindrance for animals in shallow underwater environments. Animals in such habitats may have evolved to use polarization vision for discriminating objects while ignoring the variations in illumination caused by caustics. To explore this possibility, crabs ( Carcinus maenas ) and cuttlefish ( Sepia officinalis ), both of which have polarization vision, were presented with moving stimuli overlaid with caustics. Dynamic caustics inhibited the detection of an intensity-based stimulus but not when these stimuli were polarized. This study is the first to demonstrate that polarization vision reduces the negative impacts that dynamic illumination can have on visual perception.
Publisher: The Royal Society
Date: 03-07-2017
Abstract: The costs and benefits of group living often depend on the spatial position of in iduals within groups and the ability of in iduals to occupy preferred positions. For ex le, models of predation events for moving prey groups predict higher mortality risk for in iduals at the periphery and front of groups. We investigated these predictions in sardine ( Sardinella aurita ) schools under attack from group hunting sailfish ( Istiophorus platypterus ) in the open ocean. Sailfish approached sardine schools about equally often from the front and rear, but prior to attack there was a chasing period in which sardines attempted to swim away from the predator. Consequently, all sailfish attacks were directed at the rear and peripheral positions of the school, resulting in higher predation risk for in iduals at these positions. During attacks, sailfish slash at sardines with their bill causing prey injury including scale removal and tissue damage. Sardines injured in previous attacks were more often found in the rear half of the school than in the front half. Moreover, injured fish had lower tail-beat frequencies and lagged behind uninjured fish. Injuries inflicted by sailfish bills may, therefore, hinder prey swimming speed and drive spatial sorting in prey schools through passive self-assortment. We found only partial support for the theoretical predictions from current predator–prey models, highlighting the importance of incorporating more realistic predator–prey dynamics into these models. This article is part of the themed issue ‘Physiological determinants of social behaviour in animals’.
Publisher: Springer Science and Business Media LLC
Date: 07-07-2022
DOI: 10.1038/S41559-022-01812-0
Abstract: The bio ersity of marine and coastal habitats is experiencing unprecedented change. While there are well-known drivers of these changes, such as overexploitation, climate change and pollution, there are also relatively unknown emerging issues that are poorly understood or recognized that have potentially positive or negative impacts on marine and coastal ecosystems. In this inaugural Marine and Coastal Horizon Scan, we brought together 30 scientists, policymakers and practitioners with transdisciplinary expertise in marine and coastal systems to identify new issues that are likely to have a significant impact on the functioning and conservation of marine and coastal bio ersity over the next 5-10 years. Based on a modified Delphi voting process, the final 15 issues presented were distilled from a list of 75 submitted by participants at the start of the process. These issues are grouped into three categories: ecosystem impacts, for ex le the impact of wildfires and the effect of poleward migration on equatorial bio ersity resource exploitation, including an increase in the trade of fish swim bladders and increased exploitation of marine collagens and new technologies, such as soft robotics and new biodegradable products. Our early identification of these issues and their potential impacts on marine and coastal bio ersity will support scientists, conservationists, resource managers and policymakers to address the challenges facing marine ecosystems.
Publisher: The Royal Society
Date: 04-2015
DOI: 10.1098/RSOS.140355
Abstract: The exceptional reactivity of animal collectives to predatory attacks is thought to be owing to rapid, but local, transfer of information between group members. These groups turn together in unison and produce escape waves. However, it is not clear how escape waves are created from local interactions, nor is it understood how these patterns are shaped by natural selection. By startling schools of fish with a simulated attack in an experimental arena, we demonstrate that changes in the direction and speed by a small percentage of in iduals that detect the danger initiate an escape wave. This escape wave consists of a densely packed band of in iduals that causes other school members to change direction. In the majority of cases, this wave passes through the entire group. We use a simulation model to demonstrate that this mechanism can, through local interactions alone, produce arbitrarily large escape waves. In the model, when we set the group density to that seen in real fish schools, we find that the risk to the members at the edge of the group is roughly equal to the risk of those within the group. Our experiments and modelling results provide a plausible explanation for how escape waves propagate in nature without centralized control.
Publisher: The Royal Society
Date: 12-2015
Abstract: A widespread problem in biological research is assessing whether a model adequately describes some real-world data. But even if a model captures the large-scale statistical properties of the data, should we be satisfied with it? We developed a method, inspired by Alan Turing, to assess the effectiveness of model fitting. We first built a self-propelled particle model whose properties (order and cohesion) statistically matched those of real fish schools. We then asked members of the public to play an online game (a modified Turing test) in which they attempted to distinguish between the movements of real fish schools or those generated by the model. Even though the statistical properties of the real data and the model were consistent with each other, the public could still distinguish between the two, highlighting the need for model refinement. Our results demonstrate that we can use ‘citizen science’ to cross-validate and improve model fitting not only in the field of collective behaviour, but also across a broad range of biological systems.
