Competitive Lotka–Volterra equations

The competitive Lotka–Volterra equations are a simple model of the population dynamics of species
competing for some common resource. They can be further generalised to include trophic interactions.

Contents
Overview
Two species
N species
Possible dynamics
4-dimensional example
Spatial arrangements
Background
Matrix organization
Lyapunov functions
Line systems and eigenvalues
Notes

Overview
The form is similar to the Lotka–Volterra equations for predation in that the equation for each species has
one term for self-interaction and one term for the interaction with other species. In the equations for
predation, the base population model is exponential. For the competition equations, the logistic equation is
the basis.

The logistic population model, when used by ecologists often takes the following form:

Here x is the size of the population at a given time, r is inherent per-capita growth rate, and K is the carrying
capacity.

Two species

Given two populations, x1 and x2, with logistic dynamics, the Lotka–Volterra formulation adds an additional
term to account for the species' interactions. Thus the competitive Lotka–Volterra equations are:
Here, α12 represents the effect species 2 has on the population of species 1 and α21 represents the effect
species 1 has on the population of species 2. These values do not have to be equal. Because this is the
competitive version of the model, all interactions must be harmful (competition) and therefore all α-values
are positive. Also, note that each species can have its own growth rate and carrying capacity. A complete
classification of this dynamics, even for all sign patterns of above coefficients, is available,[1][2] which is
based upon equivalence to the 3-type replicator equation.

N species

This model can be generalized to any number of species competing against each other. One can think of the
populations and growth rates as vectors and the interaction α's as a matrix. Then the equation for any species
i becomes

or, if the carrying capacity is pulled into the interaction matrix (this doesn't actually change the equations,
only how the interaction matrix is defined),

where N is the total number of interacting species. For simplicity all self-interacting terms αii are often set to
1.

Possible dynamics

The definition of a competitive Lotka-Volterra system assumes that all values in the interaction matrix are
positive or 0 (αij ≥ 0 for all i,j). If it is also assumed that the population of any species will increase in the
absence of competition unless the population is already at the carrying capacity (ri > 0 for all i), then some
definite statements can be made about the behavior of the system.

1. The populations of all species will be bounded between 0 and 1 at all times (0 ≤ xi ≤ 1, for all i)
as long as the populations started out positive.
2. Smale[3] showed that Lotka-Volterra systems that meet the above conditions and have five or
more species (N ≥ 5) can exhibit any asymptotic behavior, including a fixed point, a limit cycle,
an n-torus, or attractors.
3. Hirsch[4][5][6] proved that all of the dynamics of the attractor occur on a manifold of dimension
N-1. This essentially says that the attractor cannot have dimension greater than N-1. This is
important because a limit cycle cannot exist in fewer than two dimensions, an n-torus cannot
exist in less than n dimensions, and chaos cannot occur in less than three dimensions. So,
Hirsch proved that competitive Lotka–Volterra systems cannot exhibit a limit cycle for N < 3, or
any torus or chaos for N < 4. This is still in agreement with Smale that any dynamics can occur
for N ≥ 5.
More specifically, Hirsch showed there is an invariant set C that is homeomorphic to the (N-
1)-dimensional simplex
 and is a global attractor of every point excluding the origin. This carrying simplex contains
all of the asymptotic dynamics of the system.
4. To create a stable ecosystem the αij matrix must have all positive eigenvalues. For large N
systems Lotka-Volterra models are either unstable or have low connectivity. Kondoh[7] and
Ackland and Gallagher[8] have independently shown that large, stable Lotka-Volterra systems
arise if the elements of αij (i.e. the features of the species) can evolve in accordance with
natural selection.

4-dimensional example
A simple 4-Dimensional example of a
competitive Lotka–Volterra system has been
characterized by Vano et al.[9] Here the growth
rates and interaction matrix have been set to

The competitive Lotka–Volterra system plotted in phase

space with the x4 value represented by the color.

with for all . This system is chaotic and has a largest Lyapunov exponent of 0.0203. From the
theorems by Hirsch, it is one of the lowest-dimensional chaotic competitive Lotka–Volterra systems. The
Kaplan–Yorke dimension, a measure of the dimensionality of the attractor, is 2.074. This value is not a
whole number, indicative of the fractal structure inherent in a strange attractor. The coexisting equilibrium
point, the point at which all derivatives are equal to zero but that is not the origin, can be found by inverting
the interaction matrix and multiplying by the unit column vector, and is equal to

Note that there are always 2N equilibrium points, but all others have at least one species' population equal to
zero.
The eigenvalues of the system at this point are 0.0414±0.1903i, -0.3342, and -1.0319. This point is unstable
due to the positive value of the real part of the complex eigenvalue pair. If the real part were negative, this
point would be stable and the orbit would attract asymptotically. The transition between these two states,
where the real part of the complex eigenvalue pair is equal to zero, is called a Hopf bifurcation.

