Avian Navigation - Pigeon Homing As A Paradigm

Hans G.
Wallraff
Avian Navigation: Pigeon Homing as a Paradigm
Hans G. Wallraff
Avian Navigation:
Pigeon Homing
as a Paradigm
With 98 Figures
Dr. Hans G. Wallraff

Max Planck Institute for Ornithology
82319 Seewiesen
Germany
e-mail: wallraff@orn.mpg.de
ISBN 3-540-22385-1 Springer-Verlag Berlin Heidelberg New York

Library of Congress Control Number: 2004110945
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Preface
Bird migration includes homing performances in almost global dimensions.

After having spent the winter in South Africa, a stork or a swallow returns in
spring to its old nest in a village in Denmark. A redstart, a flycatcher or a bobolink arrives in its former breeding territory after nocturnal flights covering
thousands of kilometres. In what way do the birds solve the problems of navigation behind these performances?
Although this book deals neither with bird migration nor with storks and
flycatchers, but focuses on a domestic animal, it nevertheless deals with a central problem implicated in migrations and other long-distance flights characterizing the life cycles of many avian species. It is well known that such birds
can find their way home to their breeding sites over hundreds of kilometres of
unfamiliar territory, but little is known about how they achieve this admirable
feat. Experiments dealing with that question in detail, with navigational strategies and with use or non-use of various environmental signals have, for good
methodological reasons, predominantly been conducted with carrier pigeons.
Thus, what actually can be reported is mainly our present knowledge about pigeon homing. However, there is little doubt, and there are findings suggesting,
that the results and conclusions obtained with pigeons can largely be generalized towards an understanding of the navigational capabilities of other birds
as well. It is not the pigeon itself as a particular bird, as a companion of man
over thousands of years, that is the focus of interest of this monograph, but
specifically its role as a paradigm for the study of navigation mechanisms
enabling a bird to find its familiar home site from distant unfamiliar areas.
The book summarizes the outcome of about half a century of research. A final solution to the homing problem has not yet been achieved during this time;
substantial questions still remain. However, it seems that now, after a long period of trial-and-error searches and accumulation of initially unexplainable
observations, at least the four basic categories of exploited input signals have
been identified: the sun and the geomagnetic field (both used to determine
compass directions), the visual landscape (used to determine the current position inside a familiar area around home), and atmospheric trace gases perceived by olfaction (used to determine the position in relation to home even
from far away in unfamiliar areas). Naturally, the last category was not among
the candidate input signals hypothetically envisaged from the beginning. It entered the arena unexpectedly and achieved its central position, against understandable scepticism, under the pressure of more and more compelling empir-
VI
Preface
ical findings. Presentation of these findings, discussion of their significance

and theoretical consequence, and a proper look at the atmosphere make up the
longest chapter of the book.
Other sections proved necessary in order to show that initially favoured
physical cues, especially the earths magnetic field or the sun, are apparently
not used by birds to determine their current position with respect to home, but
are nevertheless substantial components of the homing process as a whole.
Further, it was necessary to describe the pigeons homing behaviour in some
detail and to separate its navigationally relevant parts from concomitant peculiarities which must be considered for methodological reasons and whose
disregard may lead to misinterpretation of results.
The book tries to give a state-of-the-art survey for everyone interested, but
also tries to meet the demands of a critical readership being more or less familiar with the field. Considering the former group, the main text does not aim to
report comprehensively on all pigeon homing experiments conducted over
five decades and on all debates and controversies connected with them. I focus
on those findings and ideas that I consider sound and relevant and that formed
my current picture of the field. Being selective in the interest of keeping the
text sufficiently fluent and attaining comprehensible conclusions, however, implies the danger and the possible reproach that the presented picture has been
made coherent at the cost of objectiveness. Therefore, I did not simply ignore
findings and opinions that might confuse the picture, but deal with them in a
separate section (Sect. 7.8), in a chapter on research history (Chap. 11) and in
the Appendix at the end, in which selected particulars are discussed in some
detail.
Even readers who are not ready to accept my style of presentation, my arguments and conclusions may profit from the book, as it facilitates access to the
extensive literature from which everybody can derive his or her own view of a
field and a period of research that now possibly approaches its end phase. I
may be wrong, but I feel that the types of pigeon releases, field glass observations and statistical analyses, as initiated by G. Kramer and G.V.T. Matthews in
the 1950s and described in the following pages, have largely done their task. On
the one hand, these experiments probably revealed the most fundamental peculiarities of pigeon homing (albeit many particulars are not yet clarified),
while, on the other hand, future progress in solving the remaining most fundamental questions will require more sophisticated methods intruding deeper
into the environment and into the animals neural mechanisms of signal processing. This monograph attempts to provide the material basis from which
more refined research may commence.
The book has a personal touch insofar as it includes a retrospect of my own
research over a period of about 45 years, i.e. over almost the whole period during which pigeon homing has been intensely investigated. Naturally, my mode
of looking at the accumulated material agrees most properly with the mode in
which I have previously designed experiments aimed at achieving some understanding of the pigeons homing behaviour. Therefore, and because other
Preface
VII
corresponding data are often unavailable, many of the figures show results obtained by me and by colleagues with whom I worked. I thank all those who contributed to produce the related data files, in particular Silvano Benvenuti,
Augusto Fo, Jakob Kiepenheuer, Michael Neumann, Ulrich Sinsch, Andrea
Streng and my technical assistants Rainer Wahl and Karl Wielander. I remember most gratefully the productive collaborations I had over the years with
Floriano Papi and his associates in Pisa, whose outcomes are included in the
following pages. Last, but not least, I thank Floriano for his readiness to read a
draft of the whole manuscript and for his helpful advice resulting in a number
of improvements.
Seewiesen / Starnberg, May 2004
Hans G. Wallraff
Contents
1 Introduction .................................................................................................... 1
1.1 Avian Migration and Navigation .......................................................... 1
1.2 Why Investigate Domestic Homing Pigeons? ...................................... 2
1.3 On Terms and Definitions ..................................................................... 4
2 Observation Data Used to Investigate Pigeon Homing ............................... 7
2.1 Initial Orientation .................................................................................. 7
2.2 Homing Performance .......................................................................... 11
2.3 Homing Routes .................................................................................... 12
2.4 Recoveries ............................................................................................. 15
2.5 Cage Experiments ................................................................................ 16
3 Basic Features of Pigeon Homing ................................................................ 17
3.1 Homing of Inexperienced Pigeons ...................................................... 18
3.1.1 Peculiarities of Initial Orientation ........................................... 18
3.1.2 The Problem of Motivation ...................................................... 23
3.1.3 Limited Significance of Exercise Flights at Home .................. 26
3.2 Experience Gained by Homing Flights ............................................... 29
3.2.1 Number and Range of Previous Homing Flights .................... 30
3.2.2 Directions of Previous Homing Flights ................................... 33
3.2.3 Familiarity with the Release Site or Area ................................. 36
3.3 Temporal Variability ........................................................................... 38
3.3.1 Annual Periodicity .................................................................... 38
3.3.2 Aperiodic Fluctuations ............................................................. 41
3.4 Accuracy of Homeward Orientation ................................................... 42
3.4.1 Distance of Displacement ......................................................... 42
3.4.2 Geographical Variability .......................................................... 43
3.4.3 Distraction by Landscape Configurations ............................... 44
3.4.4 Stochastic Noise ........................................................................ 46
3.5 Spatial Range of Pigeon Homing ........................................................ 46
3.6 Multiple and Mobile Home Sites ......................................................... 47
3.7 Conclusions and Perspectives ............................................................. 48
3.7.1 Home-Related and Home-Independent Components
of Initial Orientation ................................................................. 48
3.7.2 Motivation, Selection and Homing Experience ...................... 50
3.7.3 Inaccuracy and Variability of Homing Orientation ............... 51
Contents
4 Potential Input Signals Exploitable for Home-Finding ............................. 53