Publisher: Oxford University Press (OUP)
Date: 21-04-2015
DOI: 10.1093/ICB/ICV017
Abstract: Billfishes are considered among the fastest swimmers in the oceans. Despite early estimates of extremely high speeds, more recent work showed that these predators (e.g., blue marlin) spend most of their time swimming slowly, rarely exceeding 2 m s(-1). Predator-prey interactions provide a context within which one may expect maximal speeds both by predators and prey. Beyond speed, however, an important component determining the outcome of predator-prey encounters is unsteady swimming (i.e., turning and accelerating). Although large predators are faster than their small prey, the latter show higher performance in unsteady swimming. To contrast the evading behaviors of their highly maneuverable prey, sailfish and other large aquatic predators possess morphological adaptations, such as elongated bills, which can be moved more rapidly than the whole body itself, facilitating capture of the prey. Therefore, it is an open question whether such supposedly very fast swimmers do use high-speed bursts when feeding on evasive prey, in addition to using their bill for slashing prey. Here, we measured the swimming behavior of sailfish by using high-frequency accelerometry and high-speed video observations during predator-prey interactions. These measurements allowed analyses of tail beat frequencies to estimate swimming speeds. Our results suggest that sailfish burst at speeds of about 7 m s(-1) and do not exceed swimming speeds of 10 m s(-1) during predator-prey interactions. These speeds are much lower than previous estimates. In addition, the oscillations of the bill during swimming with, and without, extension of the dorsal fin (i.e., the sail) were measured. We suggest that extension of the dorsal fin may allow sailfish to improve the control of the bill and minimize its yaw, hence preventing disturbance of the prey. Therefore, sailfish, like other large predators, may rely mainly on accuracy of movement and the use of the extensions of their bodies, rather than resorting to top speeds when hunting evasive prey.
Publisher: The Royal Society
Date: 04-2017
DOI: 10.1098/RSOS.161056
Abstract: While a rich variety of self-propelled particle models propose to explain the collective motion of fish and other animals, rigorous statistical comparison between models and data remains a challenge. Plausible models should be flexible enough to capture changes in the collective behaviour of animal groups at their different developmental stages and group sizes. Here, we analyse the statistical properties of schooling fish ( Pseudomugil signifer ) through a combination of experiments and simulations. We make novel use of a Boltzmann inversion method, usually applied in molecular dynamics, to identify the effective potential of the mean force of fish interactions. Specifically, we show that larger fish have a larger repulsion zone, but stronger attraction, resulting in greater alignment in their collective motion. We model the collective dynamics of schools using a self-propelled particle model, modified to include varying particle speed and a local repulsion rule. We demonstrate that the statistical properties of the fish schools are reproduced by our model, thereby capturing a number of features of the behaviour and development of schooling fish.
Publisher: Wiley
Date: 23-01-2018
DOI: 10.1111/JZO.12534
Publisher: University of Chicago Press
Date: 06-2013
DOI: 10.1086/670242
Abstract: Explaining how in idual behavior and social interactions give rise to group-level outcomes and affect issues such as leadership is fundamental to the understanding of collective behavior. Here we examined in idual and collective behavioral dynamics in groups of humbug damselfish both before and during a collective movement. During the predeparture phase, group activity increased until the collective movement occurred. Although such movements were precipitated by one in idual, the success or failure of any attempt to instigate a collective movement was not solely dependent on this initiator's behavior but on the behavior of the group as a whole. Specifically, groups were more active and less cohesive before a successful initiation attempt than before a failed attempt. In iduals who made the most attempts to initiate a collective movement during each trial were ultimately most likely to lead the collective movement. Leadership was not related to dominance but was consistent between trials. The probability of fish recruiting to a group movement initiative was an approximately linear function of the number of fish already recruited. Overall, these results are consistent with nonselective local mimetism, with the decision to leave based on a group's, rather than any particular in idual's, readiness to leave.
