IOP PUBLISHING JOURNAL OF PHYSICS A: MATHEMATICAL AND THEORETICAL doi:10.1088/1751-8113/46/25/254023 J. Phys. A: Math. Theor. 46 (2013) 254023 (21pp) Ecological implications of eddy retention in the open ocean: a Lagrangian approach Francesco d’Ovidio 1 , Silvia De Monte 2,3,4 , Alice Della Penna 1,5 , Cedric Cotté 1,6 and Christophe Guinet 6 1 Laboratoire d’Océanographie et du Climat: Expérimentation et Approches Numériques, Institut Pierre Simon Laplace, CNRS/UPMC/IRD/MNHN, Paris, France 2 École Normale Supérieure, Unité Mixte de Recherche 7625, Écologie et Évolution, 46 rue d’Ulm, 75005 Paris, France 3 Université Pierre et Marie Curie-Paris 6, Unité Mixte de Recherche 7625, Écologie et Évolution, CC 237-7 quai Saint Bernard, 75005 Paris, France 4 Centre National de la Recherche Scientifique, Unité Mixte de Recherche 7625, Écologie et Évolution, 46 rue d’Ulm, 75005 Paris, France 5 Univ. Paris Diderot, Sorbonne Cité, Paris, France 6 Centre d’Études Biologiques de Chizé, Centre National de la Recherche Scientifique, 79360 Villiers en Bois, France E-mail: [email protected] Received 29 August 2012, in final form 13 December 2012 Published 4 June 2013 Online at stacks.iop.org/JPhysA/46/254023 Abstract The repartition of tracers in the ocean’s upper layer on the scale of a few tens of kilometres is largely determined by the horizontal transport induced by surface currents. Here we consider surface currents detected from satellite altimetry (Jason and Envisat missions) and we study how surface waters may be trapped by mesoscale eddies through a semi-Lagrangian diagnostic which combines the Lyapunov approach with Eulerian techniques. Such a diagnostic identifies the regions of the ocean’s upper layer with different retention times that appear to influence the behaviour of a tagged marine predator (an elephant seal) along a foraging trip. The comparison between predator trajectory and eddy retention time suggests that water trapping by mesoscale eddies, derived from satellite altimetry, may be an important factor for monitoring hotspots of trophic interactions in the open ocean. This article is part of a special issue of Journal of Physics A: Mathematical and Theoretical devoted to ‘Lyapunov analysis: from dynamical systems theory to applications’. PACS numbers: 47.52.+j, 92.20.jm (Some figures may appear in colour only in the online journal) 1751-8113/13/254023+21$33.00 © 2013 IOP Publishing Ltd Printed in the UK & the USA 1 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al 1. Introduction Remote sensing and in situ observations of the ocean and atmosphere show strong variability in physical and biogeochemical properties at several spatiotemporal scales. This variability has been associated with the presence in the velocity field of coherent structures that create regions of retention (tracer reservoirs) or of mixing [73]. Several diagnostics have been proposed to identify Eulerian and Lagrangian coherent structures and to detect their effect on the distribution of an advected tracer field [9]. The Lyapunov exponent, in particular, has been shown to be very effective in identifying regions of geophysical flow where the stretching rates are maximal, which are in turn candidate locations of tracer frontal structures and transport barriers. The structuring effect of the velocity field over physical or biogeochemical tracers is dramatically evident at the ocean surface on scales of tens to hundreds of kilometres, where satellite images of surface chlorophyll concentration or sea surface temperature display strong contrasts [44, 42] reminiscent of meteorological images of weather patterns. This regime, sometimes referred to as the ocean weather, covers the so-called mesoscale and part of the sub-mesoscale, so that in the following we will refer to it as the (sub-)mesoscale [40]. In this regime, the ocean surface velocity field is dominated by the presence of coherent eddies [17], i.e. axis symmetrical patches of strong vorticity with lifetimes ranging from several weeks to more than one year and typical radii of tens/hundreds of kilometres. The core and the periphery of these structures act in a very different way on the tracer’s redistribution. At the core of the eddies, water parcels tend to recirculate and are only marginally mixed with intruding ambient water. Therefore, under some conditions, eddy cores retain water parcels—and hence their biotic or abiotic content—for timescales of weeks/months, and sometimes even longer [11, 53, 55, 38, 16, 15, 64, 45]. At their peripheries, where strain prevails over vorticity, eddies have the inverse effect and enhance water mixing. In this case, tracer anomalies are redistributed by the eddy field in the form of filaments and become eventually mixed with the ambient waters on relatively fast timescales (days to weeks). The different structuring role of eddy cores and eddy peripheries is well captured by Lagrangian diagnostics such as finite-size or finite-time Lyapunov exponents, which analyze properties of the velocity field along particle trajectories. Such diagnostics typically identify lobular ridges of maximal stretching rates that delimit the eddy core. Depending on the intensity and the temporal variability of the eddy, different situations can arise. In the limiting case of no temporal variability, closed orbits are also material lines. In this case, Lyapunov-detected transport barriers associated with eddies form a closed curve, so that the eddy core is not permeable to lateral advection. When the velocity field changes in time, such closed barriers break into interleaved spirals along which ambient water can be exchanged with the interior of the eddy. Idealized quasi-geostrophic models of oceanic mesoscale circulation have shown that eddies can systematically trap and transport water parcels for timescales comparable to their lifetime, shielding their content from dispersion [11]. In the real ocean, however, eddies strongly interact with jets, topography and with each other. As a result, trapping of water parcels is a more transient phenomenon than in idealized models and it typically depends on a parcel being stirred by an ensemble of structures. Lagrangian and Eulerian analyses based on multisatellite data [39, 13, 23, 15] indicate that 10–100 km wide lobular patches of trapped tracers arise due to stirring by mesoscale currents, with lifetime shorter than the lifetime of eddies (days/weeks compared to weeks/months and more). Primary producers—typically phytoplankton, whose doubling time is of the order of a day—have been shown to be affected by such patches, which may act as fluid dynamical niches [21]. 2 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al Patchiness in the ocean is known to affect not only primary producers, but also higher trophic levels. It is indeed conjectured that mesoscale turbulence structures the distribution of higher trophic levels from zooplankton and fish [32, 65, 61] up to top predators (e.g., [72, 26, 68, 18, 46]). However, grazers and predators need some time to grow and aggregate over patches of high phytoplanktonic concentration. Therefore, in order to be found in association with eddy cores, their distribution may require retention times longer than those sufficient for confining primary producers. In particular, they should be mostly affected by water that remained confined for a time comparable with or longer than the duration of a bloom (about 30 days), which is the period during which phytoplanktonic biomass is accumulated. The situation of isolated and strong mesoscale eddies, where such a long retention condition may be met, is not typical in the ocean but does occur (e.g., [17]). This is the case for instance of the eddies shed by energetic currents like the Gulf Stream and the Antarctic Circumpolar Current (so-called Rings), or for eddies trapped by the topography like the first Alboran gyre in the West Mediterranean and the Great Whirl off the Somali coast [58]. Indeed, these eddies have been found to be associated with spot-like anomalies of physical or biogeochemical properties like sea surface temperature, chlorophyll and even with peculiar phytoplanktonic community structures [6, 45, 21]. Due to their long retention times and their observed structuring role on primary producers, it is natural to ask whether eddies with a long retention time are also hotspots of trophic interactions. In this paper, we aim at exploring the role of trapping by altimetry-derived mesoscale eddy cores on trophic interactions. In order to achieve this, we start by quantifying retention times from real data. Lagrangian diagnostics like the