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Fractals have been applied to describe the complexity of behavioral displays in a range of organisms. Recent work suggests that they may represent a promising tool in the quantification of subtle behavioral responses in marine mammals under chronic exposure to disturbance. This paper aims at introducing the still seldom used fractals to the broader community of marine mammal scientists. We first briefly rehearse some of the fundamental principles behind fractal theory and review the previous uses of fractals in marine mammal science. We subsequently introduce two methods that may be used to assess the complexity of marine mammal diving patterns, and we apply them to the temporal dynamics of the diving patterns of killer whales in the presence and absence of sea kayaks, the sequential behavior of harbor and gray seals in environments with distinct levels of anthropogenic influence, and southern right whales with and without calves. We discuss the ecological relevance of identifying fractal properties in marine mammal behavior, and the potential strength of the fractal behavioral parameters in comparison to more standard behavioral metrics. We finally briefly address the relevance fractal methods may have for the design and implementation of management and conservation strategies. © 2017 Society for Marine Mammalogy.


Klaassen W.,University of Groningen | Spilmont N.,University of Lille Nord de France | Spilmont N.,CNRS Laboratory of Oceanology and Geosciences
Estuarine, Coastal and Shelf Science | Year: 2012

Carbon dioxide (CO 2) exchanges between a tidal flat (Wadden Sea, Netherlands) and the atmosphere were measured during a three-year survey. CO 2 exchanges were monitored during 1-2 days each month between September 2006 and September 2009 using a flux chamber. The flux of CO 2 was separated into two fluxes: the dark flux and the gross light flux, with the dark flux representing the flux during darkness and the gross light flux the difference between the net CO 2 flux (measured under light) and the dark flux. It was argued that dark and light fluxes may deviate from respiration and photosynthesis, as the fluxes between wet tidal sediment and the atmosphere are affected by the partial pressure of CO 2 in pore water, which is only gradually changed by sources and sinks of CO 2 in the sediment. Light and dark fluxes were empirically related to environmental parameters in order to interpolate between succeeding measurements. The dark flux appeared to increase with temperature and the light flux became more intense with increasing irradiance with signs of saturation at high light levels on many but not all measurement days. These relations with environmental parameters showed seasonal and inter-annual variability. Fluxes were negligible just after the site was emersed and it took up to 3 h of emersion until fluxes were adapted to environmental conditions. Dark and light fluxes both showed strong seasonality with high values in summer and low values in winter. The tidal flat appeared as a source of atmospheric CO 2 in the first year of measurement (+0.35 mol CO 2 m -2 yr -1) and a sink in the following two years (-1.59 and -0.72 mol CO 2 m -2 yr -1). Since the source of CO 2 was observed during an extremely warm year, we suggest that climate warming might influence the carbon budget of tidal flats. © 2011 Elsevier Ltd.


Edwards M.,Sir Alister Hardy Foundation for Ocean Science | Edwards M.,University of Plymouth | Beaugrand G.,CNRS Laboratory of Oceanology and Geosciences | Hays G.C.,University of Swansea | And 3 more authors.
Trends in Ecology and Evolution | Year: 2010

Long-term biological time-series in the oceans are relatively rare. Using the two longest of these we show how the information value of such ecological time-series increases through space and time in terms of their potential policy value. We also explore the co-evolution of these oceanic biological time-series with changing marine management drivers. Lessons learnt from reviewing these sequences of observations provide valuable context for the continuation of existing time-series and perspective for the initiation of new time-series in response to rapid global change. Concluding sections call for a more integrated approach to marine observation systems and highlight the future role of ocean observations in adaptive marine management. © 2010 Elsevier Ltd.


Frederiksen M.,University of Aarhus | Frederiksen M.,UK Center for Ecology and Hydrology | Anker-Nilssen T.,Norwegian Institute for Nature Research | Beaugrand G.,CNRS Laboratory of Oceanology and Geosciences | Wanless S.,UK Center for Ecology and Hydrology
Global Change Biology | Year: 2013

The boreal Northeast Atlantic is strongly affected by current climate change, and large shifts in abundance and distribution of many organisms have been observed, including the dominant copepod Calanus finmarchicus, which supports the grazing food web and thus many fish populations. At the same time, large-scale declines have been observed in many piscivorous seabirds, which depend on abundant small pelagic fish. Here, we combine predictions from a niche model of C. finmarchicus with long-term data on seabird breeding success to link trophic levels. The niche model shows that environmental suitability for C. finmarchicus has declined in southern areas with large breeding seabird populations (e.g. the North Sea), and predicts that this decline is likely to spread northwards during the 21st century to affect populations in Iceland and the Faroes. In a North Sea colony, breeding success of three common piscivorous seabird species [black-legged kittiwake (Rissa tridactyla), common guillemot (Uria aalge) and Atlantic puffin (Fratercula arctica)] was strongly positively correlated with local environmental suitability for C. finmarchicus, whereas this was not the case at a more northerly colony in west Norway. Large seabird populations seem only to occur where C. finmarchicus is abundant, and northward distributional shifts of common boreal seabirds are therefore expected over the coming decades. Whether or not population size can be maintained depends on the dispersal ability and inclination of these colonial breeders, and on the carrying capacity of more northerly areas in a warmer climate. © 2012 Blackwell Publishing Ltd.