4.1 Directional References ......................................................................... 53
4.2 Indicators of Position .......................................................................... 54
4.2.1 Motion-Bound Signals .............................................................. 54
4.2.2 Location-Bound Signals ........................................................... 54
5 The Role of the Sun ....................................................................................... 59
5.1 Effects of Shifting the Birds Circadian Clock .................................... 59
5.2 Linkage Between Sun Compass and Map ........................................... 64
5.3 Orientation Under Overcast Skies ...................................................... 66
6 The Role of the Geomagnetic Field .............................................................. 69
6.1 Effects of Artificial Magnetic Fields .................................................... 69
6.1.1 Magnets or Coils During Flight ................................................ 69
6.1.2 Magnetic Fields Before Release ................................................ 73
6.2 Geomagnetic Irregularities .................................................................. 77
6.2.1 Spatial Magnetic Anomalies ..................................................... 77
6.2.2 Temporal Magnetic Fluctuations ............................................. 78
6.3.1 The Interference with the Stress Issue ................................... 79
6.3.2 The Magnetic Compass Issue ................................................. 80
6.3.3 The Magnetic Map Issue ......................................................... 81
6.3.4 The Path Integration Issue ..................................................... 82
6.3.5 The Magnetic Sense Issue ....................................................... 82
6.3.6 Overall Conclusions and Outlook ............................................ 85
7 The Role of the Chemical Atmosphere ........................................................ 87
7.1 Effects of Olfactory Deprivation .......................................................... 87
7.1.1 Olfactory Nerve Section ............................................................ 88
7.1.2 Inactivation of the Olfactory Epithelium by Zinc Sulphate .... 92
7.1.3 Nasal Anaesthesia ..................................................................... 93
7.1.4 Occlusion of Nostrils ................................................................ 94
7.1.5 Insertion of Nasal Tubes ........................................................... 96
7.2 Dependence on the Air the Pigeons Breathe ...................................... 96
7.2.1 Removal of Trace Gases by Filtration ...................................... 96
7.2.2 Air from Different Environments ............................................ 99
7.3 Olfactory Misguidance ....................................................................... 101
7.3.1 Spatial Separation of Sites of Air-Smelling and Release ....... 101
7.3.2 Outward-Journey Detours ...................................................... 104
7.3.3 Natural Olfactory Misguidance .............................................. 105
7.4 Spatial Range and Variability
of Olfaction-Based Homing ............................................................... 107
7.4.1 Distance Range and Efficiency in Different Regions ............ 107
7.4.2 Temporal Variability .............................................................. 110
Contents
XI
7.5 Varying Home Site Conditions in Aviary Experiments ................... 111

7.5.1 Shielding, Deflecting and Reversing Winds .......................... 111
7.5.2 Applying Artificial Odours and Winds ................................. 118
7.5.3 A Digression to the Preferred Compass Direction (PCD) .... 119
7.6 Spatial Structures in the Chemical Atmosphere .............................. 120
7.6.1 A Working Hypothesis ........................................................... 120
7.6.2 Gradients in Ratios of Trace Gases ........................................ 123
7.6.3 Applicability of Atmospheric Gradients to Navigation ........ 128
7.6.4 Assumed Dependence on Geographical Peculiarities .......... 131
7.6.5 The Short Distance/Long Distance Problem ......................... 133
7.7 Avian Olfactory Perception: A Suitable Tool for Navigation? ......... 136
7.7.1 The Problem of Sensitivity ..................................................... 136
7.7.2 The Problem of Odour Discrimination ................................. 137
7.7.3 The Problem of Compound Selection ................................... 137
7.7.4 The Problem of Adaptation/Habituation .............................. 138
7.8 Arguments Against Olfactory Navigation and Replies ................. 138
7.8.1 Intuitive Incredibility .............................................................. 138
7.8.2 Potential Non-Specific Side Effects ........................................ 139
7.8.3 Inconsistent Results ................................................................ 140
7.9 Conclusions and Perspectives ........................................................... 143
7.9.1 Experimental Evidence of Olfactory Navigation ................... 143
7.9.2 How Does Olfactory Navigation Operate? ............................. 144
7.9.3 Do Pigeons Have an Olfactory Map? ..................................... 144
7.9.4 Unsolved Problems Remaining Challenges ....................... 146
8 The Role of the Visual Landscape............................................................... 149
8.1 The Landscape as a Home-Guiding Factor ...................................... 149
8.1.1 Non-Olfactory Homing .......................................................... 149
8.1.2 Downgrading of Visual Signals .............................................. 152
8.1.3 Homing with Olfaction and Vision Impaired ....................... 153
8.1.4 Interrelations Between Landscape and Sun Compass .......... 155
8.2 The Landscape as a Disturbing Factor .............................................. 157
9 The Neural Bases of Pigeon Homing ......................................................... 161
9.1 The Hippocampus and the Familiar-Landscape Map ..................... 161
9.2 The Hippocampus and the Olfactory Map ....................................... 165
9.3 The Piriform Cortex and the Olfactory Map .................................... 166
9.4 Roles of Other Brain Regions and Hemispheric Lateralization ...... 167
10 Homing in Other Birds ............................................................................... 169
10.1 Experimental Approaches ................................................................. 169
10.1.1 Brief Overview ......................................................................... 169
XII
Contents
10.1.2 Compass Orientation and Preferred Compass Directions ... 171