Lyapunov exponents can probe the permeability of an eddy core, but they are not informative of its content. On the other hand, Eulerian quantities like the vorticity identify the core of an eddy but not its retention time. Following an approach which has been proposed on model systems [27] and generalizing the calculation of Lehahn and co-authors [38], here we combine the Lagrangian and Eulerian approaches and define a diagnostic able to estimate the retention time of mesoscale eddies observable from satellite-derived surface currents. After discussing how the temporal variability affects retention times, we consider the case of the energetic eddies present in the meanders of the Antarctic Circumpolar Current. We locate eddies with different retention times, compute the surface which they cover climatologically and compare them with the foraging behaviour of a tagged elephant seal, which made a long foraging trip (thousands of kilometres) in a region with eddies of contrasted characteristics. Remarkably, this predator halted and foraged intensively in an eddy with large retention time (>30 days), but did not stop when crossing other eddies with retention times of a few days. Section 2 is dedicated to the introduction of the key observational and theoretical tools used in the paper, in particular, the Okubo–Weiss (OW) parameter and the finite-size Lyapunov exponents (FSLEs), and provide examples of the applications of such different diagnostics to the ocean surface velocity field derived from satellite altimetry. Section 3 introduces the retention parameter as a semi-Lagrangian diagnostic which integrates the information provided by a Lagrangian approach and the OW parameter. Section 4 deals with the coupling of marine ecosystems and of the physical variability at the sub-mesoscale, applying the retention parameter to the analysis of a foraging trip of an elephant seal through the meanders of the Antarctic Circumpolar Current East of Kerguelen Islands. 2. Surface currents and tracer distribution in the ocean 2.1. Satellite altimetry and the Lyapunov exponent calculation Since the launch of ERS-1 and Topex-Poseidon altimetry satellites in the 1990s, information on the oceanic surface circulation is almost globally accessible from remote sensing. The 3 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al -2 [cm] 80 -46 60 [days ] 0.2 -46 0.15 40 -48 0.1 -48 0 -50 0.05 lat lat 20 0 -50 -20 -52 -0.05 -52 -40 -60 -54 70 75 80 lon 85 90 95 -0.1 -0.15 -54 -80 70 75 80 70 75 80 lon 85 90 85 90 95 -0.2 -1 [days ] 0.2 -46 -46 0.15 0.1 -50 -52 lat -48 lat -48 -50 -52 0.05 -54 -54 70 75 80 lon 85 90 95 0 lon 95 Figure 1. Four different analyses of the surface velocity field in a region around the Kerguelen Islands. (a) Sea surface height (SSH) measured by satellite (AVISO, Kerguelen product) and superimposed surface velocity field, obtained by assuming that the flow is quasi-geostrophic. (b) Okubo–Weiss parameter computed on the velocity field of panel (a). Blue regions indicate points where vorticity dominates over strain (e.g. the eddy cores), red regions the opposite. The OW parameter is a Eulerian diagnostic, and its computation is based on the instantaneous flow. (c) Backward finite-size Lyapunov exponents map: higher values of the dominant exponent indicate higher stretching of water parcels, and identify potential locations of tracer fronts. (d) Distribution of a virtual, passive tracer (red dots) advected by the (time-varying) flow of panel (a) after being initialized with uniform concentration inside the region delimited by the blue lines. The tracer distribution fronts are largely aligned with the FSLE ridges (black lines). principle behind satellite altimetry is to measure the sea surface height (SSH, figure 1(a)) above a reference geoid and to compute geostrophic velocities by balancing the pressure gradient (which depends on the gradient of the SSH) with the Coriolis force (which depends on the horizontal velocity, the unknown to be retrieved). This is formally equivalent to the definition of a time-varying two-dimensional Hamiltonian system, where the SSH corresponds to the Hamiltonian. The spatiotemporal resolution of the SSH estimation—and hence of the derived velocities—depends on the number of altimetry satellites in orbit at the same time and on their orbit characteristics. The error depends on the precision of the on-board altimeters, the orbit estimation, the precision at which the shape of the reference geoid is known and the algorithms which process the satellite signals. More technical information on satellite altimetry, including the characteristics of past, current and incoming altimetry constellation, 4 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al can be found at the CNES/CLS AVISO website www.aviso.oceanobs.com [28]. Although all these parameters constantly change, altimetry is considered reliable for the detection of geostrophic circulation structures of about 70–100 km and lifetimes of at least a week—hence for mesoscale studies [36]. Note that satellite altimeters are radar instruments and are therefore unhindered by cloud coverage. The availability of two-dimensional ocean surface velocity fields on an extended and uninterrupted spatial and temporal domain (tens of years with almost a global coverage) has opened the way to the application of several mathematical approaches originally developed for the analysis of dynamical systems or idealized geophysical flows (see for instance the reviews by Wiggins [74] and Samelson [62]). The main aim of these studies is the identification of the key mesoscale properties and dynamical structures which control transport and mixing at the ocean surface and which may link the current field to the biogeochemical dynamics of the ocean. We briefly review chronologically some of these previous results, focusing on the Lyapunov exponent calculations applied to altimetry data. Following some previous applications to atmospheric or model systems (e.g., [56]), probably the first calculation of Lyapunov exponents from altimetry data has been made by Abraham and Bowen [1], who estimated mean stirring rates and showed that the ridges of Lyapunov exponents were in close agreement with fronts of sea surface temperature. In order to further explore the relation between stirring and sea surface temperature, Abraham and Bowen also advected a tracer with either a longitudinal or a latitudinal gradient, and applied to the ocean dynamics a formalism which was proposed for chaotic advected chemical fields [49]. This approach was further developed for the case of the Tasman sea [71] and then to the global oceans [70], where a strong correlation was found between Lyapunov exponents and Eulerian quantities like the strain rate and the eddy kinetic energy. A second regional study (East Mediterranean) [23] compared Lyapunov exponents and Eulerian diagnostics at submesoscale-resolving resolution, showing the superior ability of the former in detecting filaments and fronts associated with sea surface temperature. The study also concluded that the correlation between Eulerian diagnostics and Lyapunov exponents does not hold when basin-scale climatologies are computed by averaging submesoscale-resolving maps. Transport barriers derived from altimetry by computing the finite-size Lyapunov exponent have been compared to satellite-derived [39, 38] and modelled [57] chlorophyll patches. These comparisons showed the structuring role of stirring on primary production and in particular provided evidence of how the temporal variability of the mesoscale velocity field can break closed orbits associated with mesoscale eddies and create submesoscale spiralling patterns visible in satellite chlorophyll images. This difference between closed Eulerian eddies and spiralling Lagrangian structures was later described also by Beron-Vera et al [7] who compared ridges of Lyapunov exponents with the trajectory of one Lagrangian drifter. A larger number of drifter trajectories and transport barriers were then compared by Resplandy et al [58] who combined in situ, altimetry and a biogeochemical circulation model to interpret the strong pCO2 variability observed by CARIOCA drifters, in terms of stirring and transport barriers. The biogeochemical role of horizontal stirring was further investigated by Calil et al [13] and Rossi et al [60, 59] where the Lyapunov analysis was used to link the horizontal stirring dynamics to upwelling and primary production. In particular, Calil et al showed that ridges of Lyapunov exponents can be associated with hotspots of primary production in oligotrophic waters, by