Kunstformen der Natur, Ernest Haeckel
Publications
12. Flack, J. C. & de Waal, F.B.M. Context modulates signal meaning in primate communication. PNAS- Track 2, 104, 1581-1586 (2007).
A central issue in the evolution of social
complexity and the evolution of communication concerns the
capacity to communicate about increasingly abstract objects
and concepts. Many animals can communicate about immediate
behavior, but to date, none have been reported to
communicate about behavior during future interactions. In
this study, we show that a special, unidirectional,
cost-free dominance-related signal used by monkeys
(pigtailed macaques: Macaca
nemestrina) means
submission (immediate behavior) or subordination (pattern
of behavior) depending on the context of usage. We
hypothesize that to decrease receiver uncertainty that the
signal object is subordination, senders shift contextual
usage from the conflict context, where the signal evolved,
to a peaceful one, in which submission is unwarranted. We
predict and find that deceasing receiver uncertainty
through peaceful signal exchange facilitates the
development of higher quality social relationships:
Individuals exchanging the peaceful variant groom and
reconcile more frequently and fight less frequently than
individuals exchanging signals only in the conflict context
or no signals. We rule out alternative hypotheses,
including an underlying reciprocity rule, temperament, and
proximity effects. Our results suggest that primates can
communicate about behavioral patterns when these concern
relationship rules. The invention of signals decreasing
uncertainty about relationship state is likely to have been
critical for the evolution of social complexity and to the
emergence of robust power structures that feed down to
influence rapidly changing individual
behavior.
11. Flack, J.C. & Krakauer, D.C.
Encoding power in communication
networks.
American Naturalist,
168, 97-102, 2006.
In animal societies conflicts can be
resolved by combatants or through third-party intervention.
In gregarious species, conflicts among pairs can spread to
involve multiple individuals. In the case of large
conflicts, containment and termination of aggression by
third parties is important. Successful intervention relies
on consensus among combatants about the intervener’s
capacity to use force. We refer to this consensus as power.
We measure it and study how it arises, using as our model
system a pigtailed macaque (Macaca nemestrina)
society. In macaques, the
degree to which one individual perceives another as capable
of using force is communicated using a special dominance
signal. Group consensus about an individual’s
capacity to use force arises from the network of signaling
interactions. We derive a formalism to quantify consensus
in the network. We find that the power distribution is fat
tailed and power is a strong predictor of social variables
including request for support, intervention cost, and
intensity. We develop models to show how dominance
signaling strategies promote robust power distributions
despite individual signaling errors. We suggest that when
considering correlated interactions among many individuals
it can be more useful to emphasize coarse-grained
information stored at the group level— behavioral
macrostates—over detailed information at the
individual level.
10. Ay, N., Flack, J.C., & Krakauer, D.C.
Convergent complexity and robustness in
multimodal signaling networks.
Philosophical Transactions of the Royal Society, Series
B,
362, 441-447 (2007).
In animal communication, signals are
frequently emitted using different channels (e.g.
frequencies in a vocalization) and different modalities
(e.g. gestures can accompany vocalizations). We explore two
explanations that have been provided for multimodality: (i)
selection for high information transfer through dedicated
channels and (ii) increasing fault tolerance or robustness
through multichannel signals. Robustness relates to an
accurate decoding of a signal when parts of a signal are
occluded. We show analytically in simple feed-forward
neural networks that while a multichannel signal can solve
the robustness problem, a multimodal signal does so more
effectively because it can maximize the contribution made
by each channelwhile minimizing the effects of exclusion.
Multimodality refers to sets of channels where within each
set information is highly correlated.We show that the
robustness property ensures correlations among channels
producing complex, associative networks as a by-product.We
refer to this as the principle of robust overdesign.We
discuss the biological implications of this for the
evolution of combinatorial signalling systems; in
particular, how robustness promotes enough redundancy to
allow for a subsequent specialization of redundant
components into novel signals.
9. Flack, J. C., Girvan, M., de Waal, F. B. M. &
Krakauer, D. C.
Policing stabilizes construction of social
niches in primates.
Nature,
439, 426-429. (2006). [Article]
Supplementary Material
All organisms interact with their
environment, and in doing so shape it, modifying resource
availability. Termed niche construction, this process has
been studied primarily at the ecological level with an
emphasis on the consequences of construction across
generations1. We
focus on the behavioural process of construction within a
single generation, identifying the role a robustness
mechanism2—conflict management—has in
promoting interactions that build social resource networks
or social niches. Using ‘knockout’ experiments
on a large, captive group of pigtailed macaques (Macaca
nemestrina), we show that a policing function, performed
infrequently by a small subset of
individuals3,
significantly contributes to maintaining stable resource
networks in the face of chronic perturbations that arise
through conflict. When policing is absent, social niches
destabilize, with group members building smaller, less
diverse, and less integrated grooming, play, proximity and
contact-sitting networks. Instability is quantified in
terms of reduced mean degree, increased clustering, reduced
reach, and increased assortativity. Policing not only
controls conflict3–5, we find it significantly influences the
structure of networks that constitute essential social
resources in gregarious primate societies. The structure of
such networks plays a critical role in infant
survivorship6,
emergence and spread of cooperative
behaviour7,
social learning and cultural traditions8.
