Table Of ContentAnimal Behaviour: Evolution and Mechanisms
Peter Kappeler (Ed.)
Animal Behaviour:
Evolution and Mechanisms
123
Prof.Dr.PeterKappeler
UniversityofGöttingen
Dept.ofSociobiology/Anthropology
&CRCEvolutionofSocialBehavior
&GermanPrimateCenter
BehavioralEcology&SociobiologyUnit
Kellnerweg6
37077Göttingen
Germany
[email protected]
ISBN978-3-642-02623-2 e-ISBN978-3-642-02624-9
DOI10.1007/978-3-642-02624-9
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Preface
The study of animal behaviour has become one of the fastest growing bio-
logical disciplines in recent decades. This development can be easily in-
ferred, for example, from the steady increase in the total number of publi-
cations on any aspect of animal behaviour, in particular also in journals
with a more general readership (e.g. Nature, Proceedings of the Royal So-
ciety or Current Biology), the ever-increasing number of participants at in-
ternational conferences (e.g. IEC or ISBE), and from the growing numbers
of students choosing courses in this field. This development has several
causes, of which I find three particularly compelling. First, it is increas-
ingly being appreciated that behaviour is the crucial level at which an indi-
vidual’s genotype and phenotype interface with the environment. Recog-
nising behaviour as the main mechanism animals employ to ascertain their
homeostasis, growth, survival and reproduction therefore provides a deep
understanding of organismal integration and adaptation. Second, the aston-
ishing success of the study of animal behaviour also has importantly to do
with the intellectual flexibility and methodological inter-disciplinarity re-
quired for comprehensive analyses of behaviour. Today, students of behav-
iour are jacks-of-all-trades; importing, applying and improving methods
from many neighbouring disciplines, such as molecular genetics, physiol-
ogy or micro-electronics, as well as concepts and theories from less obvi-
ous sources, such as economics or sociology, for example. Finally, Charles
Darwin’s theory of natural and sexual selection provide the study of ani-
mal behaviour with a powerful and firm theoretical framework that many
closely-related disciplines (e.g. neurobiology) are lacking.
This increase in the number of studies published in a growing number of
ever more specialised journals and the application of new concepts, meth-
ods and technologies also has frustrating consequences, however. Except
perhaps for a few exceptional colleagues, no one today is really able to de-
velop and maintain an active research programme and to read all interest-
ing and important publications and books that appear every month. We are
increasingly forced to specialise and to restrict our attention to a few topics
or taxa, despite a much larger intellectual curiosity. Most readers with a
PhD will be familiar with these constraints set by increasing administrative
loads, constantly changing teaching obligations as well as new types of
VI Preface
expectations of our employers about our publication and grant acquisition
records. This tendency to specialise backfires, however, when new genera-
tions of students need to be introduced comprehensively to all aspects of
the study of animal behaviour, when they ask for background information
to pursue their own personal curiosity, or when they ask great questions in
a lecture course or seminar.
One solution to this problem is to identify a useful introductory text or
up-to-date review. In practice, however, textbooks tend to focus more on
general principles than on current research, and reviews are typically writ-
ten for specialists, and are, hence, of limited use if you want to keep
abreast the literature in a broader field. A collection of authoritative re-
views written by active leaders in their respective fields can fill this gap if
they specifically address a non-specialist readership (i.e. not the closest
peers), summarise and explain recent developments, and if they provide a
forward-looking perspective for interested students as well as their closer
colleagues. In 1978, John Krebs and Nick Davies began providing just this
type of guidance for the then latest and fastest-growing field of animal be-
haviour: behavioural ecology. Their subsequent four edited volumes have
informed and influenced several generations of students and academic
mentors alike by providing a useful and stimulating basis for graduate
seminars, a competent source of reference for non-specialists as well as a
source of inspiration for newcomers and experts alike.
The study of animal behaviour has made enormous leaps forward since
the publication of the last Krebs and Davies volume in 1997 (Blackwell).
Others have also sensed the void left by the non-continuation of their se-
ries (Danchin et al. 2008: ‘Behavioural Ecology’ Oxford University Press;
Westneat and Fox 2010: ‘Evolutionary Behavioral Ecology’ Oxford Uni-
versity Press), but the field has become so wide that no single volume can
do justice to the existing diversity of behavioural research projects any
more. A recent trend among volumes of this kind appears to have been the
concentration on a regional (i.e. either francophone or anglo-american) set
of peers and their work; perhaps because they share certain preferred for-
mats for teaching. The current volume attempts to fill a similar niche by
featuring the state of the art in the study of animal behaviour in central
continental Europe, where ethology has its deepest roots. In addition, be-
cause of space limitations, the contributions to this volume only represent
a subset of current major research topics, but cover all recent international
developments. For historical reasons, ethologist in German-speaking coun-
tries have mainly been interested in mechanisms, but the number of re-
searchers embracing ultimate questions has been growing steadily. The ti-
tle of this volume was therefore chosen to reflect this development.