10.1.3 Goal-Oriented Navigation ...................................................... 173
10.2 Homing in Natural Life ...................................................................... 176
10.2.1 Non-Migratory Excursions .................................................... 177
10.2.2 Homing as a Constituent of Bird Migration .......................... 179
11 Research History: Blind Alleys and an Unexpected Passage ................... 183
12 Overall Synthesis and Perspective ............................................................. 187
12.1 Environmental Cues Involved in Home-Finding Processes ............ 187
12.2 Two Homing Mechanisms ................................................................. 188
12.3 Challenges for Future Research ......................................................... 189
12.3.1 Unsettled Basics of Pigeon Homing ....................................... 189
12.3.2 Problems of Visual Landscape Orientation ........................... 190
12.3.3 Problems of Olfactory Navigation ......................................... 191
12.4 Outlook ............................................................................................... 192
Appendix: Notes on Selected Particulars ........................................................ 195
References ......................................................................................................... 205
Subject Index .................................................................................................... 225
Introduction
1.1
Avian Migration and Navigation
Among other aspects of bird migration, such as aerodynamics, energetics and
associated migratory strategies, including remarkable ecological adaptations,
the aspect of spatial orientation is certainly the most striking one. Consequently,
migratory orientation has been investigated in great detail.It is known that birds
can determine compass directions by means of the geomagnetic field, the stellar
sky or the sun, which are globally available as directional references. Using their
compasses, the birds can select genetically encoded directions which lead them
to their winter quarters. An endogenous time programme gives the appropriate
distance of flight in a temporal scale. Experimental data suggest that such bearing-and-distance programmes, which are roughly adapted to, and modified by,
topographical and ecological conditions of the covered regions, and which in
some species include even directional changes, are the general bases of many or
most population-specific migratory routes. Applying in this way its endogenous
intentions to the external world, even a young bird, leaving its birth place for the
first time, eventually arrives in the far distant wintering range of its population.
With larger birds,such as waterfowl or storks,which migrate in flocks during the
daytime, also guidance of juveniles by adults along traditional routes is involved.
Such guidance can be excluded in most of the smaller birds, particularly in the
many passerine species migrating predominantly at night. Detailed reports on
bird migration and migratory orientation can be found in other books (see
Alerstam 1990; Berthold 1996, 2000, 2001; Berthold et al. 2003).
In spring, the bearing-and-distance programme can be expected to operate
analogously to its operation in autumn; only the directions need to be reversed. A bird may return thereby to its population-specific breeding range. It
is hardly imaginable, however, that a redstart can reach, on this basis alone, the
village and the garden in which it had bred the year before. Migrating at night
and exposed to drift-causing winds, simply keeping the body axis on a genetically encoded compass course would be insufficient. In fact, the birds can perform better. It has been shown that birds of many species (e.g. starlings, cowbirds, blackbirds, swallows, swifts, terns, gulls, storks, shearwaters, petrels and
albatrosses) are able to find a familiar home site by goal-related navigation
from far distant unfamiliar areas to which they had been passively displaced
(for lists of such experiments and for references see Matthews 1955; Wiltschko
1 Introduction
1992; kesson 2003). To achieve this performance, the animals must have more
than a compass. Only if a bird is aware that it is north (and not south, east or
west) of home, can it usefully apply a compass in order to steer a southward
course guiding it home.
The process of birds finding their way back from a position in a distant unfamiliar area (where the animals have never been before) to a familiar geographical position (which has been established, during longer-lasting previous presence, as a home site) has often been called navigation in a narrow sense
(Sect. 1.3). There is hardly any doubt that birds commonly apply goal-related
navigation in the course of their migrations (Sect. 10.2.2) and that, without this
capability, bird migration patterns would never have achieved their present
overall character. Nevertheless, as substantial parts of migratory orientation,
homing mechanisms are usually not closely inspected in books and articles
dealing with bird migration. In fact, these mechanisms have not been closely
investigated in migratory species. However, they have been intensely investigated in a non-migratory domesticated bird. I feel confident that the knowledge gained from pigeon homing can largely be considered as knowledge
about avian homing in general and hence also of goal-finding in the course of
bird migration.
1.2
Why Investigate Domestic Homing Pigeons?
In research on avian navigation, the homing pigeon acts as a kind of laboratory
rat. Without the experiments conducted with this domestic animal, only a
small fraction of our present knowledge about the mechanisms used by birds
to find their way home from remote areas would have been achieved. The advantages of homing pigeons, as compared to any wild species, are obvious.
Hundreds or even thousands of pigeons can be kept in man-made lofts at
man-selected locations. Over the whole year the birds are easily available for
experimental use without laborious and stressful trapping procedures. Individual returns from releases can be precisely recorded, usually by only one observer. Homing pigeons are always motivated to return to their loft, not only
during the short breeding season. Due to their strong homing drive, they fly off
mostly immediately after release, thus giving opportunity to observe their initially selected courses, whereas most other birds tend to land on nearby trees
or in meadows etc. before starting homeward movements some time later.
Over short and median distances, many pigeons return in a non-stop flight, so
that the time they need for homing reflects, by and large, the length of the route
they have flown. Even many of the pigeons that fail to home provide information about their orientation: Due to their social impulses, they tend to enter
one or another of the pigeon lofts which are, in some countries, quite densely
scattered over the land. Reports by pigeon fanciers contribute instructive spatial distributions of recovery sites. Finally, pigeons are very handy, their size is
neither too small nor too large. They are large enough to be observed with bin-
1.2 Why Investigate Domestic Homing Pigeons?
oculars over approximately 2 km and large enough to carry the weight of a

