The Hidden Layer

Focusing on the statistical pattern of space use without acknowledging the biophysical model for the process will create much confusion and unnecessary controversy. Ecologists are now forced to get a better grip on concepts from statistical mechanics than earlier generations. For example, to understand the transformation from data on actual behaviour to pattern analysis of space use, the concept of the hidden layer represents the first gate to pass.

Research on animal movement and space use has always had a central place in ecology. However, as more field data, better computers and more sophisticated statistical methods have become available, some old dogma have come under attack. Specific theoretical aspects of this quest for improved model realism have emerged from the rapidly growing cooperation between biologists and physicists in the emerging field of macro-level biophysics. The so-called Lévy flight foraging hypothesis is one example. And, of course, I can’t resist mentioning the MRW theory.

A booted eagle Hieraaetus pennatus is triggering a flock of spotless starlings Sturnus unicolor to show swarming behaviour. Malaga river delta, December 2017. Photo: AOG.

In 1985 Charles Krebs described ecology as the scientific study of the interactions that determine the distribution and abundance of organisms. In an ethological context animal space use is studied on two levels – tactical and strategic. The tactical level regards understanding individual biology and behavioural ecology on a moment-to-moment temporal scale. Strategic space use adds an extra layer of complexity to the tactical behaviour. In a simplistic manner we may refer to this layer as the animal’s state at a given moment in time; for example whether it is hungry or not (e.g., in hunting mode). Strategy also involves processing of memory-based goals. Strategies executed at coarser time scales than tactics. Some of the interaction between tactics and strategy may then – under specific conditions (see below) – be transformed to dynamic models at the tactical level; so-called mechanistic models, which consists of a set of executable rules covering respective cognitive and environmental conditions. Validating the model dynamics and resulting statistical patterns against real animal data then rates the degree of model realism. For example; realistic, tactical models have been developed to cast light on the “clumping behaviour” (dense swarming) of flock of birds that are threatened by a raptor.

The myriad of rules that influence animal movement makes detailed modelling an impossible task, and would anyway only lead to a descriptive picture with no value to ecological hypothesis testing. In fact, the signature of successful modelling is simplification. Thus, only specific aspects of the reference individual’s behaviour can be included and scrutinized.

The present post addresses one particular aspect of system simplification; coarse-graining the temporal scale. This approach implies a qualitative change of how the space use system is observed and analyzed. Actually, temporal coarse-graining is forced upon us when studying animal space use from sampling an individual’s successive displacements as a series of locations (fixes) during a given period of time. During each inter-fix interval the observed displacement regards the resultant vector from a myriad of intermediate and unobserved events. What has happened to the moment-to-moment kind of behavioural ecology? It has become buried below the hidden layer.

At the surface of this hidden layer you lose sight of behavioural details (like raptor response and swarming rules) but you gain access to an alternative perspective of movement and space use. Alternative statistical descriptors are emerging at this temporally coarser scale, following the laws of statistical mechanics. What is analyzed above the hidden layer is the over-all pattern from many displacements events that are aggregated into a spatial scatter of fixes.

For example, you may coarse-grain both the temporal and spatial system dimensions, and study the aggregated distribution of fixes at the spatial scale of virtual grid cells (pixels) and temporal scale of the fix sampling period. The spatio-temporal variations in intensity of space use within the actual space-time extents then allows for modelling and hypothesis testing, but now using statistical-mechanical descriptors of space use intensity. These descriptors are either not valid below the hidden layer (e.g., the information content of local density of fixes) or they have an alternative interpretation (e.g., movement as a “step” versus movement as a resultant vector for a given interval and location). Both levels of analysis require large sets of input to allow for statistical treatment.

Why is the hidden layer concept and the statistical-mechanical approach more important to relate to today than in earlier decennia? The short answer is the realization – seeded by better and more extensive data – that animal space use involves more than a couple of universality classes of movement (see this post). In fact, in my book, papers and blog posts I have detailed eight classes, most of which are unfamiliar to you.

