Populations on the verge of extinction:

treelines and other range margins

Photo: George Shepherdc

Beáta Oborny
Biological Institiute, Loránd Eötvös University
Institute of Evolution, Loránd Eötvös Research Network
 Collaborators so far in the broader project,
Ecological Boundaries

Gábor Csányi

Cambridge University (UK)

György Szabó Róbert Juhász

MFA, KFKI Wigner Institute of Physics

Géza Meszéna, Jeromos Vukov, Michael Gastner, Beáta Oborny, Igor Kovács, Dániel Zimmermann, Ole Peters

ELTE , Santa Fe Institute (USA), Imperial College of London (UK)

 Spatial models
1. Dynamics of extinction 2. Formation of range margins across
in homogeneous space environmental gradients

Photo: George Shepherdc

 A general observation: zonation
Sharp boundaries of can emerge across
smooth environmental gradients.

Photo: George Shepherd What are the minimal conditions

for the zonation of vegetation?

Geometry and movement

of the boundary (zone).

Alexander von Humboldt

at Mt Chimborazo, 1801-1803
Painting: Friedrich Georg Weitsch
 Zooming in: complex structure, fluctuations

SW Yukon, USA Glacier National Park, USA

Photo: Danby, R.K., 2011. Map: Zheng, Y. & Malanson, G.P.,

2006. Geograph. Analysis 38: 271-287

How to delineate the range of distribution?

How to distinguish between fluctuations and trend-like shifts?
 A basic, general model of occupation of space

States: empty vs. occupied

Population dynamics Metapopulation dynamics

The components are individuals.

The components are (sub)populations.
The events are birth and death.
The events are colonization and extinction.
Non-spatial version: the logistic model.
Non-spatial version: the Levins model.

dN K−N dN
= r0 N   = cN (1 − N ) − eN
dt  K  dt

They are equivalent.

Homogeneous contact process (no gradient)

c
Colonisation:

Extinction: e

c
Rate of spreading: =
e
Homogeneous Contact Process

ContactProcess.exe
 Phase transition at a critical threshold

Steady-state occupancy

mean variance

nˆ  ( − c )  V
( − c ) −
A
V

 
c = 1.649 c = 1.649
 Applicable to various situations with local spreading.

The shapes of the curves are known near the extinction threshold ( c ).
These shapes, even the values of the exponents (in 2D), are independent of the local
details of the model.
.

Scaling laws:
Mean occupancy nˆ  ( − c )  , where  = 0.583

Variance of occup. ( − c ) − , where  = 0.35(1)

V
A
Autocorrelation distance   ( − c ) −  , where  s = 0.733(4) in space
t = 1.295(6) in time

This process belongs to a described universality class of phase transitions:

directed percolation. This could provide a general, parsimonous explanation for zones.
Homogeneous Gradient

Photo: GeorgeShepherd
 Contact process
1. homogeneous 2. gradient


c = 1.649

c c( x )
=  ( x) =
e e( x )

Unequivocally determines the Provides a good approximation,

steady-state population size except in case of strong parent-dependence and
abrupt changes in the environmental conditions
 Some objectives in gradient environments

1. Monitoring the effects 2. Predicting the expansion of

of climate change invasive species (e.g. pests)

Gypsy moth
(Lymantria dispar)

Range shifts Frontline + outposts

Are the external occurrences („outermost islands”) suitable for the delineation?

Theoretical considerations suggested that this is not a good idea, because the
correlation length and relaxation time diverge.
Large areas and long times of observation would be needed.

We tried a different approach:

▪ Approach the edge from the dense region
▪ Keep the resolution as fine as possible (no artificial coarsening)
▪ Utilize that we have info about the pattern-generating process
An alternative approach: let’s look at the boundary
between connected vs. fragmented occupancy

Percolation Outlining
with step length s the hull
Gradient Contact Process
Boundary.exe
 The hull is a (stochastic) fractal with dimension D=7/4.

This fractal structure does not change

Self-similarity when we change the slope.

steep

smooth

We can even vary the

shape of the ‚hillside’.

Gastner & Oborny, 2012, New Journal of Physics 14: 103019

The fractal structure does not depend on some local details of
the population dynamics either.

Extinction Colonization
D = 7/4
GRM
e(x) c(x)

Gastner, Oborny, Pruessner & Zimmermann, 2009, American Naturalist 174: 23-39.
 The fractal structure is not changed by a fine-grained heterogeneity in the
environment (on the top of the gradient).
D = 7/4
Environment The Rugged Slope Model
(the contrast
could also be
changed)

Occupancy
in a snapshot
The largest cluster Estimation of the fractal dimension
and the hull in case of various slopes

Long-term
occupancy

Juhász & Oborny, 2019, Ecological Complexity 42, 100814

Does the fractal structure remain the same when the environment is changing,
thus, the population is advancing or retreating?

advancing front

standing

retreating
The fractal structure remained the same when the environment was changing,
thus, the population is advancing or retreating

D = 7/4 Fast retreat Slow retreat Standing Slow advance Fast advance
front

v=-100% D=7/4 v=-5% D=7/4 v=0% D=7/4 v=5% D=7/4 v=14% D<7/4

D The trailing edge was less distorted

than the leading edge.
Even at the leading edge, the fractal
dimension remained D=7/4 when the
advance was reletively slow.
v
Zimmermann & Oborny, 2023, Ecography, accepted mscr.
The position of the hull is sharp. This is advantageous for monitoring.

Fragmented Connected

Width of the hull?

Length of the hull?

They also have

characteristic scaling laws.

Density profile of the hull

Gastner, Oborny, Pruessner & Zimmermann, 2009, American Naturalist 174: 23-39.
 Fragmented Connected
Proposal

Let us define the range’s edge as

the average position of the hull,
i.e., of the boundary between
fragmented and connected occupancy,

because...
■ it is not sensitive to the local details (pop., env.),
■ its structure remains the same during range shifts
(except for fast advancing fronts), and
■ the structure can be described by characteristic
scaling laws, thus, it is possible to check whether
we have. found it.
.

Thus, it permits comparisons between years, places or species.

An example from the Aggteleki Karszt hills, Hungary

Image and analysis: Eszter Tanács

 Suggestion for 3 research tasks
1. Gradient Random Map 2. Gradient Contact Process 3. Real-life data

A) Simulation: GRM
A) Simulation: GCP
B) Identifying the percolation clusters
B) Using 1.B-D on these data
C) Marking the hull of the giant cluster A) Downloading maps + consultation with a
C) Generalization of 2.A: any c(k) and e(k)
D) Estimation of D cartographer (with Beata)
D) Selecting one of these: the Allee effect
E) Study of the finite-size effect (changing Ly) B) Transformation of these into pixel maps
F) The efficiency of different methods (D & E) C) Image analysis -> binary maps (tree / not tree)
D) Using 1.B-D on these data
E) Treeline summary

Interest in programming and fractals Interest in dynamical systems Interest in real treelines

Aiming at a mathematical paper Aiming at a theoretical biological paper Aiming at a practical biological paper