either creating frontal regions where nutrients can be upwelled by secondary circulation or by structuring in filament patches of blooming waters. In spite of the large number of case studies in which altimetry-derived Lyapunov ridges have been shown to visually match fronts of physico-chemical or biological fields (with some exceptions: see Nencioli et al [48] for dramatic errors in stirring patterns derived from 5 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al altimetry along the coast), there are not many pieces of work where the robustness of the altimetry-derived Lyapunov analysis has been assessed. Hernández-Carrasco et al [29] tested the effects of noise and of the dynamics of unresolved scales, using data from a primitive numerical model as a benchmark. Lacorata et al [33] already in 1996 compared Lyapunov exponents computed from drifters and from model data showing agreement between the two diagnostics. However, to our knowledge, the only systematic validation of Lyapunov-derived frontal regions and in situ fronts detection has been done by Dèspres et al [20], who found that the front spatiotemporal distribution of tracers like temperature or salinity can be predicted by Lagrangian analyses of altimetry data only if information on the large-scale tracer distribution is also known. The effect of noise on altimetry-based Lyapunov calculation has been also briefly discussed by Cotté et al [18]. The spatial structure of the Lyapunov ridges was shown to be highly robust to delta-correlated, high-frequency noise (which mimics small-scale diffusion), and an analytical scaling relation integrating noise intensity in the value of the Lyapunov exponents was proposed. More recently, altimetry-derived Lyapunov exponent calculations have been used to explore the ecological role of horizontal stirring in ecological processes. Tew Kai et al [31] for the first time related Lyapunov ridges to the distribution of a top marine predator (Fregata minor) of the Mozambique Channel. Using satellites to track the position of eight frigatebirds, the authors provided evidence that these predators co-localized with Lyapunov ridges. Thus, they argued that coherent Lagrangian structures may be detected by visual, olfactory or wind cues. The co-localization between position of frigatebirds and Lyapunov ridges was recently revisited by De Monte and co-authors [19] who used higher resolution (GPS), behaviour-resolving tracking data, and showed that behavioural switches occur in the presence of Lyapunov-detected transport fronts especially when they coincide with thermal fronts. An important role of Lyapunov ridges for marine predators was further evidenced by Cotté et al [18], who analysed the relation between Mediterranean whales and ocean physics in the Mediterranean sea at various spatiotemporal scales. The authors found that whales were associated with Lyapunov ridges only during the summertime phytoplankton minimum, suggesting that patchiness in food resources induced by stirring may be especially critical for marine predators during oligotrophic conditions. In the same year, d’Ovidio et al [21] compared remotely sensed dominant phytoplankton types and altimetry-derived transport structures. This analysis showed that the phytoplanktonic landscape is organized in (sub-)mesoscale patches of dominant types separated by physical fronts. These fronts, induced by horizontal stirring, are identified from altimetry data through a Lyapunov analysis. The authors named ‘fluid-dynamical niches’ such water patches delimited by Lyapunov ridges and supporting different dominant types. The concept of a water patch which maintains its integrity for several weeks and longer has been also exploited by biogeochemical campaigns, which have tracked and sampled such semi-closed systems, viewing them as megacosms. This approach has allowed for instance the characterization of the biogeochemistry and ecology of an artificially fertilized water patch from the beginning of the bloom to the export of organic material to the deep ocean at the end of the bloom [69, 12]. 2.2. Eulerian and Lagrangian diagnostics We introduce here the formal definitions of the diagnostics that will be used in the following sections, and that are possibly the most widespread tools for detecting the structures generated in the ocean by horizontal transport. For a more extensive introduction to these and other statistics of a velocity field, we refer to [9] and [23]. 6 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al Given a streamfunction ψ (x, y, t ) := g/ f H, where g is the acceleration due to gravity, f is the Coriolis parameter which takes into account the Earth’s angular velocity and H is the SSH with respect to a reference geoid, the geostrophic velocities are defined as ∂ψ , (1) vx (x, y, t ) = − ∂y ∂ψ , (2) ∂x where x, y and t are respectively longitude, latitude and time. It is easy to check that such a velocity field has zero divergence for all the scale lengths at which f (and g) can be considered constant. The OW parameter compares the intensity of the strain to the vorticity: vy (x, y, t ) = OW(t, x, y) := s(t, x, y)2 − ω(t, x, y)2 , where s is the strain and ω is the vorticity. The strain and vorticity are obtained from the derivative of the velocity field as ∂vy 2 ∂vy ∂vx 2 ∂vx 2 − + s = + , ∂x ∂y ∂x ∂y (3) (4) ∂vy ∂vx 2 ω = . (5) − ∂x ∂y It can be shown that the OW parameter is equal to the determinant of the Jacobian of the velocity field multiplied by −4. When calculated at a stagnation point of the velocity field, it is hence negative for an elliptic point (with two complex conjugated eigenvalues) and positive for a hyperbolic point (with two real and opposite eigenvalues). The OW parameter can therefore be seen as a way to extend the properties of a stagnation point to its neighbourhood. The region of the plane (x, y) where OW(t, x, y) < 0, and hence recirculation prevails over stretching, can be taken as an operative definition of an eddy core [17, 30, 47, 23]. The isolines of the OW parameter are used as a tool for identifying the Eulerian eddy boundaries. The OW parameter cannot however account for the generation of the complex filamentous structures induced by chaotic stirring on a tracer field, since these structures depend on the temporal variability of the velocity field. In order to study these filaments, it is necessary to move from the Eulerian framework to a Lagrangian one, using statistics computed over particle trajectories, such as the Lyapunov exponent. This way, one can reconstruct the advection history of a tracer field, integrating spatial and temporal information along the trajectory of a material point. This approach has another advantage. Tracer filaments produced by chaotic stirring and retrieved by Lagrangian techniques occur typically at spatial scales smaller than the size of mesoscale eddies. Therefore, Lagrangian techniques are a way to explore a regime which is not directly resolved by altimetry data. We recall here some general concepts related to Lyapunov exponents and fluid dynamics applications. More details can be found for instance in [37] and in the tutorial by S Shadden (http://mmae.iit.edu/shadden/LCS-tutorial/). The trajectory of a material point advected by the flow can be obtained by solving the differential equations (1) for the boundary condition x(t0 ) = (x0 , y0 ), which provides the position of the Lagrangian particle at time t0 . Given that δ0 represents the distance between x(t0 ) and a nearby point and that δ represents the separation at a time t of the trajectories originating at those points, Lyapunov exponents are formally defined [52] by the double limit δ 1 (6) λ = lim lim ln . t→∞ δ0 →0 t δ0 2 7 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al This expression allows us to discriminate chaotic and non-chaotic domains of ergodic dynamical systems. However, the limit is exactly determined only when the velocity field is known at all times and all points in the plane. In order to apply the Lyapunov approach to a field that, as derived from altimetry data, has a limited spatiotemporal resolution, the relaxation of one of the two limits in the definition of the Lyapunov exponents is necessary. One can either prescribe the integration time t leading to the finite-time Lyapunov exponent (FTLE) or a spatial scale (the ratio δ/δ0 ) leading to the finite-size Lyapunov exponent (FSLE). Both these quantities approach, in the limit of complete time-scale separation, the Lyapunov exponent defined by equation (6). In this work, we will use the dominant FSLE which has been introduced in [4, 3, 9]. (For more