small transmitter for telemetry or some other apparatus apt to record their
flight paths. They are small and tame enough to be confined in large numbers
together in manageable crates for transport.
These outstanding advantages resulted in in-depth investigations on homing capabilities of birds being almost exclusively conducted with homing pigeons. Thus, the question arises whether pigeon homing can be seen as a
model case of bird homing in general. At least there are good reasons to assume that it can. It appears extremely unlikely that, in the course of domestication from the rock pigeon (Columba livia) and selective breeding, an entirely
novel navigation mechanism that did not exist before in the genus Columba,
has been implanted into the strains of homing pigeons. The fact that the rock
pigeon is usually sedentary implies no argument against the assumption that
navigation mechanisms detected in homing pigeons can indicate mechanisms
used by other birds during migration. Closely related species (e.g. Columba
palumbus) do migrate, and Berthold et al. (1990) have shown that offspring of
partially migratory birds of a given species can be genetically transformed, by
means of selective breeding, to either full migrants or non-migrants within
only a few generations. It is uncertain whether selection performed by man
noteworthily affected the navigational capabilities of pigeons. Much more obvious is improvement of the homing drive, which appears less highly developed in the rock pigeon (Alleva et al. 1975; Visalberghi et al. 1978) and in other
wild birds (e.g. Able et al. 1984). In pigeon races, the breeder is concerned with
getting a maximum number of pigeons homed at maximum speed. The fastest
homers are then preferentially selected for breeding. Homing speed in such
races, however, is not primarily a result of outstanding navigation but rather of
motivation, physical strength and persistence. The birds are usually trained to
return in large flocks from always the same direction. Thus, the ability to find
their way home from novel directions and sites by making individual decisions
is not a criterion for selection.
Of course, there may be differences among pigeons and other species in
many details of homing behaviour, for instance in the motivation to fly home
at all, in the hierarchy in which different navigational cues are used or in the effectiveness with which such cues are exploited. It seems most likely, however,
that once we understand pigeon homing we are able to understand the principles of avian goal-oriented navigation in general. Nevertheless, this assumption remains merely an assumption unless it can be actually shown that at least
the most substantial conclusions drawn from experiments with homing pigeons can in fact be generalized and are valid for other bird species as well.
Some supporting evidence is already available (Chap. 10).
Although the carrier pigeon is a unique subject for research on animal homing, the huge amount of data obtainable from millions of homing flights performed in the course of pigeon races has only a minor value as a source of information in the context of this research. As mentioned above, races take place
in large flocks, usually comprising thousands of birds, so that, in a statistical
1 Introduction
sense, an individual bird is not an independent unit. Moreover, the pigeons of a

given region are usually displaced towards a standard compass direction with
distances increasing in the course of the season. Only homing speeds are measured. By multiple correlation analyses using such data (e.g. Dornfeldt 1996) it
is hardly possible to isolate causal relationships concerning navigational processes. To analyse these processes, it is necessary to conduct specifically designed experiments using individually flying pigeons released at sites in varying directions around home. Thus, researchers have no choice but to install
their own scientific pigeon lofts in order to gather suitable data.
1.3
On Terms and Definitions
Homing and navigation are the key terms used in this book. It is self-evident
that a precondition for homing is the creation of a home to which an animal is
willing to return from a distant site, whether it has moved to this site voluntarily or been displaced passively by man. Any adult bird usually makes its
breeding territory a home site. Domestic pigeons feel bound to a loft in which
they were living over several weeks postfledging (Sect. 3.1). Any act of return
movement to this site may be called homing, but not any such act is commonly
called navigation. If a pigeon or a starling returns from an accustomed feeding
area some hundred metres or even a few kilometres away, we may hardly speak
of navigation. However, if an albatross returns from a foraging flight covering
some thousand kilometres, its homing to a small island in the midst of the
ocean is undoubtedly an outcome of a remarkable navigational performance.
So what is navigation? I refrain from giving a sharp definition. Attempts to do
so, as attempted earlier, have never been successful. Usage of the term has even
been broadened in recent years (navigation through the brain, the Internet
etc.). Nevertheless, in the present context and in a narrower sense, the term
true navigation is frequently used for goal-related orientation towards a given
position on earth, usually a home site, towards which no direct sensory contact
is available at the current position of an animal. In its strictest sense, the term
is only applied if not even familiar landmarks surrounding the goal can be recognized, or are used, which potentially can guide to the goal. Such guidance by
familiar landmarks is often called piloting.
Another pair of terms to mention are compass and map. These are tools used
by human navigators and it is frequently implicated that animals also make use
of such instruments. In fact, applying these terms in the field of animal orientation and navigation is very convenient, as they describe particular functions
in a vivid and succinct way. While speaking about compasses and maps, however, we should be aware that we are using anthropomorphic metaphors. A flying bird might perhaps have an equivalent of a map exhibiting explored topographical features, but certainly it does not have a chart showing isolines of
physical quantities that might correspond to a grid of coordinates. However, an
animal navigator must have some representations in the brain referring to spa-
1.3 On Terms and Definitions
tial structures in the environment and must evaluate them in a way so that the
outcome is comparable to the outcome of our use of such instruments. Thus, in
order to avoid long-winded circumscriptions of neural processes which are
largely unknown, I shall use the terms compass and map as metaphoric shortcuts of such processes. As I do so with some reluctance, however, the terms
should always be thought of as between quotation marks, although they are
mostly not labelled that way because of their frequent occurrence.
Further words of caution concern the attributes often preceding the word
map. If I speak of a gradient map (Sect. 4.2.2), for instance, I do not presume
that a pigeon is in any way aware of spatial gradients or that such gradients are
mapped to some brain region. The term merely implies that the birds navigational system exploits scalar values of relevant environmental parameters that
increase or decrease more or less monotonically over some distance in a given
direction. The bird is thought to make use of gradients that could be drawn (by
man) on a map, but the bird is not thought to have such a map (Sect. 7.9.3).
The attributes connected with a map may belong to quite different categories.
They may refer to the physical or sensory nature of input signals (magnetic
map, olfactory map), to the spatial structure of evaluated cues (gradient map,
mosaic map, topographical map), to the application of true navigation as opposed to piloting (navigational map), to the spatial experience of an individual
bird (familiar area map) or to a psychological category (cognitive map). Concerning the latter, I do not take a position on whether or not pigeons have a
cognitive map (Sects. 4.2.2 and 9.1) (cf. Bennett 1996; Mackintosh 2002).
Generally, I have no ambition to get into debates on terminology. Classifications of terms and definitions rarely find unrestricted acceptance by all researchers working on different topics within a given field. Lines of demarcation separating natural phenomena from each other may appear appropriate
in one context but are often inappropriate when seen from another point of
view or when thus far unconsidered circumstances have to be included. By and
large, my notions are roughly, but not always literally, consistent with the terminological classifications as proposed by Papi (1992b) and Able (2000).
Observation Data Used

