Escape path complexity and its context dependency in Pacific blue-eyes (Pseudomugil signifer)

The escape trajectories animals take following a predatory attack appear to show high degrees of apparent 'randomness' - a property that has been described as 'protean behaviour'. Here we present a method of quantifying the escape trajectories of individual animals using a path complexity approach. When fish (Pseudomugil signifer) were attacked either on their own or in groups, we find that an individual's path rapidly increases in entropy (our measure of complexity) following the attack. For individuals on their own, this entropy remains elevated (indicating a more random path) for a sustained period (10 seconds) after the attack, whilst it falls more quickly for individuals in groups. The entropy of the path is context dependent. When attacks towards single fish come from greater distances, a fish's path shows less complexity compared to attacks that come from short range. This context dependency effect did not exist, however, when individuals were in groups. Nor did the path complexity of individuals in groups depend on a fish's local density of neighbours. We separate out the components of speed and direction changes to determine which of these components contributes to the overall increase in path complexity following an attack. We found that both speed and direction measures contribute similarly to an individual's path's complexity in absolute terms. Our work highlights the adaptive behavioural tactics that animals use to avoid predators and also provides a novel method for quantifying the escape trajectories of animals.Comment: 9 page

Social recognition and social attraction in group-living fishes

This work was supported by a grant from the Australian Research Council, Discovery Project DP190100660, focusing on the social behavior of animals.Social aggregation is a widespread and important phenomenon among fishes. Understanding the questions of why and how aggregations form and are subsequently maintained is a central goal for behavioral ecologists. Research in this field has shown that aggregations are typically structured, non-random associations. This indicates that fish are able to differentiate between potential group-mates and that this ability mediates their association preferences, and, ultimately, the composition of their groups. In this review, we examine the characteristics that influence the expression of social attraction among fishes, before going on to describe the recognition mechanisms that underpin social attraction. Finally, we highlight a number of outstanding questions in the field with a view to generating a more complete understanding of social aggregation in fishes.Publisher PDFPeer reviewe

To clean or not to clean: cleaning mutualism breakdown in a tidal environment

The dynamics and prevalence of mutualistic interactions, which are responsible for the maintenance and structuring of all ecological communities, are vulnerable to changes in abiotic and biotic environmental conditions. Mutualistic outcomes can quickly shift from cooperation to conflict, but it unclear how resilient and stable mutualistic outcomes are to more variable conditions. Tidally controlled coral atoll lagoons that experience extreme diurnal environmental shifts thus provide a model from which to test plasticity in mutualistic behavior of dedicated (formerly obligate) cleaner fish, which acquire all their food resources through client interactions. Here, we investigated cleaning patterns of a model cleaner fish species, the bluestreak wrasse (Labroides dimidiatus), in an isolated tidal lagoon on the Great Barrier Reef. Under tidally restricted conditions, uniquely both adults and juveniles were part‐time facultative cleaners, pecking on Isopora palifera coral. The mutualism was not completely abandoned, with adults also wandering across the reef in search of clients, rather than waiting at fixed site cleaning stations, a behavior not yet observed at any other reef. Contrary to well‐established patterns for this cleaner, juveniles appeared to exploit the system, by biting (“cheating”) their clients more frequently than adults. We show for the first time, that within this variable tidal environment, where mutualistic cleaning might not represent a stable food source, the prevalence and dynamics of this mutualism may be breaking down (through increased cheating and partial abandonment). Environmental variability could thus reduce the pervasiveness of mutualisms within our ecosystems, ultimately reducing the stability of the system

Mid-sized groups perform best in a collective decision task in sticklebacks

This study was supported by Australian Research Council (grant no. DP160103905).Numerous studies have reported functional improvements in collective behaviour with increasing group size, however, the possibility that such improvements may saturate or even decline as group size continues to grow have seldom been tested experimentally. Here, we tested the ability of solitary three-spined sticklebacks and those in groups, ranging from 2 to 29 fish, to leave an unfavourable patch of habitat. Our results replicate the findings of previous studies at low group sizes, with the fish initially showing a reduction in their latency to leave the unfavourable habitat as group size increased. As group size continued to increase, however, latency to leave the habitat increased, so that the functional relationship between group size and latency to depart was U-shaped. Our results suggest an optimum group size in this context of between 12 and 20 fish. Underlying this group-level trend was a similar U-shaped relationship between group size and the first fish to leave the habitat, suggesting that at larger group sizes, social conformity to the behaviour of the majority can stifle the ability of fish to innovate-in this case, to induce a collective movement from the unfavourable habitat.PostprintPeer reviewe

Morphological differences between habitats are associated with physiological and behavioural trade-offs in stickleback (Gasterosteus aculeatus)