To understand space use that is influenced by spatial memory and scale-free movement, statistical mechanical modelling is a prerequisite for realistic representation of such complex systems, unless you limit your perspective to a short-term behavioural bout within a very localized arena. In other words, “a single piece of a jigsaw-puzzle of space use dynamics”. For example, if you zoom closely into a small segment of a circle you observe an approximately straight line. Take a step outwards, and you are facing the qualitatively different geometry – the mathematics of a curve and finally a full circle. Stubbornly staying within the linear framework when analyzing more extensive objects than what you observe at fine scales will force you into a corner filled with paradoxes.

Fine-grained and coarse-grained analyses of animal space use are complementary approaches to the same system.


MRW and Ecology- Part VII: Testing Habitat Familiarity

Consider having a series of GPS fixes, and you wonder if the individual was utilizing familiar space during your observation period – or started building site familiarity around the time when you started collecting data. Simulation studies of Multi-scaled random walk (MRW) shows how you may cast light on this important ecological aspect of space use.

First, you should of course test for compliance with the MRW assumptions, (a) site fidelity with no “distance penalty” on return events, (b) scale-free space use over the spatial range that is covered by your data, and (c) uniform space utilization on average over this scale range. One single test in the MRW Simulator, the A(N) regression, cast light on all these aspects. First, you seek to optimize pixel resolution for the analysis (estimating the Characteristic scale of space use, CSSU). Next, if you find “Home range ghost” compliance; i.e., incidence I expands proportionally with square root of sample size of fixes, your data supports (a) spatial memory utilization with no distance penalty due to sub-diffusive and non-asymptotic area expansion, (b) scale-free space use due to linearity of the log[I(N)] scatter plot, and (c) equal inter-scale weight of space use due to slope ≈ 0.5.

Supposing your data confirmed MRW, how to test for time-dependent strength of habitat familiarity? Consider the following simulation example, mimicking space use during a season and under constant environmental conditions.

The red dots show log(N,I) for various sample sizes up to the total set of 11,000 fixes. Each dot represents the average I for respective N of the two methods continuous sampling and frequency sampling (counteracting autocorrelation effect; see a previous post). However, analyzing the first 1,000 fixes separately (black dots) consistently revealed a more sloppy space use in terms of aggregated incidence at a given N, relative to the total season. The next 1,000 fixes, however, was compliant with the total series both with respect to slope and y-intercept (CSSU) (green dots).

The reason for the discrepancy in space use during the initial period of fix sampling* was in the present scenario the actual simulation condition; site familiarity was set to develop “from scratch” simultaneously with the onset of fix collection. I define strength of site familiarity as proportional with the total path length from which the model animal collects a previous location to return to**. In the start of the sampling period, the underlying path is short in comparison to the total path that was traversed during the total season, and – crucially – return steps targeted previous locations from the actual simulation period only, and not locations prior to to this start time. In other words, the animal was assumed to settle down in the area at the point in time when the simulation commenced.

To conclude, if your data shows CSSU and slope of similar magnitude in the early and later phase of data collection, you sampled an individual with a well-established memory map of its environment during the entire observation period. The implicit assumption for this conclusion is of course that the environmental conditions was constant during the entire sampling period, including the initial phase. Using empirical rather than synthetic data means that additional tests would have to be performed to cast light on this aspect.


*) The presentation above reflects the pixel resolution that was optimized for the total series. The first 1,000 fixes showed a more coarse-grained space use, reflected in a 50% larger CSSU scale (not shown: optimal pixel size was 50% larger for this part of the series) despite constant movement speed and return rate for the entire simulation period. In this scenario a larger CSSU [coarser optimal pixel for the A(N) analysis] signals a less mature habitat utilization in the home range’s early phase. The CSSU was temporarily inflated during build-up of site familiarity, but – somewhat paradoxically – the accumulated number of fix-embedding grid cells (incidence) for a given N at this scale was smaller. These two effects, reflecting degree of habitat familiarity during home range establishment, should be considered a transient effect.

**) Two definitions should be specified:

  • I define strength of site familiarity as proportional with the total path length from which the model animal collects a previous location to return to.
  • I define strength of site fidelity as proportional with the return frequency.

Both definitions rest on the assumptions of no distance penalty on return targets and no time penalty on returns; i.e., infinite spatio-temporal memory horizon relative to the actual sampling period.

