Marc Naguib - The evolution of communication: signalling in a social network

Protocol by Paula Roloff-Dieck

Matrikelnr. 885694

Introduction
Everywhere on this planet, organisms communicate with each other. This communication
can be both intraspecific and interspecific. Over time, different mechanisms evolved - visual,
acoustic, chemical signals, for example. The acoustic communication is a topic which is
researched very often. Quite understandable, as for us humans, talking is the main
mechanism of communication.
Most animals developed sounds to understand each other. The sounds are used for
communication between members of a species for courtship or also territorial signalling.
There are certain signal traits that result in social interaction not only between two
individuals but between whole groups!
Birdsong, for example, has a strong effect on intraspecies interaction. Connections between
individuals form via spacial relations, but also via signals.
The song of birds often express motivation, quality and personality of a single bird. The birds
form territories next to each other, creating neighborhoods. Some individuals are more
tolerant than others, but in general, neighborhoods are beneficial because the birds do not
have to fight all the time. The territories are clear and the borders are accepted by all
neighbors. Thus, neighborhoods increase evolutionary fitness as well as affecting behaviour
when it comes to potential relationships.
In songbirds, the song functions both in male competition and in female attraction.
In the presented study three different systems in birds were observed to determine their
interactions and behaviour due to signalling in their social networks.

Nightingale
Nightingales migrate, but during breeding season, they are highly territorial.
Are there any differences in activity between different groups? Male and female nightingales
were caught to find out about their territory occupancy and mating status and their
movements were tracked. Observations showed that unmated males sing at night and
through the dawn, while mated males sing only at dawn. This is also the time when unmated
males move around most to find a new territory. This makes sense, as the peak of singing
activity from all males is the best time to prospect the area for information on territory
occupancy and mating status.
Females, on the other hand, move mainly at night when the unmated males have their peak
of singing activity, heightening their chance to find an unmated partner. Thus, both sexes
explore the social community at times when relevant information peaks
prospecting behaviour. But is this the cause or effect of signaling?
The "dialogues" between the nightingales could be described by several characteristics, or
singing assymetries:
⁃ the song rate, or how often singing would take place
⁃ song matching
⁃ song overlapping, or interrupting, of another bird's song
Different nightingale songs were recorded and played in the social community. If interrupted
by a playback, unmated males would be more affected (interruping themselves, stuttering)
than mated males, who tended to be more sure of themselves. Also, mated males interrupt
the playback more often.
Interestingly, the playbacks did not only affect the behaviour of the bird in whose territory it
was played. Males used their neighbor's response as a yard stick to estimate the degree of
danger: When there is an intruder in the community, neighbors will silently come closer to
see and assess how "dangerous" the challenger will be for them. Eavesdropping males
respond to songs stronger to overlaps and to leaders and do not discriminate when the
interaction is symmetric.
Personality traits also explain variation in territorial and spacial behaviour. Individual
nightingales were caught and put in a room. Shy ones stay on one branch (slow explorers),
while bold ones quickly examined the room (fast explorers).
Repeating the playback experiment after classifying slow and fast explorers showed that
slow explorers were more likely to leave their territory after a playback, and that the
neighbors reacted more strongly when the challenged male was a slow explorer. So, they
knew about the increased propability of the challenged male losing the confrontation and
leaving, thus letting a potentially fitter male into the territory.
Also, females were more likely to seek extra-pair copulation if their mate lost a vocal
challenge. They reassessed their mate's fitness.
So, both singnaling relations and spacial relations connect individuals. Together, they form
social interactions and networks.

The great tit

Unlike the nightingales, tits are resident. Im winter, they form social flocks, forming
territories but being quite tolerant at the same time.
One of the questions Marc Naguib and his team tried to answer was, "do territorial
intrusions affect neighborhoods in more subtle ways?"
To answer this, they radiotracked all grat tits of one forest simultaniously, observing who
was where and with whom at the same time, who moves a lot, who not, and trying to find
the correlation between social activity and boldness.
When recording the singing in the neighborhood, they found that neighbours sing at a more
dissimilar rate than non-neighbours (not in relation to space, just neighbours and non-
neighbours!), and that if an individual who sings not so often is challenged via playback, all
the neighbours start singing more often.
Strong vocal responses repel neighbouring males, and attract neighboring females.

The zebra finch

Little is known about the signaling of zebra finches. Although they are model organisms, they
are little studied in the wild. It is known that males sing - quite softly - at nest boxes, and
more boldly at communal trees, but the finer social structure has not been researched yet.

Conclusion
It is important to understand that communication is always a complex topic including spatial
behaviour and social networks. Birdsong serves as a model to understand the evolution of
communication, and the complexity of it.