details, see appendix A in [24].) Oseledec’s theorem states that if the flow is ergodic, then the limit in equation (6) is a unique λ∗ irrespective of the perturbation applied to a point [51] and for almost all initial conditions on the ergodic set. Such asymptotic value is positive for points belonging to chaotic domains. For finite times and separations, however, spatial homogeneity no longer holds. This turns out to be an advantage for geophysical applications, because the dominant FSLE assumes different values for different positions in space, revealing regions with different kinematic properties. In the case of mesoscale turbulence, it is not difficult to choose an integration time or particle separation for FTLE and FSLE such that most of the calculations occur in the linear regime. There the finite-time and the finite-size calculations provide in practice a similar outcome (compare for instance FTLE maps in [7] with FSLE maps in [39]). Although we do not use this approach here, we also mention that another use of the FSLE is the exploration of different dispersion regimes [4, 3, 9, 67]. The computation of the dominant FSLE can be performed either forward or backward (when t < 0) in time. Here, we will focus on the dominant FSLE computed backward in time, since it allows us to detect the barriers to transport in the flow. 2.3. Okubo–Weiss parameter and Lyapunov exponent applied to altimetry-derived velocities An example of a snapshot of the mesoscale velocity field derived by satellite altimetry is shown in figure 1(a) in terms of SSH (colour) and geostrophic velocity (arrows). The panel depicts the Kerguelen region (Indian sector of the Southern Ocean). The Antarctic Circumpolar Current—one of the most energetic currents of the world—is steered along its eastward flow by the Kerguelen plateau and splits into several meandering branches, creating a very contrasted mesoscale field. There is a low-energetic region on the wake of the plateau (70◦ –85◦ E) followed eastward by strong and dynamically active eddies (85◦ –100◦ E). Mesoscale eddies can be evidenced as closed streamlines with a radius of about 100 km. A map of the OW parameter for this flow is shown in figure 1(b). This image depicts the eddy field (blue spots) which is at the origin of horizontal stirring. In order to estimate the effect of these eddies over any advected tracer, a Lagrangian analysis is necessary. Figure 1(c) shows an example of FSLE calculation backward in time, where the map is obtained by measuring the rate of separation of particles initialized in nearby positions. In this example, the parameters for the computation of FSLEs are δ0 = 2.5 km and δ = 60 km. This choice is related to the spatial scales of the system: since we are interested in mesoscale structures, the final separation δ is chosen as a typical mesoscale structure’s dimension, whereas δ0 is chosen close to the resolution of the grid [22]. The timescale t for going from a separation of δ0 to δ is related to the mixing properties of different regions: typical values are about 1–4 weeks, corresponding to FSLEs in the range [0,0.14] days−1. 8 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al An intuitive understanding of the physical meaning of FSLE maxima (ridges) can be gathered by examining the advection of a passive tracer (figure 1(d)). A backward-in-time large separation of particles initialized nearby means forward-in-time convergence of water parcels that were located far apart from each other. If these water parcels had contrasted properties, such as different tracer concentrations, the confluence dynamics would result in the amplification of the tracer gradient, because the distance between the water parcels, and the tracers that they transport, is reduced. Therefore, a ridge of large exponents can be found in association with a biogeochemical front. When there is no temporal variability, the ridges of the Lyapunov map coincide with the homoclinic and heteroclinic orbits surrounding elliptic points, and thus provide a separation of the plane into elliptic or hyperbolic domains that is consistent with what one would get by connecting hyperbolic points with streamlines. 2.4. Horizontal stirring of biogeochemical tracers and Lyapunov analysis A powerful application of the Lyapunov exponent calculation in oceanography is the identification of fronts of tracers advected by the mesoscale (tens to hundreds of kilometres) velocity field. Note however that lines of large Lyapunov exponents do not necessarily have a one-toone correspondence to the fronts of a tracer. Indeed, horizontal transport can only intensify pre-existing heterogeneities, such as large-scale tracer gradients. For this reason, ridges of Lyapunov exponents (which may be considered as transport or kinematic fronts) are typically much more numerous than tracer fronts. There are also several factors which may affect the effectiveness of the Lyapunov exponents calculation for the detection of tracer fronts, namely (i) if the large-scale tracer gradient is oriented orthogonally to the stretching direction, then it is not amplified; (ii) the tracer has an active dynamics so that initial contrasts in converging waters are not preserved during advection; (iii) transport mechanisms different from stirring by mesoscale turbulence are also active (e.g., upwelling and downwelling, convection); and (iv) altimetry-derived geostrophic velocities do not perfectly represent horizontal stirring (e.g., in terms of resolution and in terms of missing ageostrophic components). These factors, which are often all present at the same time, may induce one to think that the application of the Lyapunov exponents is hopeless in any realistic case. Nevertheless, several studies (see section 2.1) have shown empirically that the aforementioned factors do not compromise the application of the Lyapunov exponent calculation, although misplacements of the fronts of the order of 10–20 km can typically arise. A comparison between a numerically advected tracer and the FSLE field of figure 1(c) is shown in figure 1(d). The tracer patch, initialized within a box, is structured in convoluted filaments typical of chaotic advection. The contours of these filaments are almost perfectly identified by ridges of FSLEs. As previously remarked, however, the Lyapunov ridges are much more numerous than the tracer filament boundaries (which depend on the specific initial conditions). Note that by visually inspecting the tightness of the spiralling ridges, it is not difficult to guess which eddies may have longer lived cores; however, the Lyapunov exponent map neither quantitatively provides this information nor indicates which points lie inside or outside an eddy core. Figure 2 shows an example of the matching between a real tracer, chlorophyll, and FSLE ridges. Even if the tracer is not conservative (that is, its local concentration changes in time) and the currents are actually affected by both ageostrophic dynamics (notably, wind-induced Ekman transport) and unresolved geostrophic velocities, then the fact that such a correspondence holds provides yet another empirical confirmation of the relevance 9 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [µg/l] 3 -46 1.5 -48 1 lat 0.75 -50 0.5 -52 0.2 -54 70 75 80 lon 85 90 95 0.1 Figure 2. Chlorophyll (Chl) distribution (from water-leaving radiance, SeaWiFS) and ridges of FSLEs, in the same region as figure 1. The FSLE ridges show remarkable agreement with Chl fronts, in spite of the fact that Chl is not a passive tracer. The possibility of matching Chl tracer fronts with FSLE lines lies in the fact that the tracer evolves on a slower timescale than the velocity field. It is worth noting that the actual distribution of Chl essentially depends of the initial location of the phytoplankton bloom, that is, in the case illustrated here, localized on the Kerguelen plateau. of Lagrangian approaches to the study of the oceanographic dynamics, at least at the (sub-)mesoscale. 