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A sympatric assemblage of morphologically similar predators is expected to exhibit fine-scale habitat segregation, or resource partitioning, to reduce the effects of direct competition. This principle has been well studied for predators in terrestrial ecosystems. In the marine environment, the fine-scale spatial segregation of sympatric species of large predators is poorly understood because detailed movement and behavioural data are often not available across multiple species within the same timeframe. The ways in which co-occurring congeneric predators separate spatially is even less well understood. Medium-sized species of skates (genus Raja) co-occur in temperate habitats of the north-east Atlantic Ocean, share similar morphologies and have distributional ranges that overlap significantly in the western English Channel ecosystem. In the present study, detailed depth time series retrieved from 89 electronic data storage tags attached to 4 species of skate were analysed to determine preferred depth ranges. The 4 species were found to segregate spatially into 2 groups, with one group having a significantly shallower core annual depth range than the other. To our knowledge, fine-scale segregation by depth has not been observed previously. Interestingly, the members of each species group appeared complementary, each group comprising species with different dietary preferences and with a larger and smaller body size. An understanding of how core depth ranges differ and how these species utilise vertical habitat could potentially enable geographic ranges around the coast to be predicted, with important implications for how these species interact with fisheries and Marine Protected Areas.

article i nfo The increasing importance of the European common cuttlefish Sepia officinalis as a commercial fishery re- source raises concerns that our limited understanding of its behaviour and movements in natural habitats may impede future management for this species. Whilst information regarding the fine-scale movements and behaviour of small marine species such as cuttlefish can be difficult to obtain, the increasing miniaturisation of electronic tags, combined with novel attachment techniques, has now enabled tracking of this cephalopod. In this study we describe the first use of electronic tags attached to adult and sub-adult S. officinalis to monitor their movements and behaviour within natural environments. Eight adults (170- 205 mm dorsal mantle length (DML)) were fitted with continuous acoustic transmitters and ten sub-adults (132-180 mm DML) were fitted with coded acoustic transmitters. All transmitters were attached to the internal cuttlebone of the living animal. Tagged animals recovered rapidly from anaesthesia and surgical pro- cedures and were found to retain tags for periods of up to 11 weeks in this short-term study. Six of the eight adult cuttlefish released into a radio-acoustic positioning (VRAP) array left the area rapidly (≤45 min) whilst two remained for multiple days (≤9 days). Short-term site fidelity at a local level was observed in only two adults, whilst larger scale movements along the coastline (≤35 km) occurred in three adults, indicating that a complex range of spatial behaviour patterns could occur among spawning adults. Similar patterns were ob- served in the space use of sub-adults monitored using a passive acoustic-receiver array on the seabed. Three sub-adults were detected within the study area over an extended period of time (≥73 days), whilst the remaining sub-adults left the study area rapidly and without return. The capacity to monitor the movements of both adult and sub-adult cuttlefish in their natural environment will facilitate identification and under- standing of ontogenic changes in behaviours and migratory patterns of wild populations. Our results high- light the potential of electronic tagging methods as tools to aid conservation and management of this important commercial fishery species in the future.

The physiology and behaviour of captive cuttlefish have been well studied over recent decades and yet very few parallel studies have investigated the movements and behaviour of free-ranging animals. One reason for this gap is that methods for longer term tracking are not well developed for the majority of species, especially those of smaller size. In this study we describe a methodology for the long-term attachment of electronic tags to free-ranging cuttlefish of a broad size range. Five common cuttlefish Sepia officinalis (122–240 mm mantle length) were fitted with temperature and depth-logging archival tags attached to the internal cuttlebone of the living animal (tag weight was < 1.8% of body weight). Tagged animals recovered rapidly from sedative and surgical procedures and resumed feeding within 24 h of tag attachment. Laboratory-kept cuttlefish maintained in a 2-m deep aquarium exhibited diel vertical movements, spending daylight hours close to the bottom and ascending into the water column at night, a behaviour which is consistent with normal diel vertical migration seen in other marine organisms such as zooplankton and fish. That regular feeding and expected patterns of depth use were observed indicates that tag attachment did not impede normal behaviours. Furthermore, tags remained attached to laboratory cuttlefish for up to 5 months, from first tagging through to natural mortality, and with the tag remaining attached to the cuttlebone post mortem. We also found that cuttlebones with attached tags remained buoyant at the water's surface post mortem, offering the unique opportunity to recover archival tags from cuttlebones washed ashore. The capacity for long-term attachment to juvenile cuttlefish and recovery following natural mortality identifies the potential of this method for recording near ‘lifetime’ behaviour and movements of cuttlefish in their natural environment.