Publisher: The Royal Society
Date: 16-11-2016
Abstract: We present evidence of a novel form of group hunting. In idual sailfish ( Istiophorus platypterus ) alternate attacks with other group members on their schooling prey ( Sardinella aurita ). While only 24% of attacks result in prey capture, multiple prey are injured in 95% of attacks, resulting in an increase of injured fish in the school with the number of attacks. How quickly prey are captured is positively correlated with the level of injury of the school, suggesting that hunters can benefit from other conspecifics' attacks on the prey. To explore this, we built a mathematical model capturing the dynamics of the hunt. We show that group hunting provides major efficiency gains (prey caught per unit time) for in iduals in groups of up to 70 members. We also demonstrate that a free riding strategy, where some in iduals wait until the prey are sufficiently injured before attacking, is only beneficial if the cost of attacking is high, and only then when waiting times are short. Our findings provide evidence that cooperative benefits can be realized through the facilitative effects of in iduals' hunting actions without spatial coordination of attacks. Such ‘proto-cooperation’ may be the pre-cursor to more complex group-hunting strategies.
Publisher: The Royal Society
Date: 22-05-2019
Abstract: In the wild, prey species often live in the vicinity of predators, rendering the ability to assess risk on a moment-to-moment basis crucial to survival. Visual cues are important as they allow prey to assess predator species, size, proximity and behaviour. However, few studies have explicitly examined prey's ability to assess risk based on predator behaviour and orientation. Using mosquitofish, Gambusia holbrooki , and their predator, jade perch, Scortum barcoo , under controlled conditions, we provide some of the first fine-scale characterization of how prey adapt their behaviour according to their continuous assessment of risk based on both predator behaviour and angular distance to the predator's mouth. When these predators were inactive and posed less of an immediate threat, prey within the attack cone of the predator showed reductions in speed and acceleration characteristic of predator-inspection behaviour. However, when predators became active, prey swam faster with greater acceleration and were closer together within the attack cone of predators. Most importantly, this study provides evidence that prey do not adopt a uniform response to the presence of a predator. Instead, we demonstrate that prey are capable of rapidly and dynamically updating their assessment of risk and showing fine-scale adjustments to their behaviour.
Publisher: Springer Science and Business Media LLC
Date: 08-12-2013
Publisher: Cold Spring Harbor Laboratory
Date: 20-04-2020
DOI: 10.1101/2020.04.20.050245
Abstract: Grouping is a widespread form of predator defense, with in iduals in groups often performing evasive collective movements in response to predators’ attacks. In iduals in these groups use behavioral rules to coordinate their movements, with visual cues about neighbors’ positions and orientations informing movement decisions. Although the exact visual cues in iduals use to coordinate their movements with neighbors have not yet been decoded, some studies have suggested that stripes, lines or other body patterns may act as conspicuous conveyors of movement information that could promote coordinated group movement, or promote dazzle camouflage, thereby confusing predators. We used phylogenetic logistic regressions to test whether the contrasting achromatic stripes present in four different taxa vulnerable to predation, including species within two orders of birds (Anseriformes and Charadriiformes), a suborder of Artiodactyla (the ruminants) and several orders of marine fish (predominantly Perciformes) were associated with group living. Contrasting patterns were significantly more prevalent in social species, and tended to be absent in solitary species or species less vulnerable to predation. We suggest that stripes taking the form of light-colored lines on dark backgrounds, or vice vers a, provide a widespread mechanism across taxa that serves either to inform conspecifics of neighbors’ directional movement, or to confuse predators, when moving in groups. Detection and processing of patterns and of motion in the visual channel is essentially colourblind. That erse animal taxa with widely different vision systems (including di-, tri- and tetrachromats) appear to have converged on a similar use of achromatic patterns is therefore expected given signal-detection theory. This hypothesis would explain the convergent evolution of conspicuous achromatic patterns as an antipredator mechanism in numerous vertebrate species.
Publisher: The Royal Society
Date: 28-12-2016
Publisher: Wiley
Date: 02-06-2014
DOI: 10.1111/ETH.12262
Publisher: American Association for the Advancement of Science (AAAS)
Date: 04-12-2020
Abstract: The rules of interaction that animals use to coordinate their movements are heritable and can evolve rapidly.