3. Time-varying velocity fields: estimating the timescale of eddy trapping Let us first discuss the idealized case of a time-invariant streamfunction (this can be a good approximation of a long-lived eddy trapped by the bathymetry). The stagnation points of the velocity field (equilibria of equations (1)) play a structuring role for the circulation. In particular, the invariant manifolds stemming from hyperbolic, saddle points act as transport barriers and are thus able to constrain the displacement of particles advected by the surface currents. Imagine that a tracer is present in the water at concentration c(x, y), and that it is conservative (the tracer is not produced or consumed) and passive (it does not affect the velocity field). If diffusion is negligible, as when considering scales above a few kilometres in the ocean, its dynamics is described by the advection equation ∂c = −v · ∇c. (7) ∂t When the velocity field is independent of time, the tracer cannot cross a streamline (line of constant ψ), since ∇c ⊥ v. On the other hand, the invariant manifolds (homoclinic or heteroclinic orbits) of the saddle points are streamlines that connect equilibria, and can enclose regions that contain elliptical stagnation points. These domains correspond to oceanic eddies, within which the water circulates, and are separated by the free water that flows outside their boundaries. It is easily shown that if the velocity field is time-independent, then the total 10 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al concentration C = c dω of a tracer within a region , whose contour ∂ is given by streamlines, is conserved: ∂c dC = dω = − ∇(vc) dω = − (vc) · n ds = 0. (8) dt ∂t ∂ If applied to a homoclinic orbit, such as the contour of certain eddies in time-invariant fields, this means that the tracer content of the region it contours is constant in time. As soon as the field changes in time, however, the correspondence between streamlines and particle trajectories does not hold any longer and particles can escape from eddy cores. Still, the structure of a time-invariant field is maintained up to a certain approximation (related to the observation time scale), so that the regions that have high retention times (typically, eddy cores) and other regions (typically, eddy peripheries) where instead mixing is dominant exist. We follow here some previous works that have addressed the issue of distinguishing retention zones with different mixing properties in idealized time-varying vector fields [35, 27, 34, 50, 66, 43]. We adopt a hybrid Lagrangian and Eulerian approach to give a definition of the time during which particles remain trapped within eddies. In this section, we introduce a retention parameter (RP) and discuss its relationship with the OW parameter and FSLEs. In section 4, we will then propose an application to the case study of a foraging trip of a tagged top marine predator. With RP at a given time t0 and at a point (x0 , y0 ) we indicate the number of days a water parcel at that given location has remained confined within an eddy core, that we define as a region where the OW parameter is negative. This amounts to computing the OW parameter OW(t, x, y), with t ∈ I = {t0 −tback , t0 }, along the trajectory (x(t ), y(t )) obtained by integrating backwards in time for a time tback the velocity field with boundary values (t0 , x0 , y0 ), and finding the time when OW changes sign, indicating that the particle has left the eddy core. If we define te = max {t ∈ I|OW(t, x(t ), y(t )) = 0} the first singular value of the OW parameter, then we can formally define the RP as RP(t0 , x0 , y0 ) = t0 − te , or equivalently, and in analogy to [27], as t0 RP(t0 , x0 , y0 ) = dt. (9) (10) te Note that this definition is different from the ‘ellipticity time field’ proposed by Haller [27], since the latter was based on a measure of the ellipticity properties along the trajectory, and the integration takes place at all times that the trajectory spends in a region of given topological properties. In our case, instead, we decided to base our definition on an Eulerian statistics, the OW parameter, since this is what is most commonly used in oceanography for identifying eddy cores (see section 2.4). Also, in the definition we propose here, we do not allow a particle to leave and re-enter eddy cores. The rationale behind this choice is the following: we assume that as soon as a water parcel gets in a hyperbolic region (i.e. outside eddy cores), its content is stretched and mixed, so that its specific content is lost. Figure 3 illustrates an example of the calculation of the RP (with a maximum integration time tback = 60 days) computed for two positions in space. In both cases, the backtrajectories have been obtained by backward integration in the same time interval, and plotted together with the sign of the OW parameter calculated along them. Note that the blue segment (negative OW) corresponds to the time in which the particle is circulating in an eddy core, that is, the value of the RP. 11 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al Figure 3. Illustration of the rationale behind the definition of the retention parameter: such a diagnostic is meant to quantify the amount of time a water parcel has remained in a confined region of the ocean. Backtrajectories (tback = 60 days) originating from two points (red dots) situated at time t0 in nearby eddies are represented together with the sign of the OW parameter calculated along them (blue, negative; red, positive). The values of the retention parameters associated with these two points are displayed, showing that the point with higher retention parameter is associated with a trajectory spinning around an eddy core, whereas a smaller retention parameter is associated with a water parcel that has been more recently trapped within an eddy. 3.1. Retention times of altimetry-detected mesoscale eddies What are the retention times of oceanic mesoscale eddies? We address this question by constructing some instantaneous and climatological maps of the RP for the Kerguelen region. In order to evidence how properties of oceanic eddies are influenced by the temporal variability of the velocity field, for any time t0 at which we compute a map, we artificially tune the timescale of the currents derived from satellite observation. We rescale the time variable so as to generate a family of fields v(t, t0 , ) = v(t0 − (t0 − t )) such that ∈ [0, 1] sets the variability timescale: when it is zero, the velocity field is frozen at all times under the condition observed at time t0 : v(t, t0 , 0) = v(t0 ) ∀t. All eddy cores have the same, maximal, value of RP (equal to tback ), since the water within them recirculates indefinitely. If is unitary, then the field has its natural temporal variability: v(t, t0 , 1) = v(t ). Such time rescaling does not affect Eulerian measures at time t0 , which do not depend on the state of the velocity field in the future or in the past. Lagrangian diagnostics, however, are modified, since the trajectories are now computed over a field that is slowed down according to the parameter . Figures 4(a)–(c) display the dominant FSLE for an observed velocity field timed at different values of . As expected, the increased exchange of water between an eddy and its surroundings reduces the retention time of most eddies identified by the Eulerian statistics. We find however a surprising result. In some cases, the variability in time can generate retention regions that were not present in the time-invariant case. A possibility may be an eddy at the end of its lifetime. Backward trajectories for the time-invariant case only experience the weak vorticity 12 J. Phys. A: Math. Theor. 46 (2013) 254023 −46 (a) F d’Ovidio et al (b) FSLE [days−1] (c) −48 0.1 0.05 latitude −50 0 (e) −46 (d) RP [days] (f ) 60 −48 40 20 −50 77 79 81 77 79 81 77 79 81 0 longitude Figure 4. (Semi-)Lagrangian diagnostics of an altimetry-derived velocity field with artificially tunable temporal variability, as explained in the text. (a)–(c) Dominant finite-size Lyapunov exponent (δ0 = 1 km, δ = 40 km); (d)–( f ) retention parameter computed for tback = 60 days. From left to right the variability in time increases: (a), (d) = 0, (b), (e) = 0.5, (c), ( f ) = 1. In the last case, the temporal variability reflects the variation of the measured velocity field. Slow variations of the velocity field do not substantially alter the landscape one would obtain for a frozen field, where transport barriers contour recirculation regions. Time variability instead can erase, or even generate (e.g. around 50.5◦ South, 78.5◦ East), retentive regions, such processes being related to the opening and folding of transport barriers. associated with this vanishing eddy and escape the eddy, hence resulting in a short retention time; trajectories constructed in the time-dependent case however also experience the stronger vorticity associated with the eddy at a younger life stage and therefore may recirculate in the eddy for a longer time, thus resulting in a larger retention time. The complex interplay of eddy core intensity and temporal variability is further confirmed by a climatologic analysis (figure 5). The comparison of patterns of retention and patterns of OW, averaged in