The MRW Simulator: Importing Your Own GPS Data

You have a large database of GPS fixes, and you wonder if your animals have utilized their habitat in accordance to standard theory of mechanistic movement (the null hypothesis) or in compliance with the MRW theory (the alternative hypothesis). The MRW Simulator is tailormade for this kind of test.  If MRW is verified you may proceed with various analyses of behavioural ecology under the alternative statistical-mechanical theory. The initial test procedure is simple: (1) import your data, (2) prepare for a test of model compliance by applying one or more built-in algorithms, and (3) import the generated data tables for statistical test into third party packages (R, Excel, etc.).

You can import data to the MRW Simulator by preparing a two-column text file, using comma or TAB as delimiter between the two coordinate values for successive locations.

By default you should use the file name import.txt, but other names are also allowed (given the correct data structure). Place the file in the data folder (…/mov) and choose the menu “File | Import data from txt or csv”.

You are asked to define the file name for the imported data. By default, the name is set to “seed1.txt”. During importing the original series is centred on coordinate (0,0), the middle of the arena window. The arena size for the analysis is automatically adjusted to twice the space needed to display the set of the imported fixes.

After import, you set a couple of check boxes on the MRW Simulator’s user interface in accordance to User guide before clicking the run button (the MRW simulator re-formulates your imported data to its own format and saves the result in the text file levy1.txt). In particular, setting simulation series length to zero and choosing “use seed1.txt” as first part of the simulated series ensures that only your own data are reformatted. Within a fraction of a second the procedure exits without adding simulated data to the series, and you are ready to perform various analytical tasks on the levy1.txt file (see menu “Analyze”).

The procedure “A(N) regression” is typically applied to analyze space use at the home range scale. It is a convenient choice to test for MRW compliance of your data.

You are asked which of the Levy*.txt files to analyze for fix-filling area as a function of sample size N (number of fixes in the Levy*.txt file). Next, the analysis is executed in accordance to the scales set in “Arena extent for analysis”, “Arena grain for analysis” and “Pixel (intra-grain resolution)” in the MRW Simulator’s user interface.

In this procedure, set extent = grain. Pixel regards a ratio; the relative resolution of grain/pixel. For example, setting pixel = 10 performs analysis at the virtual grid scale 1/10 of arena scale; i.e., 10×10 grid cells. See User guide for details.

The progress is shown below the arena window. The algorithm is counting incidence over a range of sample sizes N at the given pixel resolution; first by sequential (continuous) sampling up to Ntotal and then by frequency (uniform) sampling over the total series. Search my blog or read my book for these concepts.

The result is saved in a text file containing a table of incidence (non-empty grid cells at the given pixel resolution) as a function of sample size N under the two sampling conditions. These data may then be imported in for example Excel for graphical presentation and statistical analysis; for example, a regression.

If you find a discrepancy between the scatter from the two sampling methods (you normally do!), your data is probably serially autocorrelated. To remove autocorrelation effect, take the average incidence for respective magnitudes of N, as was explained in this post. Conveniently, the MRW Simulator does this task for you (you find the averaging table below the tables for continuous and frequency sampling). This averaging procedure also adjusts for a “drifting home range” scenario, which also produces autocorrelation.

Does the result support MRW? First, you must verify presence of a characteristic scale of space use (CSSU), which is a property of scale-free movement under influence of spatial memory under the “parallel processing” postulate.

To test for CSSU you should experiment with various pixel resolutions and see if the log[(N,incidence)] pattern converges to a slope ∼0.5 at a given scale. If so, CSSU ≈ (pixel scale)2 = c.

If you don’t find reasonably good compliance with linearity of log[I(N)] = log(c) + 0.5*log(N) or the slope exceeds 0.5, try a coarser pixel resolution. If the slope is smaller than 0.5, try a finer pixel resolution.

If this test of convergence to log-linearity with slope ≈ 0.5 fails, you have probably either supported one of the null models; i.e., Brownian motion-like or Lévy-like space use void of spatial memory influence (slope ≈1, which is quite insensitive to change of pixel scale) or the classic paradigm: home range movement under influence of a constraining border zone [I(N) showing an area asymptote rather than a power law expansion with exponent close to 0.5].

The MRW Simulator 2.0 will now be made available as a free add-on tool for all buyers of my book. If you purchase it through my shopping cart at, you will get the program and its user guide bundled with the book. Existing book owners: contact me at and I’ll fix you a personal download link – free of charge. You may purchase by invoice – see top of this page!