Reygondeau G.,IRD Montpellier | Beaugrand G.,CNRS Laboratory of Oceanology and Geosciences
Global Change Biology | Year: 2011

Calanus finmarchicus is a key-structural species of the North Atlantic polar biome. The species plays an important trophic role in subpolar and polar ecosystems as a grazer of phytoplankton and as a prey for higher trophic levels such as the larval stages of many fish species. Here, we used a recently developed ecological niche model to assess the ecological niche (sensu Hutchinson) of C. finmarchicus and characterize its spatial distribution. This model explained about 65% of the total variance of the observed spatial distribution inferred from an independent dataset (data of the continuous plankton recorder survey). Comparisons with other types of models (structured population and ecophysiological models) revealed a clear similarity between modeled spatial distributions at the scale of the North Atlantic. Contemporary models coupled with future projections indicated a progressive reduction of the spatial habitat of the species at the southern edge and a more pronounced one in the Georges Bank, the Scotian Shelf and the North Sea and a potential increase in abundance at the northern edge of its spatial distribution, especially in the Barents Sea. These major changes will probably lead to a major alteration of the trophodynamics of North Atlantic ecosystems affecting the trophodynamics and the biological carbon pump. © 2010 Blackwell Publishing Ltd.


Helaouet P.,Sir Alister Hardy Foundation for Ocean Science | Helaouet P.,CNRS Laboratory of Oceanology and Geosciences | Beaugrand G.,CNRS Laboratory of Oceanology and Geosciences | Reid P.C.,Sir Alister Hardy Foundation for Ocean Science
Progress in Oceanography | Year: 2011

Copepods represent the major part of the dry weight of the mesozooplankton in pelagic ecosystems and therefore have a central role in the secondary production of the North Atlantic Ocean. The calanoid copepod species Calanus finmarchicus is the main large copepod in subarctic waters of the North Atlantic, dominating the dry weight of the mesozooplankton in regions such as the northern North Sea and the Norwegian Sea. The objective of this work was to investigate the relationships between both the fundamental and realised niches of C. finmarchicus in order to better understand the future influence of global climate change on the abundance, the spatial distribution and the phenology of this key-structural species. Based on standardised Principal Component Analyses (PCAs), a macroecological approach was applied to determine factors affecting the spatial distribution of C. finmarchicus and to characterise its realised niche. Second, an ecophysiological model was used to calculate the Potential Egg Production Rate (PEPR) of C. finmarchicus and the centre of its fundamental niche. Relationships between the two niches were then investigated by correlation analysis. We found a close relationship between the fundamental and realised niches of C. finmarchicus at spatial, monthly and decadal scales. While the species is at the centre of its niche in the subarctic gyre, our joint macroecological and macrophysiological analyses show that it is at the edge of its niche in the North Sea, making the species in this region more vulnerable to temperature changes. © 2010 Elsevier Ltd.


Chapperon C.,Flinders University | Seuront L.,Flinders University | Seuront L.,South Australian Research And Development Institute | Seuront L.,CNRS Laboratory of Oceanology and Geosciences
Functional Ecology | Year: 2011

1.Behavioural adaptations of ectotherms to thermally heterogeneous environments are still overlooked in the literature despite the fact that organismal behaviour could enhance survival in the warming world. This is particularly critical in the intertidal where most ectotherms live at, or near to the upper limit of thermal tolerance. 2.This study investigated (i) the environmental factors determining the body temperatures of the intertidal gastropod Nerita atramentosa, (ii) the space-time variability in environmental and individual body temperatures and (iii) the potential variability in N. atramentosa thermoregulatory behaviours, i.e. microhabitat selection and aggregation. 3.Thermal imaging was used to assess the body temperatures of N. atramentosa and surrounding substrata over two seasons (autumn and summer), at two shore levels (low- vs. high-shore levels) within two habitats of different topographic complexity (rock platform and boulders) on the same rocky shore. 4.Snail body and substratum temperatures were significantly and positively correlated within each habitat at both seasons. Substratum temperature may thus be considered as a primary driver of body temperatures of organisms that attach to a substratum. Substratum temperature and other variables such as solar irradiance critically need to be integrated in climate-change models that use single climatic variables (e.g. air temperature) that are not necessarily correlated with individual body temperatures in nature. 5.The high space-time variability in both substratum and body temperatures reinforces the growing evidence that small spatial scale variations may surpass those observed at larger spatial scales. 6.Nerita atramentosa thermoregulatory behaviour under high thermal stress appeared to be habitat specific. 7.The small spatial scale heterogeneity in environmental and individual temperatures and in thermoregulatory behaviours has stressed the need to focus on body temperature patterns at the niche level and to integrate the organismal behaviour in climate-change models. © 2011 The Authors. Functional Ecology © 2011 British Ecological Society.