time for one year, clearly shows that there is no clear association between regions of largest negative OW and regions of strongest retention. Indeed, attempting to produce a scatter plot does not provide any compact relation between the two quantities (not shown). Figure 5 suggests that retention times in excess of one and even two months—therefore likely able to structure secondary and higher production, as discussed in the introduction—are rare but possible. We quantify their occurrence by considering all the daily maps of retention for one year in the Kerguelen region and constructing the cumulative probability of retention times. We perform the calculation both for = 0 and = 1 and plot the results in figure 6. This distribution indicates that cores older than one month cover about 3% of the ocean 13 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [d-2] 0.3 [d] 30 -46 -46 25 lat 20 15 -50 10 -52 -48 0.1 lat -48 0.2 0 -50 -0.1 -52 5 -0.2 -54 70 -54 75 80 lon 85 90 0 95 70 75 80 lon 85 90 -0.3 95 Figure 5. Comparison between the retention parameter (left) and Okubo–Weiss parameter (right) averaged over one year and computed on the same region of the ocean. The patterns do not show a clear correlation between regions of largest negative Okubo–Weiss and highly retentive areas. 0 0 10 10 -1 Area covered Area covered 10 10-1 10-2 -2 10 0 -3 10 20 30 retention > days 40 50 60 10 0 10 20 30 retention > days 40 50 60 Figure 6. Cumulative distribution of the retention parameter for no time variability of the velocity field = 0 (left) and for natural time variability = 1. The importance of the time variability of the velocity field for retention issues is underlined by the fact that in the case of natural temporal variability eddy cores that are older than one month cover about 3% of the ocean surface around the Kerguelen region, whereas in the case of a frozen velocity field, one gets approximately double the value. surface around the Kerguelen region. This value would approximately double if the temporal variability of the velocity field were absent. Considering that an eddy core is about 100 km wide and supposing that the eddies are randomly distributed, this means that a linear path encounters on average one core with RP larger than 1 month every 3000 km. 4. Eddy retention and trophic interaction: an elephant seal case study The observation of top marine predator displacements is an important tool of marine ecology. High trophic levels indeed play the role of ‘integrators’ of the food chain, since their ecology is altered by misfunctioning of the ecosystem [14]. If the underlying ecological processes, which relate the predator behavioural response to changes in its environment, are known, one can 14 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [m] 0 -46 -1000 -48 lat -2000 -50 -3000 -52 -4000 -54 70 75 80 lon 85 90 95 -5000 Figure 7. Trajectory of an elephant seal trip, starting from the Kerguelen Islands on 23 January 2009 and ending when the recording stopped on 14 March 2009, superimposed on the bathymetry of the Southern Indian Ocean. The trajectory becomes more tortuous and the animal slows down when it comes close to an eddy. These features of the azimuthal trajectory of elephant seals is typically associated with an increase in capture events, indicating intensive foraging activity [26, 54]. The three red dots indicate the position of the elephant seal at three dates two weeks apart (1 February, 14 February and 1 March 2009). hope to infer from physical features the modification in the predator habitat, and consequently of its trophic function. This however requires to clarify not only the statistical correlations with environmental patterns at the large scale, such as for instance with sea surface temperature in migrating animals [8], but also at the finer scale on which the animal perceives the physical features of its habitat [19]. In this section, we provide an example of the use of previously discussed diagnostics of horizontal stirring, the FSLE and the RP, for investigating a top predator’s behaviour at the (sub)mesoscale. 4.1. How do elephant seals move in a turbulent field? Elephant seal (Mirounga leonina) populations of the Kerguelen Islands (49◦ South, 69◦ East) have been the object of long-term demographic monitoring that has been complemented in recent years by satellite tracking at increasing temporal and spatial resolutions. Here, we analyse the post-moulting trip of an adult female equipped with a ARGOS tag, who left the Kerguelen Islands on January 2009. The animal position is measured with a resolution of 1 km and 1/4 day, for a total duration of the recording of 79 days [25]. Elephant seals, and particularly females, forage in the open ocean, where they alternate hundreds of metres deep diving in search for fish or squid, and short breathing intervals at the surface (with a total period of about 20 min). After moulting on land, females leave their colony to engage in solitary trips, lasting several months, where they typically head towards a zone of strong mesoscale activity between the subtropical and the subpolar fronts (see figure 7 15 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al −48 (a) −48 (b) −48.5 (c) −49 −49 −49.5 −50 −50 −50.5 OW −2 [days ] 2 0 latitude 78 79 80 78 79 80 −2 90 −48 (d ) −48 (e) −48.5 (f ) −49 −49 −49.5 −50 −50 −50.5 78 79 80 78 79 80 90 −48.5 (i ) −49 −49 −49.5 −50 −50 −50.5 80 78 79 80 FSLE [days−1] 0.05 −48 (h) 79 92 0.1 −48 (g) 78 91 91 92 0 RP [days] 50 90 91 92 0 longitude Figure 8. Physical diagnostics derived by remote-sensing of sea surface height for three days during the elephant seal trip (see figure 7 for their locations along the trajectory). The dot indicates the position of the animal along the trajectory on a given day (specified in the panel title). (a)–(c) Okubo–Weiss parameter: eddy cores are characterized by negative values. (d)–( f ) Finitesize Lyapunov exponents (δ0 = 1 km, δ = 40 km): larger values identify stronger transport barriers. (g)–(i) Retention parameter: measures the number of days the trajectory stemming from a point and computed backwards in time remains within the same eddy, that is larger for longer-lasting structures. for an overview of the trip analysed here) [5]. This region is the target for foraging trips not only of elephant seals, but of many predator species [10]. The region displays highly variable physical structures, including eddies and filaments, that we characterize here by both Eulerian diagnostics (OW) and by Lagrangian methods (FSLE and RP). On the timescale of the predator’s trip, these structures appear and vanish. At the mesoscale, the time variability of the velocity field is evident irrespective of the method one chooses for looking at the structures generated by transport. Figure 8 illustrates the three aforementioned diagnostics at three different times during the elephant seal trip, two weeks apart from each other. The red dot in each panel indicates the position of the animal at the time when the diagnostic was computed. The tagged animal remained for more than two weeks close to a mesoscale eddy that is clearly evidenced both by the OW parameter (figures 8(a) and (b)) and the FSLEs (figures 8(d) and (e)). The submesoscale fronts generated by the temporal variability of the turbulent field are evidenced by the FSLEs, whereas the OW parameter only identifies the cores of the recirculation regions. In spite of the more accurate spatiotemporal resolution achievable by the use of Lagrangian diagnostics, that exploit at best the spatiotemporal variability of the velocity field, the FSLE 16 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al map appears to bear more information than one would need, and may not be the key prompt to understand the predator’s habitat use. Indeed, the case study reported here shows that even if the elephant seal seems to concentrate its foraging activity on a mesoscale eddy identifiable by the OW parameter, and to follow the submesoscale structures evidenced by the FSLEs, nevertheless not all eddies or all fronts are equivalent. For instance, figures 8(c), ( f ) and (i) display the animal crossing an eddy, revealed by both the OW parameter and by the FSLEs, without halting. We suggest here to use the RP introduced in section 2.2 as a tool to discriminate between eddies that may or may not be interesting for foraging. Indeed, figures 8(g)–(i) are consistent with the hypothesis that the elephant seal spends longer time in eddies with higher retention. The RP is of course not the only feature to be considered in evaluating the potential productivity of oceanic