The use of archival depth telemetry as a means of remotely assessing the reproductive rates of free-ranging fishes is explored. This is achieved by electronically tracking the vertical movements of individual female small spotted catsharks Scyliorhinus canicula in the natural environment, whilst simultaneously evaluating the temporal and vertical distributions of egg-laying in this species. Distinctive patterns of short-term (0·3-3·7 h), shallow-water activity are documented in the time-depth profiles of female S. canicula that occur at an appropriate depth (1·0-2·3 m) and periodicity (every 10-12 days) to represent egg-laying behaviour. Putative egg-laying behaviour was exhibited simultaneously by two individually tracked female S. canicula during late-spring and early-summer. The results highlight that, provided species behaviour is suitable and complementary methods such as previous data, laboratory experiments and field surveys can be used to validate the patterns observed, archival depth telemetry offers an unobtrusive means by which egg production and egg-laying behaviour of free-living fishes can be estimated. As precise information regarding life-history parameters is difficult to obtain for free-ranging fish species, this technique could be used to improve the parameterization of species demographic models that are relevant to the management of wild fish populations.

Within-species sexual segregation is a widespread phenomenon among vertebrates, but its causes remain a topic of much debate. Female avoidance of male coercive mating attempts has the potential to influence the social structure of animal populations, yet it has been largely overlooked as a driver of sexual separation. Indeed, its potential role in long-term structuring of natural populations has not been studied. Here we use a comparative approach to examine the suitability of multiple hypotheses forwarded to account for sexual segregation (i.e., activity budget, predation risk, thermal niche–fecundity, and social factors) as drivers underlying sex-specific habitat use in a monomorphic model vertebrate, the small-spotted catshark, Scyliorhinus canicula. Using this hypothesis-driven approach, we show that year-round sexual habitat segregation in S. canicula can be accounted for directly by female avoidance of male sexual harassment. Long-term electronic tracking reveals that sperm-storing female catsharks ...

Microsatellite markers were characterised in the small-spotted catshark (Scyliorhinus canicula) from unpublished sequences generated from expressed sequence tags and bacterial artificial chromosomes. All seventeen of the loci screened were found to be polymorphic within the population of catsharks tested. The number of alleles identified varied between two and twelve per locus, and the expected heterozygosity ranged from 0.40 to 0.83. Evidence of null alleles was detected in two loci, while one locus showed significant deviation from expectations of Hardy–Weinberg equilibrium. There was no evidence of significant linkage between any loci. Cross-amplification was attempted in six other species of elasmobranch and success generally followed levels of taxonomic relatedness. These markers should be a powerful tool for population genetic and molecular studies of the small-spotted catshark, and may prove useful in other shark species.

Commercial fishing and climate change have influenced the composition of marine fish assemblages worldwide, but we require a better understanding of their relative influence on long-term changes in species abundance and body-size distributions. In this study, we investigated long-term (1911‐2007) variability within a demersal fish assemblage in the western English Channel. The region has been subject to commercial fisheries throughout most of the past century, and has undergone interannual changes in sea temperature of over 2.01C. We focussed on a core 30 species that comprised 99% of total individuals sampled in the assemblage. Analyses showed that temporal trends in the abundance of smaller multispecies size classes followed thermal regime changes, but that there were persistent declines in abundance of larger size classes. Consistent with these results, larger-growing individual species had the greatest declines in body size, and the most constant declines in abundance, while abundance changes of smaller-growing species were more closely linked to preceding sea temperatures. Together these analyses are suggestive of dichotomous size-dependent responses of species to long-term climate change and commercial fishing over a century scale. Small species had rapid responses to the prevailing thermal environment, suggesting their life history traits predisposed populations to respond quickly to changing climates. Larger species declined in abundance and size, reflecting expectations from sustained size-selective overharvesting. These results demonstrate the importance of considering species traits when developing indicators of human and climatic impacts on marine fauna.