F.S. and A.J.W.W. were supported by the Australian Research Council, M.M.W. was supported by The University of St Andrews and R.S.J. and J.T. were supported by Coventry UniversityLocal specialization can be advantageous for individuals and may increase the resilience of the species to environmental change. However, there may be trade-offs between morphological responses and physiological performance and behaviour. Our aim was to test whether habitat-specific morphology of stickleback (Gasterosteus aculeatus) interacts with physiological performance and behaviour at different salinities. We rejected the hypothesis that deeper body shape of fish from habitats with high predation pressure led to decreases in locomotor performance. However, there was a trade-off between deeper body shape and muscle quality. Muscle of deeper-bodied fish produced less force than that of shallow-bodied saltmarsh fish. Nonetheless, saltmarsh fish had lower swimming performance, presumably because of lower muscle mass overall coupled with smaller caudal peduncles and larger heads. Saltmarsh fish performed better in saline water (20 ppt) relative to freshwater and relative to fish from freshwater habitats. However, exposure to salinity affected shoaling behaviour of fish from all habitats and shoals moved faster and closer together compared with freshwater. We show that habitat modification can alter phenotypes of native species, but local morphological specialization is associated with trade-offs that may reduce its benefits.Publisher PDFPeer reviewe

Quantifying the structure and dynamics of fish shoals under predation threat in three dimensions

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 at- tacked 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 individuals 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 individuals' 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, sup- port for classic theoretical predictions, and also lends new insights into predatory-prey interactions in three-dimensional environments

Local interactions and global properties of wild, free-ranging stickleback shoals

Funding: Australian Research Council. A.J.W.W. and T.M.S. were supported by a Discovery Project Grant from the Australian Research Council. D.J.T.S. and J.E.H.-R. were supported by a Knut & Alice Wallenberg Foundation Grant.Collective motion describes the global properties of moving groups of animals and the self-organized, coordinated patterns of individual behaviour that produce them. We examined the group-level patterns and local interactions between individuals 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 individual. Fish also show the highest alignment with these laterally placed individuals, 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 individuals 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 PDFPeer reviewe

Body size affects the strength of social interactions and spatial organization of a schooling fish (Pseudomugil signifer)

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

Consistency of Leadership in Shoals of Mosquitofish (Gambusia holbrooki) in Novel and in Familiar Environments

In social animal groups, an individual's spatial position is a major determinant of both predation risk and foraging rewards. Additionally, the occupation of positions in the front of moving groups is generally assumed to correlate with the initiation of group movements. However, whether some individuals are predisposed to consistently occupy certain positions and, in some instances, to consistently lead groups over time is as yet unresolved in many species. Using the mosquitofish (Gambusia holbrooki), we examined the consistency of individuals' spatial positions within a moving group over successive trials. We found that certain individuals consistently occupied front positions in moving groups and also that it was typically these individuals that initiated group decisions. The number of individuals involved in leading the group varied according to the amount of information held by group members, with a greater number of changes in leadership in a novel compared to a relatively familiar environment. Finally, our results show that the occupation of lead positions in moving groups was not explained by characteristics such as dominance, size or sex, suggesting that certain individuals are predisposed to leadership roles. This suggests that being a leader or a follower may to some extent be an intrinsic property of the individual

Opportunity Mars Rover mission: Overview and selected results from Purgatory ripple to traverses to Endeavour crater

Opportunity has been traversing the Meridiani plains since 25 January 2004 (sol 1), acquiring numerous observations of the atmosphere, soils, and rocks. This paper provides an overview of key discoveries between sols 511 and 2300, complementing earlier papers covering results from the initial phases of the mission. Key new results include (1) atmospheric argon measurements that demonstrate the importance of atmospheric transport to and from the winter carbon dioxide polar ice caps; (2) observations showing that aeolian ripples covering the plains were generated by easterly winds during an epoch with enhanced Hadley cell circulation; (3) the discovery and characterization of cobbles and boulders that include iron and stony‐iron meteorites and Martian impact ejecta; (4) measurements of wall rock strata within Erebus and Victoria craters that provide compelling evidence of formation by aeolian sand deposition, with local reworking within ephemeral lakes; (5) determination that the stratigraphy exposed in the walls of Victoria and Endurance craters show an enrichment of chlorine and depletion of magnesium and sulfur with increasing depth. This result implies that regional‐scale aqueous alteration took place before formation of these craters. Most recently, Opportunity has been traversing toward the ancient Endeavour crater. Orbital data show that clay minerals are exposed on its rim. Hydrated sulfate minerals are exposed in plains rocks adjacent to the rim, unlike the surfaces of plains outcrops observed thus far by Opportunity. With continued mechanical health, Opportunity will reach terrains on and around Endeavour’s rim that will be markedly different from anything examined to date.Additional co-authors: RM Haberle, KE Herkenhoff, JA Herman, KD Iagnemma, BL Jolliff, JR Johnson, G Klingelhöfer, AH Knoll, AT Knudson, R Li, SM McLennan, DW Mittlefehldt, RV Morris, TJ Parker, MS Rice, LA Soderblom, SW Squyres, RJ Sullivan, MJ Wolf