A detailed study of the parameter dependence of the dynamics was performed by Roques and Chekroun
in.[10] The authors observed that interaction and growth parameters leading respectively to extinction of
three species, or coexistence of two, three or four species, are for the most part arranged in large regions
with clear boundaries. As predicted by the theory, chaos was also found; taking place however over much
smaller islands of the parameter space which makes difficult the identification of their location by a random
search algorithm.[9] These regions where chaos occurs are, in the three cases analyzed in,[10] situated at the
interface between a non-chaotic four species region and a region where extinction occurs. This implies a
high sensitivity of biodiversity with respect to parameter variations in the chaotic regions. Additionally, in
regions where extinction occurs which are adjacent to chaotic regions, the computation of local Lyapunov
exponents [11] revealed that a possible cause of extinction is the overly strong fluctuations in species
abundances induced by local chaos.

Spatial arrangements

Background

There are many situations where the strength of species'

interactions depends on the physical distance of
separation. Imagine bee colonies in a field. They will
compete for food strongly with the colonies located near
to them, weakly with further colonies, and not at all
with colonies that are far away. This doesn't mean,
however, that those far colonies can be ignored. There is
a transitive effect that permeates through the system. If
colony A interacts with colony B, and B with C, then C
affects A through B. Therefore, if the competitive An illustration of spatial structure in nature. The
Lotka–Volterra equations are to be used for modeling strength of the interaction between bee colonies is
such a system, they must incorporate this spatial a function of their proximity. Colonies A and B
structure. interact, as do colonies B and C. A and C do not
interact directly, but affect each other through
colony B.
Matrix organization

One possible way to incorporate this spatial structure is to modify the nature of the Lotka–Volterra equations
to something like a reaction-diffusion system. It is much easier, however, to keep the format of the equations
the same and instead modify the interaction matrix. For simplicity, consider a five species example where all
of the species are aligned on a circle, and each interacts only with the two neighbors on either side with
strength α−1 and α1 respectively. Thus, species 3 interacts only with species 2 and 4, species 1 interacts only
with species 2 and 5, etc. The interaction matrix will now be
If each species is identical in its interactions with neighboring species, then each row of the matrix is just a
permutation of the first row. A simple, but non-realistic, example of this type of system has been
characterized by Sprott et al.[12] The coexisting equilibrium point for these systems has a very simple form
given by the inverse of the sum of the row

Lyapunov functions

A Lyapunov function is a function of the system f = f(x) whose existence in a system demonstrates stability.
It is often useful to imagine a Lyapunov function as the energy of the system. If the derivative of the
function is equal to zero for some orbit not including the equilibrium point, then that orbit is a stable
attractor, but it must be either a limit cycle or n-torus - but not a strange attractor (this is because the largest
Lyapunov exponent of a limit cycle and n-torus are zero while that of a strange attractor is positive). If the
derivative is less than zero everywhere except the equilibrium point, then the equilibrium point is a stable
fixed point attractor. When searching a dynamical system for non-fixed point attractors, the existence of a
Lyapunov function can help eliminate regions of parameter space where these dynamics are impossible.

The spatial system introduced above has a Lyapunov function that has been explored by Wildenberg et
al.[13] If all species are identical in their spatial interactions, then the interaction matrix is circulant. The
eigenvalues of a circulant matrix are given by[14]

for k = 0N − 1 and where the Nth root of unity. Here cj is the jth value in the first row of the
circulant matrix.

The Lyapunov function exists if the real part of the eigenvalues are positive (Re(λk > 0 for k = 0, …, N/2).
Consider the system where α−2 = a, α−1 = b, α1 = c, and α2 = d. The Lyapunov function exists if

for k = 0, … ,N − 1. Now, instead of having to integrate the system over thousands of time steps to see if any
dynamics other than a fixed point attractor exist, one need only determine if the Lyapunov function exists
(note: the absence of the Lyapunov function doesn't guarantee a limit cycle, torus, or chaos).

Example: Let α−2 = 0.451, α−1 = 0.5, and α2 = 0.237. If α1 = 0.5 then all eigenvalues are negative and the
only attractor is a fixed point. If α1 = 0.852 then the real part of one of the complex eigenvalue pair becomes
positive and there is a strange attractor. The disappearance of this Lyapunov function coincides with a Hopf
bifurcation.

Line systems and eigenvalues

It is also possible to arrange the species into a
line.[13] The interaction matrix for this system is
very similar to that of a circle except the
interaction terms in the lower left and upper
right of the matrix are deleted (those that
describe the interactions between species 1 and
N, etc.).

This change eliminates the Lyapunov function The eigenvalues of a circle, short line, and long line plotted
described above for the system on a circle, but in the complex plane
most likely there are other Lyapunov functions
that have not been discovered.