Understanding animal distributions and habitat utilisation is vital for the management of populations, especially those of endangered species. However, this information is not available for the majority of marine species and is difficult to obtain for those with low population densities. The common skate, Dipturus batis, was once abundant and widespread in the North-East Atlantic but is now thought to be locally extinct in the Irish Sea and in the central and southern North Sea, and is listed as Critically Endangered on the IUCN Red List. The constraints of skate body morphology on locomotory mode assume low levels of activity with long periods spent resting on the seabed, therefore predicting a high degree of site fidelity. To investigate this hypothesis we tagged 8 common skate (two male and six female, mass range: 10.9–63.5 kg) with depth and temperature-logging data storage tags off the west coast of Scotland in May 2008. All 6 tags attached to females were recovered after 1–9 months at liberty. All 6 individuals exhibited pronounced site fidelity to highly localised areas. Within these local areas however, time-depth profiles were dominated by periods of high activity, with vertical movements of > 100 m being conducted on a regular, sometimes daily, basis. Intra-individual plasticity was observed in vertical activity patterns with individuals switching between low and high activity patterns. Smaller skate were generally less active and occupied deeper depths. Limited short-term horizontal movements in preferred habitats supporting apparently high foraging activity highlights the need for spatial management of ‘refugial’ populations of this once widespread fish, that appears now largely extirpated from European waters.

the tendency for marine fish to refuge in natural and artificial structures underwater is well documented, but the incidence of this behaviour among predatory sharks is not widely known. a sub-adult male nursehound scyliorhinus stellaris was tracked by intermittent acoustic telemetry over 168 days in a tidal sea lough. this individual, in-between undertaking nocturnal foraging excursions, refuged in at least five different narrow-entrance holes. refuge location was validated by underwater and surface observations during which time other nursehounds (sub-adult male and female), were also seen refuging, sometimes together. this indicates nursehound display philopatric behaviour centred on aggregation in ‘home’ refuges that, in this case, were labyrinthine rock systems.

Efficient searching is crucial for timely location of food and other resources. Recent studies show that diverse living animals use a theoretically optimal scale-free random search for sparse resources known as a Lévy walk, but little is known of the origins and evolution of foraging behavior and the search strategies of extinct organisms. Here, using simulations of self-avoiding trace fossil trails, we show that randomly introduced strophotaxis (U-turns)--initiated by obstructions such as self-trail avoidance or innate cueing--leads to random looping patterns with clustering across increasing scales that is consistent with the presence of Lévy walks. This predicts that optimal Lévy searches may emerge from simple behaviors observed in fossil trails. We then analyzed fossilized trails of benthic marine organisms by using a novel path analysis technique and find the first evidence, to our knowledge, of Lévy-like search strategies in extinct animals. Our results show that simple search behaviors of extinct animals in heterogeneous environments give rise to hierarchically nested Brownian walk clusters that converge to optimal Lévy patterns. Primary productivity collapse and large-scale food scarcity characterizing mass extinctions evident in the fossil record may have triggered adaptation of optimal Lévy-like searches. The findings suggest that Lévy-like behavior has been used by foragers since at least the Eocene but may have a more ancient origin, which might explain recent widespread observations of such patterns among modern taxa.

International audience; The decisions animals make about how long to wait between activities can determine the success of diverse behaviours such as foraging, group formation or risk avoidance. Remarkably, for diverse animal species, including humans, spontaneous patterns of waiting times show random 'burstiness' that appears scale-invariant across a broad set of scales. However, a general theory linking this phenomenon across the animal kingdom currently lacks an ecological basis. Here, we demonstrate from tracking the activities of 15 sympatric predator species (cephalopods, sharks, skates and teleosts) under natural and controlled conditions that bursty waiting times are an intrinsic spontaneous behaviour well approximated by heavy-tailed (power-law) models over data ranges up to four orders of magnitude. Scaling exponents quantifying ratios of frequent short to rare very long waits are species-specific, being determined by traits such as foraging mode (active versus ambush predation), body size and prey preference. A stochastic-deterministic decision model reproduced the empirical waiting time scaling and species-specific exponents, indicating that apparently complex scaling can emerge from simple decisions. Results indicate temporal power-law scaling is a behavioural 'rule of thumb' that is tuned to species' ecological traits, implying a common pattern may have naturally evolved that optimizes move-wait decisions in less predictable natural environments.