mesoscale eddies. Indeed, one primary factor affecting the biomass transfer to higher trophic levels is the origin of the water mass trapped within the eddy, and as a consequence the type of ecosystem it is able to sustain [21]. It is however notable that the elephant seal halts where the RP has high values, returning towards the highly retentive eddy after having moved towards a more permeable one, and then following the filament evident in figure 8(h). These features stand out clearly when looking at the retention time, but not in the FSLE and OW map. 5. Discussion and conclusions The application of dynamical systems theory to the two-dimensional velocity field of oceanic currents, obtained by reprocessing altimetric measures, revealed a powerful approach to identify and elucidate some key mechanisms involved in (sub)mesoscale circulation and environmental structuring. The finite-size Lyapunov exponent evidences the transport barriers that control the horizontal exchange of water in and out of eddy cores. However, the Lyapunov exponent does not provide information about the time that a water parcel has spent inside an eddy. This information is of critical relevance for the ecology of the open ocean, where patchiness in the distribution of the biomass along all the trophic chains is typically found. Here, we study the retention of oceanic mesoscale eddies by introducing a novel diagnostic, the retention parameter (RP), that estimates the time a water parcel has been recirculating inside an eddy core. Along the line of Lyapunov exponent computation, this diagnostic is based on the evaluation of a property of the velocity field, the Okubo–Weiss (OW) parameter, along the trajectory that emanates backwards in time from a given point. Negative values of the OW parameter are classically used to identify eddy cores. The retention time of an eddy is hence obtained by measuring along a particle trajectory the time of persistence of a negative OW value. The RP characterizes eddy cores, whereas the analysis based on Lyapunov exponents reveals a multitude of filaments, which in some cases enclose hotspots of retention. Eddy cores with retention times relevant for structuring higher trophic levels (of one month or larger) appear to be relatively rare: in the interfrontal region east of Kerguelen, they occupy about 3% of the ocean surface. If such regions have supported high primary production during the blooming season, it is reasonable to think that high-retention regions may support higher trophic levels as well. Since marine predator displacements are able to easily overcome the distance between eddies [8], the active search of highly retentive regions should be rewarding in particular when resources are rare and patchy (likely in post-bloom, oligotrophic conditions [18]). We have provided a preliminary confirmation of this hypothesis by considering the case study of a foraging trip 17 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al performed by an elephant seal in the Austral summer. This animal crossed several eddies, but showed a transition from straight movement to intensive searching only for an eddy with large retention time. If the horizontal current field had no temporal variability, then the closed orbits encircling mesoscale eddies would be material lines and therefore all eddy cores would have infinite retention times. The analysis of eddy retention that we have performed has led us naturally to a formalism by which the temporal variability of the velocity field, which is at the origin of finite retention time, can be introduced as a tunable perturbation over a time-invariant velocity field. This approach may be useful for performing perturbative analyses of eddy permeability on altimetry data and to better understand which terms in the temporal variability lead to the breaking of eddy cores, that are isolated in time-invariant fields. Approaches like Melnikov theory may be promising in this regard [63, 2]. Finally, here we focused on the impact of horizontal stirring in structuring the environment that sustains marine ecosystems. Several other physical processes however exist which control patchiness and dispersion of biotic tracers at various scales (e.g., [41]). A complete analysis of retention processes would need to include these other mechanisms together with the horizontal dynamics of eddy cores. Acknowledgments The altimeter products were produced by Ssalto/Duacs and distributed by Aviso with support from Cnes. Chlorophyll images were obtained by the GLOBCOLOUR project. ADP thanks the Fondation Bettencourt-Schueller (through the program Frontières du Vivant) and Fondazione CRT (through the program Marco Polo) for their financial support. The elephant seal work was conducted as part of the Institut Paul Emile Victor national research program (no. 109, H Weimerskirch and the observatory Mammifères Explorateurs du Milieu Océanique, MEMO SOERE CTD 02). This work was carried out within the framework of the ANR Blanc MYCTO-3D-MAP and ANR VMC 07: IPSOS-SEAL programs, OSTST program ALTIMECO and CNES-TOSCA program (Élephants de mer océanographes). CG also thanks the Total Foundation for financial support. References [1] Abraham E R and Bowen M M 2002 Chaotic stirring by a mesoscale surface-ocean flow Chaos 12 373–81 [2] Aharon R, Rom-Kedar V and Gildor H 2012 When complexity leads to simplicity: ocean surface mixing simplified by vertical convection Phys. Fluids 24 056603–13 [3] Artale V, Boffetta G, Celani A, Cencini M and Vulpiani A 1997 Dispersion of passive tracers in closed basins: beyond the diffusion coefficient Phys. Fluids 9 3162 [4] Aurell E, Boffetta G, Crisanti A, Paladin G and Vulpiani A 1997 Predictability in the large: an extension of the concept of Lyapunov exponent J. Phys. A: Math. Gen. 30 1 [5] Bailleul F, Authier M, Ducatez S, Roquet F, Charrassin J-B, Cherel Y and Guinet C 2010 Looking at the unseen: combining animal bio-logging and stable isotopes to reveal a shift in the ecological niche of a deep diving predator Ecography 33 709–19 [6] Benitez-Nelson C R and McGillicuddy D J Jr 2008 Mesoscale physical–biological–biogeochemical linkages in the open ocean: an introduction to the results of the e-flux and EDDIES programs Deep-Sea Res. II 55 1133–8 [7] Beron-Vera F J, Olascoaga M J and Goni G J 2008 Oceanic mesoscale eddies as revealed by Lagrangian coherent structures Geophys. Res. Lett. 35 L12603 [8] Block B A et al 2011 Tracking apex marine predator movements in a dynamic ocean Nature 475 86–90 [9] Boffetta G, Lacorata G, Redaelli G and Vulpiani A 2001 Detecting barriers to transport: a review of different techniques Physica D 159 58–70 18 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [10] Bost C, Cotté C, Bailleul F, Cherel Y, Charrassin J, Guinet C, Ainley D and Weimerskirch H 2009 The importance of oceanographic fronts to marine birds and mammals of the southern oceans J. Mar. Syst. 78 363–76 [11] Bracco A, Provenzale A and Scheuring I 2000 Mesoscale vortices and the paradox of the plankton Proc. R. Soc. B: Biol. Sci. 267 1795–800 [12] Buesseler K O 2012 Biogeochemistry: the great iron dump Nature 487 305–6 [13] Calil P H R and Richards K J 2010 Transient upwelling hot spots in the oligotrophic north pacific J. Geophys. Res. 115 C02003 [14] Camphuysen C J 2006 Top Predators in Marine Ecosystems: Their Role in Monitoring and Management (Cambridge: Cambridge University Press) [15] Chelton D B, Gaube P, Schlax M G, Early J J and Samelson R M 2011 The influence of nonlinear mesoscale eddies on near-surface oceanic chlorophyll Science 334 328–32 [16] Chelton D B, Schlax M G and Samelson R M 2011 Global observations of nonlinear mesoscale eddies Prog. Oceanogr. 91 167–216 [17] Chelton D B, Schlax M G, Samelson R M and Szoeke R A d 2007 Global observations of large oceanic eddies Geophys. Res. Lett. 34 L15606 [18] Cotté C, d’Ovidio F, Chaigneau A, Lévy M, Taupier-Letage I, Mate B and Guinet C 2011 Scale-dependent interactions of Mediterranean whales with marine dynamics Limnol. Oceanogr. 