224 Fractals and Multifractals in Ecology and Aquatic Science
biomass distributions characterized by their high-order stochasticity, while in low hydrody-
namic conditions, as those encountered in the stable waters of the Ligurian Sea, phytoplankton
distribution could be rather characterized by a low-order deterministic behavior.
Although our results suggest that temperature, salinity, and phytoplankton biomass exhibit a
higher dimensionality in high hydrodynamic conditions, we cannot conclude, on the basis of the
three previously used analysis techniques, the existence of low-order deterministic chaos, but only
to a lower dimensionality in low hydrodynamic conditions.
6.1.3.4 chaos, attractors, and Fractals
Fractals can be temporal, spatial, or phase-space manifestations of chaos in nonlinear dynamic sys-
tems. Fractals in phase-space can either be attractors themselves—that is, strange attractors, such
as the bifurcation diagram of the logistic equation or the Hénon map—or they can constitute the
dividing line between separate attractor basins in phase-space (see, for example, Peitgen and Saupe
1988; Peitgen et al. 1992). The study of attractors is important because the geometry of an attractor
frequently captures much of the underlying dynamics and allows one-dimensional (fractal) descrip-
tion. We will nevertheless see hereafter that the geometry of strange attractors can be so complex
that it becomes impossible to describe them in terms of fractal dimensions and (low-order) deter-
ministic chaos. In particular, this statement precludes the introduction of the multifractal, high-
order stochastic framework.
6.1.4 ch a o s i n Ec o l o g i c a l sc i E n c E s
Since the seminal studies of chaos in discrete time models in population ecology (May 1974, 1975,
1976), the issue of chaotic dynamics in ecological systems has been widely controversial (Hassell
et al. 1976; Berryman and Millstein 1989; Pool 1989). Chaos in ecology has nevertheless been the
subject of an increasing amount of literature. In theoretical ecology, there are many examples of
temporal population models that exhibit chaos. The interaction of three variables in a predator–prey
nutrient system (Kot et al. 1992) is now a well-studied chaotic system, as chaotic dynamics expected
through a trophic coupling of three species (Hastings and Powell 1991). Recently, an ocean ecosys-
tem model also exhibited chaotic properties related to external seasonal forcing (Popova et al. 1997).
In particular, the issues raised by chaos theory in ecology have been the subject of several reviews
(May 1980, 1987; Godfrey and Blythe 1991; Ellner 1992; Logan and Allen 1992; Hastings et al.
1993; Little et al. 1996).
As briey suggested in the above section, the compelling reasons for the emerging chaos theory
to ecology are based on the hope that complex systems could be explained by relatively low-order
processes. This leads to the development of a suite of algorithms aimed at the detection of chaotic
behavior and the classication of system dynamics; see, for example, Hastings et al. (1993) and Ellner
and Turchin (1995) for reviews. While such approaches have been applied to a wide variety of time
series (Farmer and Sidorowich 1987; Ellner 1992; Theiler et al. 1992) to detect dynamic spatial chaos
(Rubin 1992; Rand 1994; Solé and Bascompte 1995), the development of nonlinear thinking to aquatic
ecology has a more recent history. Only a few studies have been devoted to detecting chaotic signature
in both marine time series and transects, and led to controversial results. Sugihara and May (1990b)
found evidence for chaotic dynamics in time series of weekly diatom counts, and Scheffer (1991)
argued that chaotic deterministic dynamics should be commonplace in plankton communities. Ascioti
et al. (1993), Strutton et al. (1996, 1997) and Seuront (2004), however, did not nd any evidence of
chaotic dynamics in both zooplankton and phytoplankton time series, phytoplankton transects, and
temperature, salinity, and in vivo uorescence time series, respectively. Ascioti et al. (1993) found a
signicant level of predictability of zooplankton abundance from that of phytoplankton, indicative of
a deterministic trophic link. A recent application of the nearest-neighbor algorithm to time series of a
range of physical and biological variables for the north Pacic Ocean (Hsieh et al. 2005) showed that
physical variables were characterized by a high dimensionality (E bounded between 13 and 20) and
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