The MRW Simulator – Finally Available!

Back in 1997 I started programming the foundation for a personal simulation environment for Multi-scaled random Walk, the MRW Simulator. Through countless updates over these 20 years the program has gradually matured into a version which finally is ready for limited distribution towards peers in the field of animal space use research.

The MRW Simulator is a Windows©-compliant tool to generate various classes of animal movement (self-produced data series) or to import existing data series. The generated or imported data – consisting of a sequence of (x,y) coordinates – may then be subject to various kinds of statistical protocols through simple menu clicks. The generated text files are then typically exported for detailed analyses and presentation of results in other applications, like the R package or Excel©.

While R is based on an interpreted language, the MRW Simulator is a fully complied program. Thus, movement paths of length up to 20 million steps may be simulated within minutes of execution time, rather than multi-hours or days. A multi-scaled analysis of data over a substantial scale range is almost forbidden in an interpreted system due to the algorithm’s long execution period. In the MRW Simulator such analyses are performed in a fraction of this time. Thus, R and the MRW Simulator may supplement each other. R is strong on statistics and algorithmic freedom; the MRW Simulator is strong on time–effective execution of a small set of basic but typically time-consuming algorithms.

The opening screen contains menus (1), a window where the simulated or imported set of fixes are displayed (2) and various command buttons, check boxes and information fields (314).

To get your first experience with the system, try out the most basic setting for a simulation. First, choose among classes of movement; Levy walk/MRW, Correlated random walk, and Composite random walk (superposition of two correlated random walks) (3). The difference between LW and MRW is explained below.

For your first test, choose Levy walk / MRW (3), with default setting for fractal dimension (D=1) and maximum displacement length between successive steps (truncation=1,000,000 length units). D=1 simulates the condition where the animal on average utilizes its environment with similar scale-free weight at each intermediate scale from unit step length to maximum step (setting 1<D<=2 skews space use towards finer-scale space use on expense of coarser scales, again in average terms).

In a column of text fields (4) you may define conditions like series length, properties for the simulated path, size of the arena and grid resolution for the subsequent analysis. For example, the difference between Levy walk and MRW is given by setting a return frequency >0 for MRW (implying targeted return events to previous locations at the chosen average frequency). For this first run, just keep the default values.

Later you will learn how to additionally modify the conditions by including a pre-defined series of coordinates (in a file called seed*.txt, where * regards an incremental number) (5). At this stage, just keep default settings.

By default the simulation runs in a homogeneous environment. The set of “Habitat heterogeneity” fields (6) allows defining the corners of a rectangle where the model animal behaves in a more “fine-grained” manner by reducing average movement speed. Other ecological aspects may also be defined, like a method to account for temporal and local resource exhaustion. As a start, just keep defaults.

Now, click the “Single-series” command button (7). You should see a number of fixes appearing as dots in the arena window.

The number of fixes reflect the ratio of total series length and the observation interval on this series; i.e., “Number of fixes” (Norig= 1,000,000) multiplied by an average “Observation frequency” (p=0.001). This leads to an observed series length – a path sample – of ca 1,000 fixes; which are displayed in the observation window.

Before moving on to your first data analysis, observe that the simulation’s default settings are defined by “schemes”, which can be pre-loaded from a dropdown menu (8). You may also run a number of replicate simulations in an automated sequence (9). The arena may be copied to the clipboard (10) for subsequent pasting into other applications like a Word document, an Excel sheet, etc.

The “Data path” field (11) displays the folder where the system saves and retrieves data. By default, the data resides in a subfolder, “\mov”, under the location of the MRW simulator’s EXE file. This location is set during program setup.

The field “Fractal resolution range” (12) defines the scale range over which a subsequent analysis of the scatter of fixes – selected from the Analysis menu – will be performed by the so-called box counting method.

The field “A(N)” (13) shows the progress of another analysis, total area (incidence) as a function of sample size, N.

The counter (14) is automatically incremented each time you click the “Singe-series” button (7). TIP: To repeat (and overwrite) an existing series, edit the counter number (14) to one decrement below the actual series. For example, to re-execute data series number 5, edit the counter field to “4” before clicking the button (7). To re-execute series 1, edit the field to “-1” (the number zero is reserved as the initial setting number).