Commentary on niche paper: Carr, G.
The police are a bunch of
monkeys,
The Economist. January 26, 2006.
Commentary on niche paper: Laland, K.
Animal Behaviour: Old World Monkeys Build New
World Order,
Current Biology, 16: R291-R292.
Commentary on niche paper: Smaglik, P.
Making the paper.
Nature, 439.
8. Flack, J. C., Krakauer, D. C. & de Waal, F. B.
M.
Robustness mechanisms in primate societies: A
perturbation study.
Proceedings of the Royal Society of London, Series
B
272, 1091-1099 (2005).
Appendix
Conflict management mechanisms have a
direct, critical effect on system robustness because they
mitigate conflict intensity and help repair damaged
relationships. However, robustness mechanisms can also have
indirect effects on system integrity by facilitating
interactions among components.We explore the indirect role
that conflict management mechanisms play in the maintenance
of social system robustness, using a perturbation technique
to ‘knockout’ components responsible for
effective conflict management. We explore the effects of
knockout on pigtailed macaque (Macaca nemestrina) social
organization, using a captive group of 84 individuals. This
system is ideal in addressing this question because there
is heterogeneity in performance of conflict management.
Consequently, conflict managers can be easily removed
without disrupting other control structures.We find that
powerful conflict managers are essential in maintaining
social order for the benefit of all members of society.We
show that knockout of components responsible for conflict
management results in system destabilization by
significantly increasing mean levels of conflict and
aggression, decreasing socio-positive interaction and
decreasing the operation of repair
mechanisms.
7. Flack, J. C., de Waal, F. B. M. & Krakauer, D.
C.
Social structure, robustness, and policing
cost in a cognitively sophisticated
species.
American Naturalist,
165, E126-E139 (2005).
Conflict management is one of the primary
requirements for social complexity. Of the many forms of
conflict management, one of the rarest and most interesting
is third-party policing, or intervening impartially to
control conflict. Third-party policing should be hard to
evolve because policers personally pay a cost for
intervening, while the benefits are diffused over the whole
group. In this study we investigate the incidence and costs
of policing in a primate society. We report quantitative
evidence of non–kin policing in the nonhuman primate,
the pigtailed macaque. We find that policing is effective
at reducing the intensity of or terminating conflict when
performed by the most powerful individuals.We define a
measure, social power consensus, that predicts effective
low-cost interventions by powerful individuals and
ineffective, relatively costly interventions by low-power
individuals. Finally, we develop a simple probabilistic
model to explore whether the degree to which policing can
effectively reduce the societal cost of conflict is
dependent on variance in the distribution of power. Our
data and simple model suggest that third-party policing
effectiveness and cost are dependent on power structure and
might emerge only in societies with high variance in
power.
Commentary
on policing paper: Anderson, J. (2005).
Animal behavior: Pigtailed
police.
Current Biology,
15, R427-R429.
6. Flack, J. C., Jeannotte, L. A. & de Waal, F. B.
M.
Play signaling and the perception of social
rules by juvenile chimpanzees (Pan
troglodytes).
Journal of Comparative Psychology,
118, 149-159 (2004).
Prescriptive social rules are enforced
statistical regularities. The authors investigated whether
juvenile chimpanzees (Pan
troglodytes)
recognize and use enforced statistical regularities to
guide dyadic play behavior. They hypothesized (a) that
proximity of adults, especially mothers of younger play
partners, to play bouts will increase the play signaling of
older partners and (b) that when juvenile–juvenile
play bouts occur in proximity to adults, older partners
will play at a lower intensity than when no adults are
present. They found that older and younger partners
increase their play signaling in the presence of the
mothers of younger partners, particularly as the intensity
of play bouts increases. In contrast to their hypothesis,
older partners played more roughly when the mothers of
younger partners were in proximity. These results suggest
that juvenile chimpanzees increase play signaling to
prevent termination of the play bouts by mothers of younger
partners.
5. Flack, J.C. & de Waal, F.B.M. (2004). Dominance
style, social power, and conflict management in macaque
societies: A conceptual framework. In:
How Societies are Built: The Macaque
Model,
eds. B. Thierry, B. Chapais, W. Kaumanns, M. Singh.
Cambridge: Cambridge University Press.
4.