to Investigate Pigeon Homing
Many of the figures in this book show diagrams belonging to a small set of
graph types referring to a small set of data types from which conclusions on
the pigeons navigational performances and mechanisms are deduced. The
reader should first become familiar with these types of data. Illustrating them
by means of examples implies a kind of preview on the pigeons general homing behaviour which will be inspected in more detail in the following chapter.
An experiment on pigeon homing (in brief: a release) is usually conducted in
the following way. A number of pigeons, individually labelled by numbered
and coloured rings and selected according to age, homing experience, group
membership, etc., are caught in their home loft (often at night, when they are
sitting quietly) and confined in crates or baskets. In the morning they are
transported (mostly inside a car, prevented from viewing the outside) to the
release site where they remain sitting in the crates until each bird, after the
vanishing of its foreflier, has individually been released. A simultaneous release of different groups of pigeons means actually that members of the different groups are released alternately in succession. Release sites are, as far as possible, in open landscapes allowing unobstructed observation in all directions.
Each bird is tossed into the air and observed with binoculars until it vanishes
from sight. The time of release is recorded and another observer records the
individual birds time of arrival at the loft. Depending on the particular purpose and design of a given experiment, the procedural details are sometimes
modified in one way or another. The following criteria are used to determine
the pigeons performances.
2.1
Initial Orientation
The most often reported data are the vanishing bearings of individually flying
pigeons. At the time of vanishing, a pigeon is approximately 1.62.2 km distant
from the starting point, the exact distance depending on atmospheric conditions, background (sky or landscape), colour of the pigeon and quality of the
observer. From the individual bearings a mean vector can be calculated
(Batschelet 1981) which gives the mean direction (deviation clockwise from
north = N , from homeward = H ) and, with its length a, a reciprocal measure
of angular dispersion (Fig. 2.1). Length a is variable between 1 (all bearings in
one direction) and 0 (bearings uniformly distributed). Most important, partic-
2 Observation Data Used to Investigate Pigeon Homing
n Fig. 2.1. An example of initial bearings (peripheral dots) of individually released pigeons. The
direction towards the home loft is marked by a double-headed arrow. From the ten bearings, a
mean vector is calculated whose length a is indicated by the length of the central arrow (possible
maximum 1=radius of the circle) and whose direction is N , (deviation clockwise from north)
and H (deviation from home), respectively. The homeward component c H is the rectangular
projection onto the axis pointing towards home. Values in the example are: home = 135 from
north, N = 86 o , H = 49 o , a = 0.84 , c H = + 0.54. Note for the following circular diagrams (if not
otherwise stated): the double-headed arrow points always to the home loft. Peripheral symbols
mark the vanishing bearings of individual pigeons or mean bearings computed from a sample of
such primary bearings (in the case of second-order presentations; see Fig. 2.4, below). Mean vectors drawn as central arrows refer to a scale with value 1 equal to the radius of the circle
ularly when a number of symmetrically arranged releases are combined

and/or differently treated pigeons are to be compared, is the homeward component c H . It results from a rectangular projection of the mean vector onto the
axis running through release site and home site (c H = a cos H ). This component, variable between +1 and 1, depends on both direction and length of the
mean vector. It is positive if the deviation from home is less than 90.
Figure 2.2 shows examples of vanishing bearings of pigeons displaced from
six different lofts to the same release site over distances of 191345 km. The
birds had never before been displaced from their home (so-called first-flight
pigeons). It is obvious that those from the three northern lofts preferred northerly directions, whereas most birds coming from a southern home site vanished in the southern semicircle. It is also obvious, however, that the mean deviation from home as well as the angular dispersion of the bearings varied
greatly. Such patterns of initial orientation are very typical. The various factors
determining the shape of a particular pattern are discussed below. Low return
rates in first-fliers displaced over such long distances are also typical.
About a quarter of the initial flights are fairly straight, so that the pigeons vanish from sight within a minimally possible period of time of 12 min (Fig. 2.3).
More frequently, the birds first fly some loops or circles (see insert in Fig. 2.3)
and often they depart on a more or less meandering route. In rare cases, they remain more than 10 min within the range of visual observation.The vanishing in-
2.1 Initial Orientation
n Fig. 2.2. Vanishing bearings of inexperienced pigeons from six home lofts in Germany released at two neighbouring sites near Giessen. Percentage of birds returned is given inside the
circles. (Modified from Wallraff 1970a)
n Fig. 2.3. Vanishing intervals of 1,476 inexperienced (first-flight) pigeons from three Bavarian
lofts observed in the course of 189 releases. Median = 50% and other percentiles are indicated
along the top (unpublished summary from my data files). Insert shows a number of flight paths
tracked by visual triangulation from three observation points (modified from Spott 1993)
10
terval represents an indirect measure of the decidedness of initial orientation.