The data file containing “observed” fixes resides in the \mov folder (see above), with name “levy*.txt”. (* = 1, 2, 3, …). It contains three columns of data; x-coordinate, y-coordinate, and inter-step distance.

The MRW Simulator 2.0 will now be made available as a free add-on tool for all buyers of my book. If you purchase it through my shopping cart at, you will get the program and its user guide bundled with the book. Existing book owners: contact me at and I’ll fix you a personal download link – free of charge. You may purchase by invoice – see top of this page!

In the next blog post I’ll show some of the menu procedures of the MRW Simulator, including how to import you own GPS space use series for analysis on-the-fly.

MRW and Ecology – Part VI: The Statistical Property of Return Events

Animals that combine scale-free space use with targeted returns to previous locations generate a self-organized kind of home range. In short, the home range becomes an emergent property from such self-reinforcing revisits. Obviously, any space use pattern from complex processes outside the domain of Markov (mechanistic) theory needs to be analyzed using methods that are coherent with this kind of behaviour. Below I exemplify further the versatility of the MRW approach to adjust for serial auto-correlation (see Part III). I also show the quite surprising model property that the sub-set of inter-fix displacement lengths for return events seems to have a similar statistical distribution as the over-all pattern of exploratory step lengths. This additional emergent property of space use may lead to methods to test a wide range of behaviour-ecological hypotheses, for example to which extent an animal calculates on an energy cost with respect to distance to potential target locations for returns..

In ecological research it is traditionally considered logical that an animal considers a return to a distant familiar location to be less preferred than revisiting closer locations. On the other hand, by default (a priori) the MRW model does not include such a distance penalty on long-distance returns. Recently, the realism of this model premise has gained empirical support from studies on bison and toads (Merkle et al. 2014,2017; Marchand et al. 2017). In the MRW model’s standard version, a given return step is targeting any previous locations with equal probability except for the additive effect of number of previous visits to a given site, which increases the statistical probability for future revisits (self-reinforcing site fidelity). The implicit assumption is that the added energetic cost from long distance returns either is negligible relative to other parts of the energy budget, or the fitness value from keeping in touch with familiar locations regardless of current distance far supersedes the energy consideration. While this property regards a homogeneous environment it is trivial to adjust to a heterogeneous scenario without loss of the general principle. In this post I present more details on the return step property of MRW from a theoretical angle, as a starting point to test the model’s default condition on real data.

First, consider that the robustness of the MRW-based method to estimate an individual’s characteristic scale of space use (CSSU) within a given time and space extent is key to understand the energy aspect of return events as outlined above. The property of return events imposes a characteristic scale; i.e., CSSU, on space use, despite the scale-free nature of exploratory steps. For a given period, CSSU is a combined function of average movement speed and average return frequency. In a previous post I proposed how CSSU may be estimated even in auto-correlated (“over-sampled”) data series of location fixes. In this post I present a pilot analysis which strengthens this approach.

Consider the two simulated Home range ghost results to the right; incidence, I, as a function of number of fixes, N. The first set of fixes (circles) regards weakly auto-correlated series of fixes from return rate 1:10 and fix sampling at 1:100, while the second series (squares) resulted from a strongly autocorrelated path sampling (return rate 1:100 and sampling at 1:10). As was shown in Part III of this set of blog posts, by performing the “averaging trick” on log(I,N) from frequency and continuous sampling (open symbols for respective sets) the average log-log slope remains close to z=0.5 (area expanding proportionally with square root of sample size) even for the strongly auto-correlated series. The slight deviance from z=0.5 in the two series should be considered normal variability to be expected from one simulated series to the next (averaging over large sets of series would bring z closer to 0.5).

Critically, the present result also shows compliance with the expected change of the characteristic scale of space use (CSSU, represented by the parameter c in the Home range ghost formula I=cN0.5) as a function of the ratio between frequency of return events relative to exploratory moves (assuming constant average movement speed). In other words, observation frequency, which represents a sub-set of all displacements along a path (sampling of fixes) should not influence the CCSU estimate despite influencing the degree of auto-correlation. According to MRW theory, fewer returns during a constant average movement speed lead to larger CSSU*. In the analysis of the present two series, ten times smaller return rate led to an optimized unit pixel size (I≡1) of magnitude √10 = 3.3 times larger than for the weakly autocorrelated series with higher return frequency.