However, all authors were requested to address both ultimate and proxi-
Preface VII
mate aspects in the presentation of their respective topics, and some chap-
ters also deal with one or both of Tinbergen’s other questions.
The contributions to this volume are organised into four broad sections.
Note, however, that several chapters would also fit comfortably under a
different header, as indicated by numerous cross-references among chap-
ters. Communication and cognition continue to be central topics in the
study of animal behaviour. In chapter 1, Martin Schaefer reviews evolu-
tionary and functional aspects of visual communication. He emphasises the
fact that visual signals not only play important roles in several functional
domains of animals, but also in the communication between plants and
animals. Using examples from both areas, he discusses models of signal
design and evolution, concluding that environmental and other ecological
factors need to be considered explicitly for a more comprehensive under-
standing of communication systems.
Claudia Fichtel and Marta Manser deal with vocal communication in
chapter 2. They focus on communication beyond the traditional sender-
receiver paradigm and argue forcefully that bystanders also perceive vocal
signals exchanged among members of social groups. This point is under-
scored by their review of empirical studies on audience effects and eaves-
dropping. Furthermore, group coordination provides a particular context,
where individuals have to address several or all members of their social
unit simultaneously, and where the traditional dyadic communication
model fails. This review should therefore inspire much exciting new re-
search on communication from a network perspective.
In chapter 3, Dustin Penn and Joachim Frommen address kin recogni-
tion as a functionally important aspect of social and sexual behaviour. Fol-
lowing a much-needed conceptual clarification of the main concepts, they
focus on the various mechanisms and signals involved in the recognition of
kin. They also discuss central theoretical aspects in the evolution of kin
recognition mechanisms as well as their genetic underpinnings and conse-
quences. Their contribution concludes with a forward-looking perspective,
identifying main problems and areas of kin recognition research requiring
further work.
In chapter 4, Mario Pahl, Jürgen Tautz and Shaowu Zhang use the hon-
eybee as a model system to illustrate the fascinating sensory and cognitive
abilities of animals with small nervous systems. They introduce the honey-
bee’s sensory world and summarise experimental work on their various
cognitive abilities, including categorisation, rule learning and context-
dependent learning. These sometimes stunning abilities provide instructive
examples of how domain-specific cognitive faculties are linked to the spe-
cific ecological and social challenges these social insects face.
VIII Preface
In the final chapter of this section, Kurt Kotrschal, Isabella Scheiber
and Katharina Hirschenhauser use the highly-structured societies of grey-
lag geese to provide a fascinating comparative perspective on social cogni-
tion. These geese show striking convergence in several fundamental fea-
tures of their social systems with many mammals, including a female-
bonded clan structure and elaborate patterns of mutual social support.
Mates paired for life form the basic social units in this species and they
exhibit stunning hormonal synchrony. They contribute to social stability
through a particular pattern of social support that is reminiscent of some
other vertebrate societies with individualised long-term relationships.
Contributions to the second part revolve around the two central prob-
lems facing members of animal societies: conflict and cooperation. In
chapter 6, Jürgen Heinze reviews sources of conflict and conflict resolu-
tion in social insect societies. Long regarded as harmonious superorgan-
isms, societies of eusocial animals have only recently been recognised as
harbouring internal strife as well. Using the queen-worker conflict about
sex allocation and the struggle for dominance as examples, Heinze illus-
trates the sources and nature of such conflicts and reviews the behavioural
mechanisms used to minimise or to resolve them. The important role
played by inclusive fitness considerations in these conflicts highlights the
interaction between ultimate and proximate factors in this context.
In chapter 7, Judith Korb turns to cooperation with social insect socie-
ties as an evolutionary puzzle. Her review illuminates how relatedness and
mechanisms that make cheating costly act in concert to minimise cheating
and to stabilise cooperation. She goes on to show that similar mechanisms
favour cooperation at other levels of biological organisation and advocates
the use of a multilevel selection approach to study this and other evolu-
tionary problems at all levels of the biological hierarchy.
The subsequent chapter by Redouan Bshary is concerned with another
extreme: cooperation among unrelated individuals. He introduces game
theory as a particularly powerful and biologically satisfying approach to
studying the problem of cooperation because it can explicitly deal with the
many contingencies of real life. Using the well-studied example of marine
cleaning mutualism between members of different species, he goes on to
show how this approach is both necessary and useful in explaining coop-
eration. The tragedy of the commons provides another example where this
approach might be fruitfully applied.