Many sharks and skates are particularly vulnerable to overfishing because of their large size, slow growth, late maturity and low fecundity. In Europe dramatic population declines have taken place in common skate (Dipturus batisL.), one of the largest demersal fish in regional shelf seas, leading to extirpations from substantial parts of its former range. Here we report the discovery of cryptic species in common skate collected from the northeast Atlantic continental shelf. Data from nuclear microsatellite markers indicated two clearly distinct clades and phylogenetic analysis of mitochondrial DNA sequences demonstrated monophyly of each one of them. Capture locations showed evidence of strong spatial segregation, with one taxon occurring mainly in waters off the southern British Isles and around Rockall, while the other was restricted to more northerly shelf waters. These apparently cryptic species showed overlapping substrate and depth preferences, but distributional limits were closely related to temperature gradients, potentially indicating thermal limits to their distributions. This discovery of hidden diversity within a large, critically endangered marine vertebrate demonstrates how marine biodiversity can be underestimated, even in such a relatively well-studied and heavily exploited region.

An optimal search theory, the so-called Levy-flight foraging hypothesis1, predicts that predators should adopt search strategies known as Levy flights where prey is sparse and distributed unpredictably, but that Brownian movement is sufficiently efficient for locating abundant prey2, 3, 4. Empirical studies have generated controversy because the accuracy of statistical methods that have been used to identify Levy behaviour has recently been questioned5, 6. Consequently, whether foragers exhibit Levy flights in the wild remains unclear. Crucially, moreover, it has not been tested whether observed movement patterns across natural landscapes having different expected resource distributions conform to the theory’s central predictions. Here we use maximum-likelihood methods to test for Levy patterns in relation to environmental gradients in the largest animal movement data set assembled for this purpose. Strong support was found for Levy search patterns across 14 species of open-ocean predatory fish (sharks, tuna, billfish and ocean sunfish), with some individuals switching between Levy and Brownian movement as they traversed different habitat types. We tested the spatial occurrence of these two principal patterns and found Levy behaviour to be associated with less productive waters (sparser prey) and Brownian movements to be associated with productive shelf or convergence-front habitats (abundant prey). These results are consistent with the Levy-flight foraging hypothesis1, 7, supporting the contention8, 9 that organism search strategies naturally evolved in such a way that they exploit optimal Levy patterns.

Sexual segregation occurs when members of a species separate such that the sexes live apart, either singly or in single-sex groups. It can be broadly categorised into two types: habitat segregation and social segregation. Sexual segregation is a behavioural phenomenon that is widespread in the animal kingdom yet the underlying causes remain poorly understood. Sexual segregation has been widely studied among terrestrial mammals such as ungulates, but it has been less well documented in the marine environment. This chapter clarifies terms and concepts which have emerged from the investigation of sexual segregation in terrestrial ecology and examines how a similar methodological approach may be complicated by differences of marine species. Here we discuss the behavioural patterns of sexual segregation among marine fish, reptile, bird and mammal species. Five hypotheses have been forwarded to account for sexual segregation, largely emerging from investigation of sexual segregation in terrestrial ungulates: the predation risk, forage selection, activity budget, thermal niche-fecundity and social factors hypotheses. These mechanisms are reviewed following careful assessment of their applicability to marine vertebrate species and case studies of marine vertebrates which support each mechanism recounted. Rigorous testing of all hypotheses is lacking from both the terrestrial and marine vertebrate literature and those analyses which have been attempted are often confounded by factors such as sexual body-size dimorphism. In this context, we indicate the value of studying model species which are monomorphic with respect to body size and discuss possible underlying causes for sexual segregation in this species. We also discuss why it is important to understand sexual segregation, for example, by illustrating how differential exploitation of the sexes by humans can lead to population decline.