In the Figure above, the two CSSU scales have both been rescaled to c=1 ([log(c)=0], but respective series’ unit scale (I=1) is de facto correctly found to be very different in absolute terms.In the present examples, CSSU was estimated to c1 = 1252 area units for the high frequency return scenario and c2 = 4002 area units for the second series with fewer returns (and stronger degree of autocorrelation).

To conclude, after optimizing pixel size in respective series by analyzing I(N) over a range of pixel resolutions as previously described in my book and other posts, this preliminary analysis verifies a strong coherence between return step frequency and the magnitude of CSSU in accordance to the theoretical parameter prediction, despite strong difference in degree of serial autocorrelation in the sample of relocations. On other words, the CSSU estimate is quite resilient to the researcher’s choice of fix sampling scheme.

However, another aspect of the return step component of may turn out to be valuable to test the opposing energy hypotheses with respect to distance penalty, as outlined above.

Quite surprisingly I must admit, even considering the implicit “no distance penalty” model design, the tail part of the step length distribution of returns is quite similar to the tail of observed step lengths (fixes) that are sampled from the total series of steps!

The example series above with the weakest degree of auto-correlation (circles in the top Figure) shows similar functional form between the over-all distribution of binned step lengths [log(L); red circles below] and return distances (open symbols)**.

As expected from the weakly autocorrelated series, the fit to the power law function with Levy exponent β=2 of the exploratory steps is showing a clear “hump” in the extreme part of the Log(L) distribution of fixes, due to influence from intermediate return events.

For the more strongly autocorrelated series with N=10,000 fixes from a total series of 100,000 steps and a lower return frequency 1:100 (Figure below) we see – as theoretically expected – a more subdued hump for the fixes, due to less influence from return events***. The hump would be even less pronounced if the fix sampling frequency had been even larger (Gautestad and Mysterud 2013; in particular Figure A2 in Supplementary material).


Again the tail distribution of return lengths – where the total set of 1,000 events is shown as triangles – is similar to the the over-all distribution of fixes (1,000 first and 1,000 last of the N=10,000 fixes, shown as red and green circles). The median length for return steps is larger under this scenario (740 length units, versus 262) due to a ten times lower return frequency in relative terms. On the other hand, the median length for the actual set of fixes is strongly reduced as a consequence of the ten times larger fix sampling frequency.

To summarize, while the estimate of CSSU is quite resilient to fix sampling frequency, the (observed) median step length of fixes and (unobserved) length of return steps are influenced by fix sampling rate and return rate, respectively. Despite independence between the median length for observed series and hidden return lengths, both aspects of movement show a similar distribution of lengths.

Finally, what if the return step targets had not been set a priori to be independent on distance; i.e., by invoking distance penalty on return events? I have not tested this aspect yet in a modified MRW simulation model, but intuitively I predict the distribution of return steps to morph towards a negative exponential function rather than a power law, as in the exploratory kind of moves. As aconsequence, the “hump” effect in the distribution of fixes should also be more subdued. Hence, by testing the difference in functional form of return steps and step lengths of observed fixes, one may have a method to test empirically the energy hypothesis that was outlined above.

The challenge, of course, is to develop a method to distinguish between exploratory moves and return events in empirical data. In simulation data it is simple to filter out the returns; in true space use data it is necessary to distinguish returns from path crossing by chance. More on this methodology in an upcoming post.


*) Thus, the ratio returns/exploratory moves have a similar influence on CSSU as a change in average movement speed where the speed is expressed as the average staying time in a given grid cell. In Gautestad and Mysterud 2010, Eq. 4, we defined the expected length of step x, Lx, as a function of a scaling parameter for movement speed δ and fractal dimension of the path, d:

Lx = (δ[1 − Rnd])−1/d         (Eq. 4)

where Rnd is  a random number 0 ≤Rnd < 1 and δ is a scaling parameter. In some sense δ may be interpreted as a parameter for expected staying time in a given patch, since larger δ implies smaller Lx and thus increased local fix contagion.
Gautestad and Mysterud 2010, p2744

Thus, by defining the space use’s fractal dimension D as D≡d, we have the relationship with CSSU’s Home range ghost parameter, c, and movement speed:

c ∝ 1/√δ  |  D = 1          (Eq 5).

