The Mysterious Taylor’s Power Law – Part I

Interesting statistical properties of population dispersion emerge from the simulations of the Zoomer model, which represents the population level variant of my Multi-scaled random walk model (MRW). The Zoomer model seems to offer a potential to cast new light on a particular scaling pattern, Taylor’s power law (Taylor 1961, 1986), which has been a nagging stone in the shoe for population ecology for more than 50 years. Taylor’s power law is one of the most widely tested empirical patterns in ecology and is the subject of an estimated thousand papers (Eisler et al. 2008)!

Despite this effort, a consensus to explain it has still not been obtained (Kendal and Jørgensen 2011). The scale-free pattern has been observed far beyond animal populations:

Taylor’s law is remarkable in that it is evident over the scale of a single chromosome (Kendal 2003, 2004) to the lungs of mice (Kendal and frost 1987), a farmer’s field (Kendal 2002), and upward to the breadth of the British Isles (Taylor and Taylor 1977). How the macroscopic fluctuations that characterize many of these observations might arise as a consequence of thermodynamic mechanisms is an intriguing question.
W. S. Kendal and B. Jørgensen, 2011.

In an ecological context, Taylor’s power law describes how variance (V) of population abundance (M) in a time series or over a spatial transect tends to comply empirically with V(M)=aMb, with b>>1 (in real population data b is typically close to 2, but sometimes even larger). In a spatial transect, the general mathematical modelling framework for population dispersion at fine scales predicts b≈1, which complies with a Poisson (“classically random”) distribution of individuals. In this framework b≈2 requires population sampling at relatively large scales.  At these coarser scales local variability in growth rate may trivially lead to high degree of heterogeneous dispersion due to larger degree of local growth independence between neighbourhood sites at these scales (environmental stochasticity). 

However, there are many empirical examples where population dispersion with b=2 has been observed also at fine spatial scales, in conflict with key model assumptions for local population mixing whether the environment is homogeneous or heterogeneous. In my book I describe these paradoxes in greater detail.


Figure 41 in my book: Illustration of the Z-paradox in a diagram of logarithmically transformed sample variance (log(V)) as a function of logarithmically transformed mean number of animals (log(M)) pr. sample unit at a given grid resolution. Sufficient population mixing to satisfy ergodicity is assumed. M is then changed by changing grid resolution. For a random distribution of individuals in a homogeneous environment, log(V) = log(M) (slope =1 in the diagram, where the slope is represented by the exponent b in Taylor’s power law). In a heterogeneous environment variance for a given M is then typicslly larger (for example, of magnitude described by the parallel shift from A to D in the illustration). This is expressed by the parameter a in Taylor’s power law (see main text), but b is unchanged upon such rescaling. Thus, by changing grain size for sample units in this manner, variance is still expected to change proportionally with this size, since M changes proportionally (say, from A to B or from D to C, or vice versa). If successive neighbourhood grid cells are not spatially correlated with respect to M, the line that is labelled “slope = 2” represents the Z-paradox, since variance is changing non-proportionally from A to C (or C to A) rather than the expected A to B (or C to D) under change of grid resolution.

In addition to the b≈2 issue even towards finer spatial resolutions, one aspect of Taylor’s power law has by and large been ignored: first, the V(M) pattern unexpectedly emerges in a consistent manner with respect to constancy of b also when abundance is measured at different spatial resolutions within a given spatial extent (Taylor 1986). Larger spatial units (grain size) lead to proportionally larger M on average. Second, and even more mysteriously, in my book I show by simulations and by empirical example that the respective “grains” do not need to be located side-by-side when lumped together to form larger scale analyses. Thus, appearance of the scaling law with b≈2 even in spatially non-auto-correlated data appears even more paradoxical in the context of constant b upon rescaling.

In my book I describe this inter-scale constancy of b≈2 (spatial self-similarity) as the “Z-paradox”, and propose – with support from simulations using the zoomer model – that a parallel-processing kind of population kinetics may contribute to solve it. I also provide a pilot test on sycamore aphids Drepanosiphum platanoides, which shows compliance with predictions from the Zoomer model and thereby also with basic MRW model properties.