In chapter 9, Gerald Kerth looks at group-decision making as a particu-
lar example of group-level cooperation that is potentially hampered by in-
ter-individual conflict. How animals reconcile the potentially conflicting
demands of group performance and individual interests has been subject to
much recent modelling. A concise review of these models is combined
Preface IX
with the results of empirical studies, including Kerth’s studies of Bechstein
bat societies, to identify fundamental rules in animal groups of varying
complexity and composition. The specific suggestions for future research
in this field offered by Kerth will promote a more comprehensive approach
towards studying group-decisions by incorporating additional intrinsic and
social variables known to shape other aspects of social behaviour.
In the final contribution to this section, Fritz Trillmich reviews the scope
for cooperation and conflict in the context of parental care. Successful re-
production is often jeopardised by conflicts between mates, between par-
ents and offspring, and among siblings. Using primarily examples from
mammals, where some of these conflicts are particularly pronounced,
Trillmich uses Tinbergen’s four questions to examine the nature and
causes of these conflicts, the leeway for cooperation as well as the mecha-
nisms modulating parental care. His chapter provides yet another example
of the benefits of addressing a particular problem from all four perspec-
tives, demonstrating in this case how consideration of all levels contributes
to a much richer understanding of a complex problem.
The third part of this volume revolves around problems of sex and re-
production. Chapter 11 by Wolf Blanckenhorn provides a broad opening
perspective on these topics by outlining formal approaches to the study of
sexual and natural selection, both in the wild and in the laboratory. He em-
phasises the benefits of using standardised selection measures in achieving
a comprehensive picture of both proximate and ultimate explanations of a
particular phenomenon. His illustration of how to obtain these measures
from a variety of sources and how to use them in meta-analyses should fa-
cilitate and encourage a much wider use of this approach in the study of
behaviour.
Chapter 12 by Nils Anthes reminds most of us that sexual conflict and
other sexual behaviours are not limited to the all too familiar separate-sex
animals. In simultaneous hermaphrodites, male and female reproductive
functions reside within the same body, providing an interesting twist for
the study of sexual strategies. Anthes reviews a growing body of literature
on hermaphrodites from an array of taxa that reveals fascinating evidence
for the occurrence of mate choice, sexual conflict and post-copulatory se-
lection in these animals. How hermaphrodites reconcile their intrinsic con-
flicts between their male and female function provides another prime ex-
ample for organismal fine-tuning in face of fundamental problems, such as
successful reproduction.
Bart Kempenaers and Emmi Schlicht deal with an intuitively much more
familiar problem in chapter 13: the optimisation of reproductive success
through extra-pair behaviour in pair-living species. Focussing on a vast lit-
erature on birds, these authors first explore sources of variation in extra-
X Preface
pair behaviour among species and populations. As extra-pair copulations
in pair-living species constitute a puzzle for evolutionary biologists, they
examine the associated potential costs and benefits from the perspectives
of both sexes, identifying effects in the expected direction, i.e. males can
increase offspring number and females offspring quality through extra-pair
activities. Kempenaers and Schlicht also explore hitherto neglected aspects
of extra-pair behaviour by illuminating its effects on the strength of sexual
selection.
In chapter 14, Bernhard Kraus and Robin Moritz focus on the polyan-
drous mating behaviour of social hymenoptera. The mating of hymenop-
teran queens with multiple males is of fundamental importance for these
animals because of its genetic consequences for their societies where con-
flict and cooperation are so finely tuned to prevailing relatedness patterns.
The authors review all major hypotheses proposed to explain the evolution
of polyandry and subsequently explore the consequences of genetic varia-
tion for the social organisation of colonies. Using the honeybee as one of
the best-studied model systems for these problems, they illustrate how ge-
netic analyses have revolutionised our understanding of social insect socie-
ties.
In chapter 15, Jutta Schneider and Lutz Frommhage provide an addi-
tional challenge to conventional sex roles by dissecting the mating strate-
gies of male spiders that invest heavily in paternity enhancement with one
(or very few) female(s), rather than trying to maximise access to many fe-
males. A small group of spiders is characterised by such exceptional
males, and therefore provides an instructive example to reflect on tradi-
tional sex roles and any sex-specific rules that have emerged from studies
of more ‘normal’ animals. The authors outline the fascinating diversity of
mating strategies among spiders and explain how aspects of their genital
morphology ultimately influence female (cannibalism) and male (monog-
yny) reproductive strategies.
In the last chapter of this part, Wolfgang Goymann and Heribert Hofer
use the theme of this volume to examine the relationships and interactions
among mating systems, social behaviour and hormones. They show that
testosterone is an important proximate factor involved in the regulation of
different mating systems across species and that it also influences individ-
ual mating decisions in some species. Additional hormones (oxytocin and
arginine vasopressin) are involved in pair-bond formation and the expres-
sion of different mating systems; in one particularly well-studied case,
changes in one hormone receptor gene have been shown to elicit a cascade
of massive changes in social behaviour. A key conclusion of this chapter
resounds the title of this volume: proximate and ultimate aspects of a par-
ticular behavioural phenomenon are best studied in combination.