106 219–32 [19] De Monte S, d’Ovidio F, Cotté C, Lévy M, Le Corre M and Weimerskirch 2012 Frigatebird behaviour at the ocean–atmosphere interface: integrating animal behaviour with multisatellite data J. R. Soc. Interface 9 3351–8 [20] Desprès A, Reverdin G and d’Ovidio F 2011 Mechanisms and spatial variability of mesoscale frontogenesis in the northwestern subpolar gyre Ocean Modelling 39 97–113 [21] d’Ovidio F, De Monte S, Alvain S, Dandonneau Y and Lévy M 2010 Fluid dynamical niches of phytoplankton types Proc. Natl Acad. Sci. USA 107 18366–70 [22] d’Ovidio F, Fernández V, Hernández-Garcı́a E and López C 2004 Mixing structures in the Mediterranean Sea from finite-size Lyapunov exponents Geophys. Res. Lett. 31 L17203 [23] d’Ovidio F, Isern-Fontanet J, López C, Hernández-Garcı́a E and Garcı́a-Ladona E 2009 Comparison between Eulerian diagnostics and finite-size Lyapunov exponents computed from altimetry in the Algerian basin Deep-Sea Res. I 56 15–31 [24] d’Ovidio F, Shuckburgh E and Legras B 2009 Local mixing events in the upper troposphere and lower stratosphere. Part I: detection with the Lyapunov diffusivity J. Atmos. Sci. 66 3678–94 [25] Dragon A C, Bar-Hen A, Monestiez P and Guinet C 2012 Horizontal and vertical movements to predict foraging success in a marine predator Mar. Ecol. Progr. Ser. 447 243–57 [26] Fauchald P, Erikstad K E and Skarsfjord H 2000 Scale-dependent predator–prey interactions: the hierarchical spatial distribution of seabirds and prey Ecology 81 773–83 [27] Haller G 2001 Lagrangian structures and the rate of strain in a partition of two-dimensional turbulence Phys. Fluids 13 3365–85 [28] AVISO 2013 Ssalto/Duacs User Handbook: M(SLA) and M(ADT) Near-Real Time and Delayed-Time Products, edition 3.4 [29] Hernández-Carrasco I, López C, Hernández-Garcı́a E and Turiel A 2011 How reliable are finite-size Lyapunov exponents for the assessment of ocean dynamics? Ocean Modelling 36 208–18 [30] Isern-Fontanet J, Garcı́a-Ladona E and Font J 2003 Identification of marine eddies from altimetric maps J. Atmos. Ocean. Technol. 20 772–8 [31] Kai E T, Rossi V, Sudre J, Weimerskirch H, Lopez C, Hernandez-Garcia E, Marsac F and Garcon V 2009 Top marine predators track Lagrangian coherent structures Proc. Natl Acad. Sci. USA 106 8245–50 [32] Labat J-P, Gasparini S, Mousseau L, Prieur L, Boutoute M and Mayzaud P 2009 Mesoscale distribution of zooplankton biomass in the Northeast Atlantic Ocean determined with an optical plankton counter: relationships with environmental structures Deep-Sea Res. I 56 1742–56 [33] Lacorata G, Purini R, Vulpiani A and Zambianchi E 1996 Dispersion of passive tracers in model flows: effects of the parametrization of small-scale processes Ann. Geophys. 14 476–84 [34] Lapeyre G, Hua B L and Legras B 2000 Comment on ‘Finding finite-time invariant manifolds in two-dimensional velocity fields’ Chaos 10 99 Lapeyre G, Hua B L and Legras B 2001 Chaos 11 427–30 [35] Lapeyre G, Klein P and Hua B L 1999 Does the tracer gradient vector align with the strain eigenvectors in 2D turbulence? Phys. Fluids 11 3729–37 [36] Le Traon P Y, Nadal F and Ducet N 1998 An improved mapping method of multisatellite altimeter data J. Atmos. Ocean. Technol. 15 522–34 [37] Legras B and Vautard R 1996 A guide to Lyapunov vectors: Proc. ECMWF Seminar on Predictability pp 143–56 19 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [38] Lehahn Y, d’Ovidio F, Lévy M, Amitai Y and Heifetz E 2011 Long range transport of a quasi isolated chlorophyll patch by an Agulhas ring Geophys. Res. Lett. 38 L16610 [39] Lehahn Y, d’Ovidio F, Lévy M and Heifetz E 2007 Stirring of the northeast Atlantic spring bloom: a Lagrangian analysis based on multisatellite data J. Geophys. Res. 112 15 [40] Lévy M 2008 The modulation of biological production by oceanic mesoscale turbulence Transport and Mixing in Geophysical Flows (Lecture Notes in Physics vol 744) ed J Weiss and A Provenzale (Berlin: Springer) pp 219–61 [41] Lévy M, Ferrari R, Franks P J S, Martin A P and Rivière P 2012 Bringing physics to life at the submesoscale Geophys. Res. Lett. 39 L14602 [42] Mahadevan A and Campbell J W 2002 Biogeochemical patchiness at the sea surface Geophys. Res. Lett. 29 4 [43] Mancho A M, Small D and Wiggins S 2006 A tutorial on dynamical systems concepts applied to Lagrangian transport in oceanic flows defined as finite time data sets: theoretical and computational issues Phys. Rep. 437 55–124 [44] Martin A 2003 Phytoplankton patchiness: the role of lateral stirring and mixing Prog. Oceanogr. 57 125–74 [45] McGillicuddy D J et al 2007 Eddy/wind interactions stimulate extraordinary mid-ocean plankton blooms Science 316 1021–6 [46] Miramontes O, Boyer D and Bartumeus F 2012 The effects of spatially heterogeneous prey distributions on detection patterns in foraging seabirds PLoS ONE 7 e34317 [47] Morrow R, Birol F, Griffin D and Sudre J 2004 Divergent pathways of cyclonic and anti-cyclonic ocean eddies Geophys. Res. Lett. 31 L24311 [48] Nencioli F, d’Ovidio F, Doglioli A M and Petrenko A A 2011 Surface coastal circulation patterns by in situ detection of Lagrangian coherent structures Geophys. Res. Lett. 38 L17604 [49] Neufeld Z, López C, Hernández-Garcı́a E and Tél T 2000 Multifractal structure of chaotically advected chemical fields Phys. Rev. E 61 3857–66 [50] Noack B R, Mezić I, Tadmor G and Banaszuk A 2004 Optimal mixing in recirculation zones Phys. Fluids 16 867–88 [51] Oseledec V 1968 Multiplicative ergodic theorem: characteristic Lyapunov exponents of dynamical systems Trans. Moscow Math. Soc. 19 197–231 [52] Ott E 2002 Chaos in Dynamical Systems 2nd edn (Cambridge: Cambridge University Press) [53] Pasquero C, Bracco A, Provenzale A and Weiss J 2007 Particle motion in a sea of eddies Lagrangian Analysis and Prediction of Coastal and Ocean Dynamics (Cambridge: Cambridge University Press) [54] Patterson T A, Thomas L, Wilcox C, Ovaskainen O and Matthiopoulos J 2008 State–space models of individual animal movement Trends Ecol. Evol. 23 87–94 [55] Perruche C, Rivière P, Lapeyre G, Carton X and Pondaven P 2011 Effects of surface quasi-geostrophic turbulence on phytoplankton competition and coexistence J. Mar. Res. 69 105–35 [56] Pierrehumbert R T and Yang H 1993 Global chaotic mixing on isentropic surfaces J. Atmos. Sci. 50 2462–80 [57] Pérez-Muñuzuri V and Huhn F 2010 The role of mesoscale eddies time and length scales on phytoplankton production Nonlinear Process. Geophys. 17 177–86 [58] Resplandy L, Vialard J, Lévy M, Aumont O and Dandonneau Y 2009 Seasonal and intraseasonal biogeochemical variability in the thermocline ridge of the southern tropical Indian Ocean J. Geophys. Res. 114 C07024 [59] Rossi V, López C, Hernández-Garcı́a E, Sudre J, Garçon V and Morel Y 2009 Surface mixing and biological activity in the four eastern boundary upwelling systems Nonlinear Process. Geophys. 16 557–68 [60] Rossi V, López C, Sudre J, Hernández-Garcı́a E and Garçon V 2008 Comparative study of mixing and biological activity of the Benguela and Canary upwelling systems Geophys. Res. Lett. 35 L11602 [61] Sabarros P S, Mnard F, Lvnez J, TewKai E and Ternon J 2009 Mesoscale eddies influence distribution and aggregation patterns of micronekton in the Mozambique Channel Mar. Ecology Prog. Ser. 395 101–7 [62] Samelson R 2013 Lagrangian motion, coherent structures, and lines of persistent material strain Annu. Rev. Mar. Sci. 5 137–63 [63] Samelson R M 1992 Fluid exchange across a meandering jet J. Phys. Oceanogr. 22 431–44 [64] Sandulescu M, López C, Hernández-Garcı́a E and Feudel U 2007 Plankton blooms in vortices: the role of biological and hydrodynamic timescales Nonlinear Process. Geophys. 14 443–54 [65] Santora J A, Sydeman W J, Schroeder I D, Wells B K and Field J C 2011 Mesoscale structure and oceanographic determinants of krill hotspots in the California Current: implications for trophic transfer and conservation Prog. Oceanogr. 91 397–409 [66] Schneider J and Tél T 2003 Extracting flow structures from tracer data Ocean Dyn. 53 64–72 [67] Schroeder K, Haza A C, Griffa A, Ozgoekmen T M, Poulain P M, Gerin R, Peggion G and Rixen M 2011 Relative dispersion in the Liguro-Provencal basin: from sub-mesoscale to mesoscale Deep-Sea Res. I 58 209–28 [68] Sims D W and Quayle V A 1998 Selective foraging behaviour of basking sharks on zooplankton in a small-scale front Nature 393 460–4 20 J. Phys. A: Math. Theor. 46 (2013) 254023 F d’Ovidio et al [69] Smetacek V et al 2012 Deep carbon export from a southern ocean iron-fertilized diatom bloom Nature 487 313–9 [70] Waugh D W and Abraham E R 2008 Stirring in the global surface ocean Geophys. Res. Lett. 35 L20605 [71] Waugh D W, Abraham E R and Bowen M M 2006 Spatial variations of stirring in the surface ocean: a case study of the Tasman sea J. Phys. Oceanogr. 36 526–42 [72] Weimerskirch H, Corre M L, Jaquemet S, Potier M and Marsac F 2004 Foraging strategy of a top predator in tropical waters: great frigatebirds in the Mozambique Channel Mar. Ecology Prog. Ser. 275 297–308 [73] Weiss J B and Provenzale A 2008 Transport And Mixing In Geophysical Flows (Berlin: Springer) [74] Wiggins S 2005 The dynamical systems approach to Lagrangian transport in oceanic flows Annu. Rev. Fluid Mech. 37 295–328 21
© Copyright 2026 Paperzz