In some follow-up posts I will illustrate more aspects of this remarkable coherence between parallel processing-based population dynamics and single-individual dispersion of relocations (e.g., GPS series). I will also connect further dots from this novel framework for animal space use towards a proposal for a general theory of complex systems from a statistical-mechanical perspective.


Kendal, W. S. 2003. Mol. Biol. Evol. 20: 579
Kendal, W. S. 2003. 2004. BMC Evol. Biol. 4:3
Kendal, W. S. 2002. Ecol. Model. 151: 261
Kendal, W. S. and P. Frost. 1987. J. Natl. Cancer Inst. 79, 1113
Kendal, W. S. and B. Jørgensen, 2011. Physical Review E 83, 066115
Taylor, L. R. 1961. Aggregation, variance and the mean. Nature 189:732-735.
Taylor, L. R. and R. A. Taylor 1977. Nature (London) 265, 415 (1977)
Taylor, L. R. 1986. Synoptic dynamics, migration and the Rothamsted insect survey. J. Anim. Ecol. 55:1-38.


A Statistical-Mechanical Perspective on Site Fidelity – Part VI

In Part III of this group of posts I described how to estimate an animal’s characteristic scale of space use, CSSU, by zooming over a range of pixel resolutions until “focus” is achieved according to the Home range ghost function I(N)=c√N. This pixel size regards the balancing point where observed inward contraction of entropy equals outward expansion of entropy, as sample size of fixes N is changed. If the power exponent satisfies 0.5 [i.e., square root expansion of I(N)], the MRW theory states that the animal – in statistical terms – had put equal weight into habitat utilization over the actual scale range of space use. In Part V I explained two new statistical-mechanical concepts, micro-and macrostates of a system, but in the context of the classical framework. In this follow-up post I’m approaching the behaviour of these key properties under the condition of complex space use, both at the “balancing scale” CSSU and by zooming over its surrounding range of scales. In other words, while the CSSU property from an I(N) regression was studied with respect to observational intensity, N, at a given spatial scale, I here describe the system at different spatial scales at a given N.

First, a summary of the standard framework. As explained in Part II in this group of posts, in a classic statistical-mechanical system in the ergodic state (non-autocorrelated fixes, in the context of a home range) there is no “surprise factor” when we are observing the system at different spatial scales. In short, the total entropy is not influenced by our observational scale. For example, if we superimpose a virtual grid upon a set of “classically distributed” GPS fixes (i.e., in compliance with the convection model for the home range-generating process) and observe the system in its ergodic state, the total entropy depends only on the spatial extent, not on the choice of grid resolution (unless the sample of fixes is serially autocorrelated, as described in another post). In the equilibrium state, the entropy for the home range equals the sum of entropy in respective grid cells multiplied with a trivial conversion factor to compensate for difference in resolution, whether we choose to study the system from a fine or coarse spatial resolution.

This property of a classically derived statistical mechanics explains why in this branch of physics – the classic Boltzmann-Gibbs statistical mechanics – you won’t find any reference to an “observer effect” on entropy from zooming over pixel scales.

Translating this standard statistical-mechanical framework to a more familiar context of statistical ecology, the sum of grid cell variance equals the total variance of the system, quite independent of the grid resolution. The variance is thus reflecting a statistically stationary space use process. Consequently, in this classical scenario stationary variance (and entropy) supports the prediction of a home range area asymptote; a stationary home range size that will be revealed in more detail as N is sufficiently increased to reduce the statistical artifact from a small sample size. For larger N, home range area is practically constant under a change of N. Hence, the embedded entropy is similarly constant. In this manner I have hereby described a bridge between statistical ecology and statistical physics under the standard framework.

superpositionofgridsHowever, more bridge-building is needed, in particular with respect to complex animal space use. In the illustration to the left (copied from Gautestad et al. 1998) I have indicated a geometric scaling of a virtual grid that is superimposed upon a set of GPS fixes; cells scale by a power of 2 in this “sandwich” of resolutions. The choice to use power of 2 means that the respective resolutions’ cell areas scale by a factor of 4 (from top level towards finer resolutions: 1, 1/4, 1/16, 1/64, etc). This geometric scaling has a purpose, because it illustrates how a multi-scaled process – “complex space use” – is postulated to scale in a similarly geometric manner.