**) Due to a return step frequency of 1:10 and actual fix sampling frequency of 1:100, the total set of return events supersedes the fix sample by a factor of 10. Thus, I have compared the distribution of of return lengths from the early part of the simulated path (open squares) with return lengths towards the end of the path (open triangles), keeping both samples at same size as the set of observed fixes. Red circles in the Figure above represent 10,000 fixes from a total series of 1 million steps. When studying the first and the last part of the 100,000 hidden return steps specifically, their distribution looks indistinguishable from the series of “observed” fixes. Triangles show the result for the first 10,000 return events, and the squares show the result from the last 10,000 returns during the total 1 of million steps.

***) In this example where observation frequency supersedes the intrinsic return frequency by a factor of 10, the first and last part of the set of fixes (red and green circles, respectively) was used for comparison with the total set of return steps (open triangles).


Gautestad, A. O., and I. Mysterud. 2010. Spatial memory, habitat auto-facilitation and the emergence of fractal home range patterns. Ecological Modelling 221:2741-2750.

Gautestad, A. O., and A. Mysterud. 2013. The Lévy flight foraging hypothesis: forgetting about memory may lead to false verification of Brownian motion. Movement Ecology 1:1-18.

Marchand, P, M. Boenke and D. M. Green. 2017. A stochastic movement model reproduces patterns of site fidelity and long-distance dispersal in a population of Fowler’s toads (Anaxyrus fowleri). Ecological Modelling 360:63–69.

Merkle, J. A., D. Fortin and J. M. Morales. 2014. A memory-based foraging tactic reveals an adaptive mechanism for restricted space use. Ecology Letters 17:924–931.

Merkle, J. A., J. R. Potts and D. Fortin. 2017. Energy benefits and emergent space use patterns of an empirically parameterized model of memory-based patch selection. Oikos 126:185–195

MRW and Ecology – Part V: Black Bear Home Ranges Revisited

Back in 1994 I enjoyed an unforgettable and extremely inspiring 2-month stay at University of Tennessee, visiting professor Stuart L. Pimm (Department of Ecology and Evolutionary Biology)  and Professor Mike L. Pelton (Department of Forestry, Wildlife and Fishery). During some hectic weeks I worked on transforming the mathematical formulation of the Zoomer model for complex population dynamics into a spatially explicit simulation model (Stuart’s lab) in parallel with interaction with many dedicated students of the biology and ecology of black bear Ursus americanus (Mike’s lab).The Zoomer model is published in my book and already commented on this blog. Regarding the stay at Mike’s lab we published a test on the bears’ general space use, where we found close compliance with the Multi-scaled random walk model, MRW (Gautestad et al. 1998). In this post I revisit the black bear data and find this model’s additional potential to cast light on behavioural ecology in a wildlife management context.

In the 1998 paper we applied the so-called re-scaled range analysis, R/SD (Mandelbrot 1983:pp247-25), to study home range area as a function of fix sample size. Details on the method and the raw data can be found in Gautestad et al. 1998. Below I revisit these bear data and re-test the Home range ghost function with the presently preferred method. Following this exercise I show a new result, which indicates that a collared bear’s characteristic scale of space use may have been inflated during the initial period of fix sampling of its path. Implicitly, the result rises the question if the experience of being collared and getting used to bearing the collar is influencing the bear’s space use towards a more coarse-grained habitat utilization on average; i.e., a larger CSSU, during the first months following the capture/release. The home range ghost regards the “paradoxical” pattern whereby home range area, A, apparently expands non-asymptotically with sample size of fixes, N, in compliance with the power law A = cNz. and z≈0.5. The parameter c is the Characteristic scale of space use, CSSU, emerging from memory-dependent tendency to return to familiar locations.

The inset exemplifies the Home range ghost for one of the black bear individuals. The apparent area asymptote [inset, showing arithmetic axes for A(N)] disappears under log-transformation of the axes; i.e., the expansion with N is non-asymptotic and power law compliant with exponent 0.5. Thus, the area expands with the square root of N.