The median period of observation is around 3 min. These vanishing intervals
are quite variable among individual pigeons and also their means are variable
between releases. Experimental treatments, however, affect them rarely to a
larger degree. Therefore vanishing times play merely a subordinate role as indicators of orientational performances. They are routinely recorded in every release, but usually cannot be used as a criterion for substantial conclusions.
The directional distributions of the pigeons bearings at departure are much
more conclusive and worthy of being analysed in more detail. Most useful for
this purpose are so-called second-order analyses combining the results of a number of releases conducted at different sites. Figure 2.4 shows two kinds of second-order distributions. Each of the arrows in the upper graphs corresponds to
n Fig. 2.4. Initial orientation of inexperienced (first-flight) pigeons from seven lofts in Ger-
many, from each loft at four symmetrically distributed sites at distances of 85216 km. Direction
towards home points upwards. Bearings of departing birds were determined 20 and 40 s after release and at time of vanishing from sight. Arrows in the upper diagrams show first-order mean
vectors per release (vector length 1 = radius; cf. Fig. 2.1), whose second-order mean length (irre=
spective of direction) is given by a. Ellipses indicate the 95 and 99% confidence ranges of the second-order bivariate mean (considering lengths and directions), which itself is represented by the
centre of the ellipse and by the three numbers (angular deviation from home, vector length,
homeward component). At the periphery of the lower diagrams, directions of the same vectors
are shown, but now with equal weight independently of their lengths, i.e. of the angular scatter
within each release. Arrows and numbers give the second-order mean vectors calculated from
these 28 directions. (Data from Wallraff 1970a, supplemented by unpubl. data)
2.2 Homing Performance
11
the central arrows shown in Figs. 2.1 and 2.2, i.e. it represents the mean vector
calculated from a sample of bearings observed at one release site ( H , a; all
=
=
home directions upwards). The second-order mean vector ( H , aH) resulting
from the 28 first-order vectors is marked by the centre of the ellipses. In the bottom graphs, each first-order vector enters with equal weight irrespective of its
length. Thus, it serves as one bearing as indicated by the peripheral symbols. A
second-order vector of that kind (central arrows) is calculated analogously to
the first-order vector in Fig. 2.1. Such second-order analyses including a number
of release sites symmetrically distributed around the home loft are, for instance,
necessary to test whether pigeons fly consistently homewards from various distant positions. At a single site, a direction preferred by a sample of pigeons could
coincide with the homeward direction simply by chance.
Figure 2.4 not only illustrates a statistical procedure, but also includes a message. Pigeons do not need some time of flying around after release to gain information about the direction of home. As little as 20 s after take-off they tend to
prefer the semicircle including the homeward direction, though their bearings
after 20 and 40 s are usually more scattered than at the time of vanishing. Greater
angular scatter results in shorter mean vectors obtained from different samples
=
of birds (Fig. 2.4 top: a = mean vector length per release independently of direction). The directions of these vectors, however, independent of their lengths, are
well homeward oriented after 20 s as they are some 2 or 5 min later (Fig. 2.4, below). Obviously, the greater initial scatter is merely a technical outcome of the
fact that the birds need some time to gain height and to orient their flight from
an arbitrary starting course to an actively intended direction. Moreover, observed bearings are rarely identical with the birds current flight direction. At a
short distance, i.e. early after release, a flight path including a considerable lateral component with respect to the observers view causes a much greater angular displacement than a linearly equally long path at a distance of 12 km. Thus,
the pigeonsdirectional tendencies are less clearly recognizable at the beginning,
but the equal vector lengths and homeward components in the lower diagrams
of Fig. 2.4 suggest that the pigeons do not need to circle around in order to obtain information about their position. Otherwise, also mean directions would
initially be more scattered and less clearly homeward-oriented. More specific results later lead to the same conclusion (Sect. 7.2.1).
2.2
Homing Performance
Another easily obtainable criterion is homing performance, i.e. homing duration or speed and homing rate. Figure 2.5 shows an example, presenting the
same data in graphically different versions. Speed is expressed in units of
beeline distance per time (km/h: Fig. 2.5A, B) and duration in time per unit of
beeline distance (min/km: Fig. 2.5C, D). The histograms result in reversely
skewed distributions. In either version, there are usually two classes of pigeons
that do not fit to the scale. One class includes birds that homed very slowly,
12
n Fig. 2.5. Homing performances of 486 inexperienced pigeons in 37 releases over 2229 km distance in southern Bavaria. The same data are shown in four kinds of diagrams (unpubl. data)
mostly on the day after the release or later. The second class concerns pigeons
that never returned at all. With greater distances of displacement, and particularly with inexperienced pigeons, the majority of the birds may belong to these
two classes. In more extreme cases, only the return rate may be available as a
measure of homing performance.
In the following sections, graphs of the type in Fig. 2.5D illustrate homing
performance. Using this type, results obtained with different experimental
treatments can be optimally compared within one diagram; the median and
other percentiles of a sample can immediately be read (see abscissa values corresponding to ordinate values 25, 50 and 75% in Fig. 2.5D). As 1 min/km
(= 60 km/h) indicates the approximate flight speed of pigeons, the abscissa
roughly indicates the factor by which the routes flown exceed the beeline distance, provided that the birds did not interrupt their flight by some period(s)
of rest.
2.3
Homing Routes
Most of the presently available data do not include any information about the
routes the pigeons have flown between vanishing a few minutes after release
and arrival at the loft. In some experiments, however, the complete flight paths,
or parts of them, were observed by one of the following four methods.
2.3 Homing Routes
13
1. Observation from airplane or helicopter. Using a slowly flying airplane