One may call it remarkable or fascinating – but this geometric scaling is what may be predicted from the parallel processing (PP) conjecture of the MRW model. According to this extended framework, entropy is postulated to be dispersed over a scale range, not just at a specific “reference” spatial scale and integrating (summing) from this reference to find the entropy at other scales. Thus, parallel processing defies standard statistical assumptions, and the system’s statistical-mechanical framework consequently needs to be extended. In both frameworks; i.e., including the extended system scenario, the log-transformed entropy is expanding proportionally with system extent. However, in the extended framework, the theory predicts the rate of expansion to be geometrically smaller (four times larger extent is needed to find a doubling of entropy, in a scale-free manner), since 50% gain of entropy is cancelled out due to the 50% parallel loss of entropy that is swamped at finer resolutions (see this post)!

  • In the standard framework, entropy expands proportionally with area or volume of the system; i.e., the system’s spatial extent.
  • In the parallel processing framework, entropy expands both with area/volume (system extent) and the scale range (system’s spatial grain) over which the process is observed.
  • The latter scale range – over which entropy is distributed uniformly (on a log-scale) – is a function of temporal observation intensity. The scale range expands with observation intensity, N, in accordance to the “inwards” and “outwards” expansion of self-organized, scale-free dispersion of incidence (cells with 1 or more fixes) at respective scales. The middle layer (in log-scale terms) is the individual’s characteristic scale of space use, CSSU!
  • Interestingly, both the standard and the PP-extended theroretical framework describe a logarithmically scaling function for the change of entropy with scale.
  • Since we assume – at this stage – a system in its ergodic state, the temporal observation intensity may be increased by increasing N from (a) increasing the observation period T at a given lag t, or (b) increasing the observation frequency 1/t within a given T (as far as the ergodic state allows for; i.e., the sample is still non-autocorrelated). Thus observed increase of entropy from a given observation intensity depends on the non-denominated product of frequency and time, 1/t*T=T/t. Hence, the model predicts a similar increase in entropy whether T is increased (T∝N) or t is decreased (1/t∝N).

For example, let us assume that we have estimated the actual CSSU as level “3” in the illustration. We have achieved this by analyzing a large set of GPS fixes, collected at a lag that satisfies serial non-autocorrelation, following the procedure in a previous blog post. For the given observation intensity (sample size N), we then expect to find compliance with a statistical fractal as we zoom towards finer or coarser scale levels below and above the CSSU level, as far as the given sample size N allows for. I have previously described (Part III of this set of posts) how entropy contraction towards lower levels equals entropy expansion towards higher levels over this scale range surrounding CSSU.

Thus, if the given N is increased sufficiently to maintain a given fractal dimension down to level “3 – 2” (three minus two) at a logarithmic scale (1/16 cell size relative to CSSU, if we use base 2 for the logarithm), we expect this pattern to simultaneously having expanded to level “3 + 2” (i.e., 16 times larger cell area relative to CSSU). This relationship – where entropy depends on observation intensity, N – clarifies that parallel processing-driven space use by a “particle” (an individual in our scenario) deviates qualitatively from classical statistical mechanics by requiring an extra dimension orthogonal on the spatial scales: the scale range per se.

The latter property may for example be observed as the Home range ghost: home range area (e.g., incidence) expanding proportionally with the square root of N. If we extend the system extent beyond the observed home range area from N fixes, we will of course not see any extra area. The additional set of grid cells are all empty. Similarly, if we expand the grid resolution towards even finer resolutions, we will similarly not find additional incidence unless N is expanded further. Waiting for larger ; i.e., stronger observational intensity, the additional set of cells are all empty. These situations have been described as “the space fill effect” and “the dilution effect”, respectively (Gautestad and Mysterud, 2012). In this blog post I’ve connected these concepts to the system’s statistical-mechanical properties, under the premise of complex (parallel processing-based) space use.


Gautestad, A. O., and I. Mysterud. 2012. The Dilution Effect and the Space Fill Effect: Seeking to Offset Statistical Artifacts When Analyzing Animal Space Use from Telemetry Fixes. Ecological Complexity 9:33-42