In the present results to the right and below the A(N) pattern was analyzed with the most recent MRW method, using incidence (number of virtual grid cells of size, I, embedding at least one location) at the estimated grid resolution of CSSU as a representative for home range utilization intensity. The respective (N,I) plots were calculated as the average from two sampling schemes; frequency and time-continuous (see this post). The advantage over R/SD is the I(N) method’s ability to estimate the parameters c (representing CSSU) and power exponent (z) even for autocorrelated series of fixes.

In the present pilot test using a subset of 15 individual data sets (the first part of the total database of 77 series) the result shows strong coherence for estimates of z between the previous R/SD method and the present I(N) method with respect to how home range area responded to sample size, N. Once more I stress the advantage of using a model which accounts for the non-trivial N-dependency in home range size estimates, making CSSU – where the N-dependency is filtered out – a superior ecological proxy relative to the traditional method, direct area estimate.

As expected from scale-free space use, the distribution of CSSU is approximately log-normal (observe the log-scaling x-axis).

On this background it’s time to move in the direction of ecological analysis of respective individual’s space use.The histogram to the right shows a strongly variable CSSU between the 15 individuals. This raises interesting ecological questions. For example, why has the leftmost individual (female F170) 1:240 magnitude of CSSU – ca 1:15 in terms of linear rather than area scale – relative to the rightmost score (female F060)? In other words, according to the present analysis F170 utilized its habitat with substantially higher intensity than F060 (intensity ∝ 1/CSSU). F170 showed a relatively small I(N) for a given N, meaning that F170 utilized its habitat in a more area-restricted manner. Unfortunately I don’t have access to individual behaviour details nor environmental GIS data for the respective space use patterns.

However, consider the relationship between sample size, N, and the CSSU (right). F060 and F170 are found as the largest and smallest CSSU in the scatterplot. The average CSSU for all 15 individuals is ca 866,000 m2 (0.866 km2). Under constant conditions one should expect CSSU to be stationary under larger sample size (larger N) and non-autocorrelated series. However, the plot indicates a negative relationship between N and CSSU in these data, which is unexpected from the standard MRW model a priori (as shown by simulated data in this post)*.

Why? Many hypotheses may be invoked and tested – here I indicate one in particular. The capturing, radio collaring and subsequent tracking might have stressed the bears for some time after release. To study this, I re-analyzed the three individuals with the largest fix sampling period, including the first 100 fixes only. As shown by open symbols, all individuals (F182, F201 and F243) showed larger CSSU during the initial period for 100 fixes following release – a period of ca 1-2 years (!) – relative to the total fix sampling period of almost four years**), and thereby became similar to the other bears with respect to average spatial scale.

Is this an indication of a more restless space use behaviour in the first period following capture, release with radio collar and subsequent telemetry tracking 1-2 times/week (distance observer-animal is unknown)? To explore this aspect, several alternative hypotheses should also be considered (the three individuals were collared in August, June and June, and were 7, 3 and 3 years of age) – but the present pilot test does at least rise an interesting hypothesis. It also illustrates how the MRW model’s CSSU parameter may be applied to cast light on a potentially concerning aspect of data collection and its interpretation in the context of wildlife management***).


*) Update: see also this post. which illustrates a transient effect in the early phase of home range establishment, and stationary CSSU for mature home range utilization.  In the present context, is it reasonable to assume mature home ranges even for the initial 1-2 years of fix sampling.

**) Alternative methods to test for inter-sample difference in space use intensity abound, but have drawbacks. For example, the mean displacement length in a set of fix sampling intervals (lags) will be influenced by intermediate return events. Further, due to the very long-tailed (leptocurtic) distribution of step lengths in data from black bear, which has been verified to comply with the MRW space use model, comparing the relative difference in median displacement size in two or more samples is subject to large error terms due to the extreme outlier issue (“occasional sallies”). CSSU resolves these obstacles.

***) It should bee mentioned that  the radio telemetry collars for black bear back in 1978 were substantially heavier than today’s standard. Triangulation also required close stalking to get a fix, in contrast to modern GPS.


Gautestad, A. O., I. Mysterud, and M. R. Pelton. 1998. Complex movement and scale-free habitat use: testing the multi-scaled home range model on black bear telemetry data. Ursus 10:219-234.

Mandelbrot, B. B. 1983, The fractal geometry of nature. New York, W. H. Freeman and Company.