Publisher: Elsevier BV
Date: 02-2017
DOI: 10.1016/J.CUB.2016.12.044
Abstract: Lateralization is widespread throughout the animal kingdom [1-7] and can increase task efficiency via shortening reaction times and saving on neural tissue [8-16]. However, lateralization might be costly because it increases predictability [17-21]. In predator-prey interactions, for ex le, predators might increase capture success because of specialization in a lateralized attack, but at the cost of increased predictability to their prey, constraining the evolution of lateralization. One unexplored mechanism for evading such costs is group hunting: this would allow in idual-level specialization, while still allowing for group-level unpredictability. We investigated this mechanism in group hunting sailfish, Istiophorus platypterus, attacking schooling sardines, Sardinella aurita. During these attacks, sailfish alternate in attacking the prey using their elongated bills to slash or tap the prey [22-24]. This rapid bill movement is either leftward or rightward. Using behavioral observations of identifiable in idual sailfish hunting in groups, we provide evidence for in idual-level attack lateralization in sailfish. More strongly lateralized in iduals had a higher capture success. Further evidence of lateralization comes from morphological analyses of sailfish bills that show strong evidence of one-sided micro-teeth abrasions. Finally, we show that attacks by single sailfish are indeed highly predictable, but predictability rapidly declines with increasing group size because of a lack of population-level lateralization. Our results present a novel benefit of group hunting: by alternating attacks, in idual-level attack lateralization can evolve, without the negative consequences of in idual-level predictability. More generally, our results suggest that group hunting in predators might provide more suitable conditions for the evolution of strategy ersity compared to solitary life.
Publisher: Elsevier BV
Date: 08-2018
Publisher: Proceedings of the National Academy of Sciences
Date: 07-11-2011
Abstract: Collective motion, where large numbers of in iduals move synchronously together, is achieved when in iduals adopt interaction rules that determine how they respond to their neighbors’ movements and positions. These rules determine how group-living animals move, make decisions, and transmit information between in iduals. Nonetheless, few studies have explicitly determined these interaction rules in moving groups, and very little is known about the interaction rules of fish. Here, we identify three key rules for the social interactions of mosquitofish ( Gambusia holbrooki ): ( i ) Attraction forces are important in maintaining group cohesion, while we find only weak evidence that fish align with their neighbor’s orientation ( ii ) repulsion is mediated principally by changes in speed ( iii ) although the positions and directions of all shoal members are highly correlated, in iduals only respond to their single nearest neighbor. The last two of these rules are different from the classical models of collective animal motion, raising new questions about how fish and other animals self-organize on the move.
Publisher: The Royal Society
Date: 07-02-2013
Abstract: How different levels of biological organization interact to shape each other's function is a central question in biology. One particularly important topic in this context is how in iduals' variation in behaviour shapes group-level characteristics. We investigated how fish that express different locomotory behaviour in an asocial context move collectively when in groups. First, we established that in idual fish have characteristic, repeatable locomotion behaviours (i.e. median speeds, variance in speeds and median turning speeds) when tested on their own. When tested in groups of two, four or eight fish, we found in iduals partly maintained their asocial median speed and median turning speed preferences, while their variance in speed preference was lost. The strength of this in iduality decreased as group size increased, with in iduals conforming to the speed of the group, while also decreasing the variability in their own speed. Further, in iduals adopted movement characteristics that were dependent on what group size they were in. This study therefore shows the influence of social context on in idual behaviour. If the results found here can be generalized across species and contexts, then although in iduality is not entirely lost in groups, social conformity and group-size-dependent effects drive how in iduals will adjust their behaviour in groups.
Publisher: Oxford University Press (OUP)
Date: 18-11-2019
Abstract: Detailed quantifications of how predators and their grouping prey interact in three dimensions (3D) remain rare. Here we record the structure and dynamics of fish shoals (Pseudomugil signifer) in 3D both with and without live predators (Philypnodon grandiceps) under controlled laboratory conditions. Shoals adopted two distinct types of shoal structure: “sphere-like” geometries at depth and flat “carpet-like” structures at the water’s surface, with shoals becoming more compact in both horizontal and vertical planes in the presence of a predator. The predators actively stalked and attacked the prey, with attacks being initiated when the shoals were not in their usual configurations. These attacks caused the shoals to break apart, but shoal reformation was rapid and involved in iduals adjusting their positions in both horizontal and vertical dimensions. Our analyses revealed that targeted prey were more isolated from other conspecifics, and were closer in terms of distance and direction to the predator compared to non-targeted prey. Moreover, which prey were targeted could largely be identified based on in iduals’ positions from a single plane. This highlights that previously proposed 2D theoretical models and their assumptions appear valid when considering how predators target groups in 3D. Our work provides experimental, and not just anecdotal, support for classic theoretical predictions and also lends new insights into predatory–prey interactions in three-dimensional environments.
Location: United Kingdom of Great Britain and Northern Ireland
Start Date: 2019
End Date: 2019
Amount: $400,000.00
Funder: Australian Research Council
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