Flack, J.C. & de Waal, F.B.M. (2002). Darwinian
building blocks of morality in nonhuman primates. Special
Issue on Evolutionary Theory.Samfundsøkonomen,
4: 29-38.
3. Flack, J.C. & de Waal, F.B.M. (2000).
'Any animal whatever': Darwinian building
blocks of morality in monkeys and apes
(includes commentaries).
Journal of Consciousness Studies,
7 (1-2): 1-65.
To what degree has biology influenced and
shaped the development of moral systems? One way to
determine the extent to which human moral systems might be
the product of natural selection is to explore behaviour in
other species that is analogous and perhaps homologous to
our own. Many non-human primates, for example, have similar
methods to humans for resolving, managing, and preventing
conflicts of interests within their groups. Such methods,
which include reciprocity and food sharing, reconciliation,
consolation, conflict intervention, and mediation, are the
very building blocks of moral systems in that they are
based on and facilitate cohesion among individuals and
reflect a concerted effort by community members to find
shared solutions to social conflict. Furthermore, these
methods of resource distribution and conflict resolution
often require or make use of capacities for empathy,
sympathy, and sometimes even community concern. Non-human
primates in societies in which such mechanisms are present
may not be exactly moral beings, but they do show signs of
a sense of social regularity that — just like the
norms and rules underlying human moral conduct —
promotes a mutually satisfactory modus
vivendi.
2. Flack, J.C. & de Waal, F.B.M. (2000).
Being nice is not a building block of
morality.
Journal of Consciousness
Studies,
7 (1-2): 67-78.
1. Pimentel, D., Wilson, C., McCullum, C., Huang, R., Dwen,
P., Flack, J., Tran, Q., Saltman, T., & Cliff, B.
(1997). Economic and environmental benefits of
biodiversity.
BioScience,
47 (11): 747-756.
In
Prep
1.
Flack, J.C. The study of animal communication networks:
Looking forward.
2.
Boehm, C. & Flack, J.C. Evolutionary perspectives:
Power and dominance in primates. In:
The Social Psychology of Power,
eds. A. Guinote & T. Vescio.
3.
Krakauer, D.C., Page, K. & Flack, J.C. Behavioral
immunology through third-party intervention.
4. Flack, J.C. & Krakauer, D.C. Principles of robust
design in evolving systems.
5. Flack, J.C, Ay, N. & Krakauer, D.C. Causality in
adaptive systems: Conceptual & methodological issues.
6. Bertschinger, N., Olbrich, E., Ay, N., Flack, J.C. &
Krakauer, D.C. The robust inferential theory of
individuality.
7. Flack, J.C. Cooperation, power, and resource equity.
8. Flack, J.C. & Krakauer, D.C. Social niche
inheritance.
9. Flack, J.C. & Krakauer, D.C. Constructing a robust
small-world: The role of behavioral neutral networks and
multiple developmental periods.
10. Krakauer, D.C. & Flack, J.C. Dynamics of war and
peace in animal societies.
11. Flack, J.C. & Krakauer, D.C. Conflict network
motifs and the spread of aggression
12. Fuentes, M., Krakauer, D.C., & Flack, J.C. Semantic
complexity and social structure: Insights from genus
Macaca
Popular
Pieces
Flack,
J.C. Complex form evolving,
Santa Fe Institute Bulletin, Winter
2007.
Flack, J.C. & de Waal, F.B.M. ‘Any Animal
Whatever’: Darwinian building blocks of morality in
monkeys and apes.
Business Ethics Quarterly,
Society for Business Ethics, University of Virginia (2004).
[Reprint of the Flack & de Waal (2000) paper published
in JCS.]
Reviews
Review of “Developing a Social Psychology of Monkeys
and Apes” by John Chadwick-Jones.
Human Ethology Bulletin,
December, 1999.
Review of “Animal Social Complexity” edited by
Frans B.M. de Waal and Peter Tyack.
Times Higher Education Supplement,
June 18, 2004.
Review of “Lords and Lemurs: Mad Scientists, Kings
with Spears, and the Survival of Diversity in
Madagascar” by Alison Jolly,
Times Higher Education Supplement,
April 22, 2005.
Review of “The First Idea: How Symbols, Language, and
Intelligence Evolved from our Primate Ancestors,” by
Stanley Greenspan & Stuart Shanker,
Times Higher Education Supplement,
September 30, 2005.
Review
of
“The Dynamic Dance: Novocal Communication in
African Great Apes,” By Barbara King,
American Journal of Primatology,
66, 389-391.
Review of "Dictionary of Animal Behavior" by David
McFarland, Oxford University Press.
Times Higher Education Supplement,
November 17, 2006.
Review of "Self-organisation and Evolution of Social
Systems" edited by Charlotte Hemelrijk, Cambridge
University Press. Forthcoming.