Sexual segregation occurs when members of a species separate such that the sexes live apart, either singly or in single-sex groups. It can be broadly categorised into two types: habitat segregation and social segregation. Sexual segregation is a behavioural phenomenon that is widespread in the animal kingdom yet the underlying causes remain poorly understood. Sexual segregation has been widely studied among terrestrial mammals such as ungulates, but it has been less well documented in the marine environment. This chapter clarifies terms and concepts which have emerged from the investigation of sexual segregation in terrestrial ecology and examines how a similar methodological approach may be complicated by differences of marine species. Here we discuss the behavioural patterns of sexual segregation among marine fish, reptile, bird and mammal species. Five hypotheses have been forwarded to account for sexual segregation, largely emerging from investigation of sexual segregation in terrestrial ungulates: the predation risk, forage selection, activity budget, thermal niche-fecundity and social factors hypotheses. These mechanisms are reviewed following careful assessment of their applicability to marine vertebrate species and case studies of marine vertebrates which support each mechanism recounted. Rigorous testing of all hypotheses is lacking from both the terrestrial and marine vertebrate literature and those analyses which have been attempted are often confounded by factors such as sexual body-size dimorphism. In this context, we indicate the value of studying model species which are monomorphic with respect to body size and discuss possible underlying causes for sexual segregation in this species. We also discuss why it is important to understand sexual segregation, for example, by illustrating how differential exploitation of the sexes by humans can lead to population decline.

Many free-ranging predators have to make foraging decisions with little, if any, knowledge of present resource distribution and availability1. The optimal search strategy they should use to maximize encounter rates with prey in heterogeneous natural environments remains a largely unresolved issue in ecology1, 2, 3. Lévy walks4 are specialized random walks giving rise to fractal movement trajectories that may represent an optimal solution for searching complex landscapes5. However, the adaptive significance of this putative strategy in response to natural prey distributions remains untested6, 7. Here we analyse over a million movement displacements recorded from animal-attached electronic tags to show that diverse marine predators—sharks, bony fishes, sea turtles and penguins—exhibit Lévy-walk-like behaviour close to a theoretical optimum2. Prey density distributions also display Lévy-like fractal patterns, suggesting response movements by predators to prey distributions. Simulations show that predators have higher encounter rates when adopting Lévy-type foraging in natural-like prey fields compared with purely random landscapes. This is consistent with the hypothesis that observed search patterns are adapted to observed statistical patterns of the landscape. This may explain why Lévy-like behaviour seems to be widespread among diverse organisms3, from microbes8 to humans9, as a 'rule' that evolved in response to patchy resource distributions.

1. It is often assumed that the timing of annual migrations of marine fish to spawning grounds occurs with very little change over time. However, it is unclear how much migration is influenced by climate change in marine species that spawn at sea but spend most time in estuarine conditions, especially as thermal regimes in estuaries may differ significantly from those in the open sea. 2. Migration phenology was studied in a population of flounder, Platichthys flesus (L.) off south-west England using high-temporal resolution trawling data over a 13-year period. 3. Flounder migrated from their estuarine habitat to spawning grounds at sea some 1-2 months earlier in years that were up to 2degreesC cooler. Flounder arrived on the spawning grounds over a shorter time period (2-6 days) when colder than normal conditions prevailed in the estuary, compared to warmer years (12-15 days). This suggests that they were responding to low temperatures by exhibiting a more synchronous, population-level early migration. 4. The timing of migration was earlier when the largest differences in temperatures between near-estuary and offshore environments occurred, differences that were related significantly to cold, negative phases of the North Atlantic Oscillation (NAO). 5. Flounder migration phenology appears to be driven to a large extent by short-term, climate-induced changes in the thermal resources of their overwintering habitat. This suggests that climate fluctuations characterizing the NAO may have significant effects on the timing of the peak abundance of fish populations generally, which, in turn, may have implications for fisheries management.