Chapter 4: Model

In this chapter I will describe the model in detail, taking the prerequisites formulated in previous
chapter as a starting point. The purpose of the model is to find out whether local interactions
between birds might lead to an increase in complexity of the used signal. In the search for global
properties based on local interactions, the model cannot be fixed beforehand; as the global
properties typically emerge only by running the simulation, the outcome cannot be anticipated.
Therefore, in the next chapter, we will investigate the behaviour of several versions of the model.
We will adjust the model in the search for a set of local interactions that can produce dialect
formation and the increase of complexity of the signal.

4.1 Environment

The environment is spatially modelled. It consists of a toroidal grid of 100x100 squares, each
representing a territory that can be occupied by an agent, to use it for reproduction. The territories
are giving equal chances to the offspring of all agents. Every territory provides equal mating
possibilities and food resources, leaving every agent with a fixed expected number of offspring per
generation.

The environment is kept as simple as possible, with no predefined barriers and no inhomogeneity
in the distribution of food in space. Barriers in space, as previously used in other dialect models
(Ellers & Slabbekoorn, 2003), will readily cause the used signals to drift apart. Here the focus will
be on social behaviour as a cause for dialect formation and signal complexity.

4.2 Agents

The agents in the model represent only male birds and reproduce asexually. They have an
unlimited memory to store the elements they hear from other songs during childhood. Their own
song will be composed from the memorised elements.

After birth, the agents go through a learning process ultimately leading to a crystallised song.
During this process, they will disperse from the territory of their parent to a new territory, which
they might occupy depending on their fitness. After settling on the territory, the agent will stay

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there until its death, which every generation might occur with a chance of 20%. As long as they do
not die, they will produce on average five offspring every generation.

In the next chapter, I will adjust this basic model and assume that the birds in addition have an
inheritable property that influences song acquisition. With this heritable property, we can measure
the selection pressure toward greater complexity of the signal. If the simulation influences the
average values for this property, a fitness advantage exists for having this property.

4.2.1 Song
The male bird’s song consists of a set of song elements. The order of the song elements in the
song does not matter and the song learning process determines the size. The song elements are
discrete and chosen from a set of 235 possible song elements. They are all equally similar, that is,
element x is just as similar to element y as to any element z.

The song learning process comprises two phases, one phase in which different elements are
acquired and stored in the memory and another phase in which the song is composed. During the
first phase, the young birds will disperse from their parent’s nest to a random territory within their
range. They travel over a maximum of 2 squares, vertically, horizontally or diagonally, ending up in
on of the 24 possible territories around their native territory.

In the first phase the birds memorise song elements with a certain probability per element per
teacher, which will decline with time. They learn the elements with a high probability from their
parent (chance = 0.8), as they will hear the most from them while stuck to the nest. They learn
considerably less from the Moore-neighbourhood of their parent (probability = 0.2) and will have
only little learning capacity left when listening to the Moore-neighbourhood
of the territory they disperse to (probability = 0.1). In the end they will have
constructed a list of known elements, their memory.

During the second phase the song is composed. In this phase the birds try
to compose a song that resembles the most frequent song in the
neighbourhood of the territory they want to settle on. If an element in the Figure 4.1:The Moore
template song is not in their memory, in their song it will be replaced by neighbourhood

another element. For example, when AB is the template and the birds has A and C in his memory,
his song will be AC. As long as they have all the elements required, they will copy the song
perfectly. However, as soon as they lack elements, they song will become different from the
template.

4.2.2 Improvisation
Due to the discrete nature of the song in our model, an explicit improvisation mechanism has to

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be implemented. The song of songbirds in nature changes as a result of errors during song learning.
Birds make small mistakes in the song copying process, which causes the songs in a population to
change gradually. In the model, the song elements cannot change gradually; they only can change
radically to another element. Where in nature copying mistakes will be small and therefore will be
hardly noticed, a changed element in the model will not be recognised anymore. Moreover, it is not
enough to put a new element in the memory in the case of a learning mistake, because it will not be
in any template that is to be copied by settlers and therefore will almost never be used.

Therefore, the birds in the model improvise on the song itself, after the song composition. The
improvisation is implemented as three different processes: lengthening, shortening and changing of
the song. 1% of the song elements will be replaced by another, arbitrary, element. With the same
chance an element is removed from the song or an element is added to the song. While the addition
chance accounts for the whole song, the deletion chance accounts for each element of the song. A
higher deletion chance prevents the song from getting any longer without selection. Moreover, a
deletion chance per whole song would stabilise the composing elements of longer songs, as the
chance of losing an element would then diminish with the song length. Song length might then be
selected for as a means of reaching mutational stability, blurring the result.

After the improvising stage, the song is crystallised and will not change anymore during the
bird’s life. The crystallised song will be used in the competition with other birds.

4.2.3 Fitness
As selection of the birds takes place in the first year of their lives and is decided on the basis of
their fitness, the fitness will only be calculated then. The fitness F is estimated by comparing their
crystallised song with the songs of the Moore-neighbourhood of the territory where the agent wants
to settle. The more equal their song is to the neighbours’ songs, the higher their fitness is. They get
the highest fitness if their song is exactly the same or when the neighbouring territory is empty:

# neighbours
{J ( x, i)}+ (8−# neighbours)
F ( x) = i = 0
8

size( song ∩ song )

x y
J ( x, y ) =
size( song ∪ song )
x y

The fitness depends on the similarity between the song of the settler and the song of each of the
neighbours of the vacant territory, which is measured by the Jaccard similarity index J (Lynch,
1996). This index is the union divided by the intersection of the two songs.

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 4.2.4 Competition
The fitness of a bird determines the chance of settling on the territory he disperses to. However,
fitness does not map directly to a survival or reproduction chance, as the bird will almost never be
the only one trying to conquer it. When he ends up on an already occupied territory, he will always
loose the competition with the resident male and die, no matter what his fitness is. When a bird
lands on an empty territory, he will enter the competition with the other agents that have landed on
the spot. The winner of the competition is chosen proportional to his fitness. The chance of being
chosen is the own fitness divided by the cumulative fitness of all the competitors. The success of a
bird then depends on two things: the amount of competitors and his fitness relative to the fitness of
the competitors.

Competition in this model is based only on cultural selection and does not select for genes. In
principle, all agents are born the same, which causes selection to be just on the position of the agent
in space. However, as the chance to learn from parents is much higher than the chance to learn from
neighbours, information is transferred vertically much more easily than horizontally.

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