The eigenvalues of the circle system plotted in the complex plane form a trefoil shape. The eigenvalues from
a short line form a sideways Y, but those of a long line begin to resemble the trefoil shape of the circle. This
could be due to the fact that a long line is indistinguishable from a circle to those species far from the ends.

Notes
1. Bomze, Immanuel M. (1983). "Lotka-Volterra equation and replicator dynamics: A two-
dimensional classification". Biological Cybernetics. Springer Science and Business Media LLC.
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ISSN 0340-1200 (https://www.worldcat.org/issn/0340-1200).
2. Bomze, Immanuel M. (1995). "Lotka-Volterra equation and replicator dynamics: new issues in
classification". Biological Cybernetics. Springer Science and Business Media LLC. 72 (5): 447–
453. doi:10.1007/bf00201420 (https://doi.org/10.1007%2Fbf00201420). ISSN 0340-1200 (http
s://www.worldcat.org/issn/0340-1200).
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Mathematical Biology. Springer Science and Business Media LLC. 3 (1): 5–7.
doi:10.1007/bf00307854 (https://doi.org/10.1007%2Fbf00307854). ISSN 0303-6812 (https://w
ww.worldcat.org/issn/0303-6812).
4. Hirsch, Morris W. (1985). "Systems of Differential Equations that are Competitive or
Cooperative II: Convergence Almost Everywhere". SIAM Journal on Mathematical Analysis.
Society for Industrial & Applied Mathematics (SIAM). 16 (3): 423–439. doi:10.1137/0516030 (h
ttps://doi.org/10.1137%2F0516030). ISSN 0036-1410 (https://www.worldcat.org/issn/0036-141
0).
5. Hirsch, M W (1988-02-01). "Systems of differential equations which are competitive or
cooperative: III. Competing species". Nonlinearity. IOP Publishing. 1 (1): 51–71.
doi:10.1088/0951-7715/1/1/003 (https://doi.org/10.1088%2F0951-7715%2F1%2F1%2F003).
ISSN 0951-7715 (https://www.worldcat.org/issn/0951-7715).
6. Hirsch, Morris W. (1990). "Systems of Differential Equations That are Competitive or
Cooperative. IV: Structural Stability in Three-Dimensional Systems". SIAM Journal on
Mathematical Analysis. Society for Industrial & Applied Mathematics (SIAM). 21 (5): 1225–
1234. doi:10.1137/0521067 (https://doi.org/10.1137%2F0521067). ISSN 0036-1410 (https://w
ww.worldcat.org/issn/0036-1410).
 7. Kondoh, M. (2003-02-28). "Foraging Adaptation and the Relationship Between Food-Web
Complexity and Stability". Science. American Association for the Advancement of Science
(AAAS). 299 (5611): 1388–1391. doi:10.1126/science.1079154 (https://doi.org/10.1126%2Fsci
ence.1079154). ISSN 0036-8075 (https://www.worldcat.org/issn/0036-8075).
8. Ackland, G. J.; Gallagher, I. D. (2004-10-08). "Stabilization of Large Generalized Lotka-Volterra
Foodwebs By Evolutionary Feedback". Physical Review Letters. American Physical Society
(APS). 93 (15): 158701. doi:10.1103/physrevlett.93.158701 (https://doi.org/10.1103%2Fphysre
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9. Vano, J A; Wildenberg, J C; Anderson, M B; Noel, J K; Sprott, J C (2006-09-15). "Chaos in
low-dimensional Lotka–Volterra models of competition". Nonlinearity. IOP Publishing. 19 (10):
2391–2404. doi:10.1088/0951-7715/19/10/006 (https://doi.org/10.1088%2F0951-7715%2F1
9%2F10%2F006). ISSN 0951-7715 (https://www.worldcat.org/issn/0951-7715).
10. Roques, Lionel; Chekroun, Mickaël D. (2011). "Probing chaos and biodiversity in a simple
competition model". Ecological Complexity. Elsevier BV. 8 (1): 98–104.
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Phenomena. Elsevier BV. 35 (1–2): 237–250. doi:10.1016/0167-2789(89)90105-x (https://doi.o
rg/10.1016%2F0167-2789%2889%2990105-x). ISSN 0167-2789 (https://www.worldcat.org/iss
n/0167-2789).
12. Sprott, J.C.; Wildenberg, J.C.; Azizi, Yousef (2005). "A simple spatiotemporal chaotic Lotka–
Volterra model". Chaos, Solitons & Fractals. Elsevier BV. 26 (4): 1035–1043.
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13. Wildenberg, J.C.; Vano, J.A.; Sprott, J.C. (2006). "Complex spatiotemporal dynamics in Lotka–
Volterra ring systems". Ecological Complexity. Elsevier BV. 3 (2): 140–147.
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University Press, Cambridge, U.K, p. 352.

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