(Hitchcock 1952) or a helicopter (Wagner 1970, 1972, 1974; Fiaschi et al.
1981), flocks of differently coloured pigeons could be visually observed
while flying over considerable distances. The method is hardly usable to follow individually flying birds and is too laborious and expensive to collect
data samples that are sufficiently large for statistical evaluations. Nevertheless, obvious responses to the local topography were observed (Sect. 3.4.3)
and altitudes of flight were found to be mostly less than 100 m, and often
less than 50 m, above ground.
2. Radio telemetry. Using ground-level receiving antennae, pigeons carrying
small radio transmitters can be followed over distances of some 1020 km
(e.g. Windsor 1975; Walcott 1977, 1978). With one receiver at the release
site, the range for vanishing bearings is thus correspondingly larger than
the visual range. Using a number of receivers distributed over an extended
area, flight paths can be determined by means of triangulation (SchmidtKoenig and Walcott 1978; see Fig. 8.3). Basically unlimited recording of
routes is possible when radio telemetry is combined with airplane tracking
(Michener and Walcott 1978). Owing to the considerable expenditure of
costs and time for each single bird, however, such techniques never found
broader application in pigeon homing research. In more recent years, a
much more elegant and efficient technique has been developed: that of satellite telemetry (e.g. Fig. 10.6). During its orbit around the earth, a satellite
determines the position of a transmitter from time to time. As the transmitters necessary for this purpose are still fairly heavy, they have so far been
used only with birds larger than pigeons. As miniaturization of devices will
probably go on, we can expect that in several years this technique will become a useful tool also in pigeon homing research. However, for many purposes the meanwhile developed position recorder (see point 4 below) may
be superior.
3. Direction recorder. Broader databases could be achieved by the development
of a miniaturized device that measures and stores, at regular intervals, the
compass direction of the pigeons body axis during the whole homing flight
(Bramanti et al. 1988; DallAntonia et al. 1992). In its newer version it weighs
only 13 g and hence can easily be carried by a pigeon. After the birds arrival
at home, the stored data can be extracted from the digital memory and flight
paths reconstructed. A source of possible error and uncertainty in such reconstructions is wind drift. Nevertheless, carefully conducted experiments
and evaluations have led to informative results, especially to findings illustrating topographical influences (e.g. Ioal et al. 1994; Bonadonna et al. 1997,
2000). For an example of routes recorded with this device see Fig. 7.13B.
4. Position recorder. A most promising new technique has recently been described by von Hnerbein et al. (2000) and Steiner et al. (2000). Birds carry a
recorder using the satellite-based global positioning system (GPS) which can
store highly accurate position fixes at intervals of seconds. Data can then be
downloaded and true flight paths plotted on a map (Fig. 2.6) and analysed. At
14
n Fig. 2.6. Examples of homing flight routes of pigeons

recorded by means of a GPS device. In this particular case,
deviations to the east from the direct route appear to be
typical. (Modified from von Hnerbein et al. 2000)
the time of writing, the weight of the device is 3335 g and hence at the limit of
being suitable for pigeons. For first applications of GPS tracking beyond
technical testing see Biro et al. (2002) and Guilford et al. (2004).
As useful as they are and probably will be, a disadvantage of the two last-mentioned techniques is the fact that the birds must be recaptured in order to gain
the recorded data and recover the fairly expensive device. Therefore, they are not
very well suited for risky experiments in which many pigeons fail to return to
their home loft. Just in the course of such experiments, however, which may, for
instance,test the influence of a particular experimental treatment,it would often
be particularly important to obtain information on the whereabouts of the
non-homers. For their localization, either satellite telemetry (point 2) or a combination with recoveries (next section) may become a suitable future method.
2.4
Recoveries
As regards observation of longer-distance movements, modern high-tech
methods are presently just at the point of replacing classical primitive sources
of information. Results shown in this book are still based on such sources.
Homing pigeons are social and hence attracted by conspecifics they meet on
their way. Thus, if they do not reach their home loft, they tend to enter another
pigeon loft. In some countries, there is a quite dense network of such lofts.
2.4 Recoveries
15
Many of the fanciers have their pigeons under daily control and detect any foreigner among them. If such a foreigner has a conspicuous address label around
its ring, possibly promising an award, the pigeon has a good chance of being
reported to its owner. In Germany, about half of the non-returning pigeons, if
labelled in such a way, have been reported (Wallraff 1989a). Recoveries show,
n Fig. 2.7. Vanishing bearings (circular diagrams) and recovery sites of pigeons from three lofts in
the north (H Hohenkirchen; W Wilhelmshaven; O Osnabrck) and one loft in the south (F
Freiburg) of a central release area near Giessen, Germany. Lines with arrowhead indicate pigeons
that homed after re-release at one or two recovery sites (a.rep after report). Most of the birds were
more or less experienced in homing over shorter distances (up to 45 km), their last previous flight
not being from the direction used here. (After Pratt and Wallraff 1958, from Kramer 1959b)
16
for instance, that little experienced pigeons, even if they never return to their
loft from a far-distant release site, have a pronounced tendency at least to approach their home and to reduce the distance from it (Fig. 2.7). However, only a
few birds were found on a straight route towards home. The recovery sites are
widely dispersed and sometimes they are biased towards a direction markedly
deviating from home. Thus, long-distance homeward orientation is far from
being precise.
2.5
Cage Experiments
Manifold attempts have been made to deduce information concerning the pigeons knowledge about its position with respect to home without or before allowing it to fly away. To get such information while the birds are confined in a
cage would be very advantageous because experimental conditions could be
manipulated much easier than for a bird in free flight and because the pigeons
could be prevented from exploring the area around. So far, however, investigations of caged birds in homing experiments have not attained the elucidating
role they have achieved in experiments on compass orientation using either
directional training or observations of migratory restlessness (e.g. Helbig
1991). Over the years, I operated five different types of orientation cages in the
context of homing, but never obtained results sufficiently satisfying to be published. Publications by others on successful experiments with caged pigeons
(Chelazzi and Pardi 1972; Kowalski 1994; Mazzotto et al. 1999) did not very
much exceed the level of technical reports and hence their contribution to our
present knowledge on the mechanisms involved in pigeon homing is limited.
Very recently, however, such cage or arena experiments have been more successful and have led to elucidating results (Gagliardo et al. 2001b; Diekamp et
al. 2002; Sect. 8.1.3).
Basic Features of Pigeon Homing
In a first approach, before coming to possible explanations, it is necessary to