Since 1967, the alumina plants in the Marseilles area (Barasse and Gardanne) have been discharging the mineral residue (i.e., red mud) resulting from the alkaline processing of bauxite into the submarine Cassidaigne canyon (north-western Mediterranean Sea) through pipes situated at 320-330 m in depth. The Barasse pipe stopped being used in 1988. From 1987 to 1996, many decrees and regulations were promulgated by the French State to rule the conditions under which the Gardanne alumina refinery was authorized to dispose of the bauxite residue in the sea. The refinery was required: (i) to study the hydrodynamic circulation in the Cassidaigne canyon to evaluate the potential dispersion and transport of fine elements discharged into the water mass and their impact on the pelagic ecosystem; (ii) to survey the marine environment every five years to control the expansion and thickness of the red mud deposit and compare the evolution of the benthic macrofauna at representative sampling sites in the environment affected by the red mud discharge with that of reference sites outside of the red mud plume; (iii) to study the effect of the discharge on fishing activities; and (iv) to investigate the toxicity of the red mud, particularly its persistence, accumulation, interaction and effect on the marine ecosystem, paying special attention to the bio-accumulation of chromium and vanadium. A Scientific Committee was created to insure an independent evaluation of the studies promised by the manufacturer in response to the State's regulations. Since the beginning of the 1960s, data have been accumulating on the structure and long-term functioning of the Cassidaigne bathyal ecosystem. This paper presents the collaborative efforts of the State-Manufacturer-Committee triplet and summarizes the main results obtained during the last period's sea campaigns (1991-2007). This paper also illustrates how national regulations concerning manufacturers, such as Gardanne alumina refinery, have provided new knowledge about the structure and functioning of a bathyal ecosystem in the Cassidaigne canyon in the north-western Mediterranean Sea. © 2009 Elsevier Ltd.


Seuront L.,CNRS Laboratory of Oceanology and Geosciences
Physica A: Statistical Mechanics and its Applications | Year: 2015

Fractal analysis is increasingly used to describe, and provide further understanding to, zooplankton swimming behavior. This may be related to the fact that fractal analysis and the related fractal dimension D have the desirable properties to be independent of measurement scale and to be very sensitive to even subtle behavioral changes that may be undetectable to other behavioral variables. As early claimed by Coughlin et al. (1992), this creates "the need for fractal analysis" in behavioral studies, which has hence the potential to become a valuable tool in zooplankton behavioral ecology. However, this paper stresses that fractal analysis, as well as the more elaborated multifractal analysis, is also a risky business that may lead to irrelevant results, without paying extreme attention to a series of both conceptual and practical steps that are all likely to bias the results of any analysis. These biases are reviewed and exemplified on the basis of the published literature, and remedial procedures are provided not only for geometric and stochastic fractal analyses, but also for the more complicated multifractal analysis. The concept of multifractals is finally introduced as a direct, objective and quantitative tool to identify models of motion behavior, such as Brownian motion, fractional Brownian motion, ballistic motion, Lévy flight/walk and multifractal random walk. I finally briefly review the state of this emerging field in zooplankton behavioral research. © 2015 Elsevier B.V.


Beaugrand G.,CNRS Laboratory of Oceanology and Geosciences
Marine Ecology Progress Series | Year: 2012

Evidence of global warming is now unequivocal, and studies suggest that it has started to influence natural systems of the planet, including the oceans. However, in the marine environment, it is well-known that species and ecosystems can also be influenced by natural sources of large-scale hydro-climatological variability. The North Atlantic Oscillation (NAO) was negatively correlated with the mean abundance of one of the subarctic key species Calanus fin- marchicus in the North Sea. This correlation was thought to have broken down in 1996, however, the timing has never been tested statistically. The present study revisits this unanticipated change and reveals that the correlation did not break down in 1996 as originally proposed but earlier, at the time of an abrupt ecosystem shift in the North Sea in the 1980s. Furthermore, the analyses demonstrate that the correlation between the NAO and C. finmarchicus abundance is modulated by the thermal regime of the North Sea, which in turn covaries positively with global temperature anomalies. This study thereby provides evidence that global climate change is likely to alter some empirical relationships found in the past between species abundance or the ecosystem state and large-scale natural sources of hydro-climatological variability. A theory is proposed to explain how this might happen. These unanticipated changes, also called 'surprises' in climatic research, are a direct consequence of the complexity of both climatic and biological systems. In this period of rapid climate change, it is therefore hazardous to integrate meteo-oceanic indices such as the NAO in models used in the management of living resources, as it has been sometimes attempted in the past. © Inter-Research 2012.

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