convey some familiarity with the phenomena that have to be explained. It is
necessary to describe the kinds of available observation data and a number of
peculiarities that characterize the pigeons homing behaviour. In this descriptive part, we do not yet deal with experiments that aim to manipulate specific
environmental cues or associated processes of signal perception and evaluation, which potentially might be involved in home-finding. However, such a
distinction between description and experimentation is somewhat arbitrary
and does not follow a sharp borderline. In a strict sense, every man-made displacement of a pigeon from its loft to a remote site is an experiment conducted
under variable circumstances determined by home site and holding conditions, release site, season, weather, previous experiences of the birds etc. Some
conclusions about properties of mechanisms and factors involved can already
be drawn from such simple non-experimental experiments, which cannot be
clearly separated from the more specific experiments reported in later sections. Also, I shall sometimes refer to a later section, because the respective
phenomenon is more closely described there in an explanatory context.
This chapter deals with homing behaviour which embraces more than oriented activities leading a displaced pigeon as fast as possible back to its home
loft. It embraces motivation to home at all, factors distracting the birds from an
optimal course, and unexplained oriented activities that most likely are not
part of the home-finding process, but cannot easily be separated from it. Also,
it is necessary to deal with the fact that an individual pigeon is not an invariable subject that shows in equivalent situations reliably equivalent behaviour.
Rather, its behaviour in later homing flights is influenced by experiences the
bird has made in previous flights, so that its individual life history co-determines its homing behaviour. It is important to consider all these concomitant
circumstances when planning experiments and interpreting their results.
Readers primarily interested in navigational mechanisms, but not so much
in behavioural particularities and methodological difficulties, may skip the
following pages, may go directly to the concluding Section 3.7 and may possibly come back to one or another particular issue when referred to in a later
context.
18
3 Basic Features of Pigeon Homing
3.1
Homing of Inexperienced Pigeons
Pigeons displaced from their home loft for the first time (first-flight pigeons)
are best suited to the investigation of the primary abilities of homeward orientation from distant sites, because these inexperienced birds are not yet influenced by any learning processes that take place, or may take place, during
homing flights over remote territories (Sect. 3.2). In the experiments described
below, the pigeons were, if not otherwise stated, at least 4 months old. At this
age, several ontogenetic processes (e.g. use of olfactory map factors and sun
compass, decreased sensitivity to disturbing transport conditions) appear to
be completed (e.g. Wiltschko and Wiltschko 1981, 1985; Gagliardo et al. 1988,
2001a; see also Wallraff 2000b). In most cases, the first-fliers were not older
than 6 months. Within this range of 46 months, no age-dependent improvement of performances was apparent (Hoffmann 1959; Wallraff, unpubl. data).
There are some indications that pigeons displaced for the first time in their
second year of life (now paired, breeding and more firmly fixed to their loft)
tend to perform better than first-year birds (unpubl. data), but such experiments were rarely conducted, because homing pigeons are usually kept for
homing experiments and not simply to become older.
3.1.1
Peculiarities of Initial Orientation
Kramer (1957) observed that first-flight pigeons from a given loft do not home
equally well from everywhere. Initial orientation as well as homing success of
pigeons housed in Wilhelmshaven was much better from 95 km south than
from 88 km east, from where also recoveries were not generally homeward oriented (Fig. 3.1). Ten years later many more data were available, now from five
lofts in the same region of northwestern Germany and always from sites symmetrically distributed around home (Fig. 3.2). Their pooled vanishing bearings were directly oriented towards home on average, but were widely scattered around the circle. This overall pattern, however, does not reflect the
behaviour of birds of a particular loft at a particular release site. At a single site,
angular scatter is mostly much smaller (and thus, mean vector longer), but, on
the other hand, the mean direction differs mostly from the direct course towards home, often quite considerably and significantly. These marked deviations from the grand mean, specific for a particular home site/release site combination, can be read from the central vectors in Fig. 3.2, most of which are
fairly long but whose directions are widely scattered. Individual examples are
shown in Fig. 2.2. Other examples (Fig. 3.3) may demonstrate that such deviations are really site-specific: at some sites very consistently reproduced in repeated releases, at others at least within a wider sector.
When these very common release-site biases were first described (Kramer
1959b; Wallraff 1959b), but often also in later years (e.g. Keeton 1974a; Ganz-
3.1 Homing of Inexperienced Pigeons
19
n Fig. 3.1. Results of two simultaneous releases of first-flight pigeons displaced from Wilhelms-
haven (W.) in Germany to a site 95 km south (L.) or 88 km east (St.), respectively: vanishing bearings, recoveries and numbers of birds returned/released. 12 on the circular diagrams indicates
wind speed in Beaufort. (Kramer 1957)
horn 1990; Wiltschko 1993; Wiltschko and Wiltschko 1998), they were interpreted as systematic navigational errors and hence as a possible guide to the
map component in pigeon homing (Keeton 1973). The initial bearings were
seen as the directions in which the birds expect their home loft. However, empirical findings raise doubts whether this seemingly reasonable conclusion is
justified.
Figure 3.4 shows, for pigeons from a loft near Wrzburg in Germany, mean
vanishing bearings observed at 36 centrally symmetrical sites at distances of
30300 km. In general, the arrows are most clearly homeward-oriented at sites
southeast and east of home. In the other areas they tend to be shorter and their
directions are more variable. When the mean vectors or directions are summarized, both a general tendency towards home and a generally preferred compass direction become obvious (large circular diagrams in Fig. 3.4). Either directional preference is highly significant and expressed with similar strength.
These results tell us that releases at a single site, or at several arbitrarily selected sites, would not give reliable information on the birds ability to orient
homeward. At a southeastern site, for instance, the pigeons would appear perfectly homeward-oriented. They are, however, also perfectly oriented towards
their generally preferred compass direction, so that it is impossible to separate
20
n Fig. 3.2. Summary of vanishing bearings of 578 pigeons from 5 lofts in northern Germany at
their very first release. Bearings are pooled with home pointing upwards. Outer circle indicates a
uniform distribution, i.e. 578 birds/24 sectors = 24.1 birds per sector. Mean vectors in the centre
(length 1 = radius of the median circle) refer to the 20 samples as indicated on the map (distances
towards home 85165 km). Radius of innermost circle and length of inner arrow represent a = 0.29
which is the length of the mean vector of the total; its direction points exactly towards home. Positions of release sites and home sites are shown together with the respective numbers of birds.
Home sites: Ho Hohenkirchen; Wi Wilhelmshaven; No Nordenham; Os Osnabrck (two lofts, 4 km
distant from each other). (Wallraff 1967)
the two directional tendencies. On the other hand, if the pigeons vanish greatly
scattered in westerly directions at a northwestern site, we must not rashly conclude that they erroneously suppose to be somewhere east of home. The ten-
21
n Fig. 3.3. Examples of site-specific initial orientation of inexperienced pigeons. Each central ar-
row represents the mean vector resulting from one release per month regularly distributed over a
year (710 vanishing bearings per release). The centres of the 95+99% confidence ellipses correspond to the end points of second-order mean vectors resulting from 12 single vectors; their numerical values are given inside the circles (angular deviation from home/vector length/homeward
component). On the periphery, mean directions per release are repeated and the second-order
mean vectors resulting from them (neglecting first-order vector length) are given numerically.
Filled symbols refer to AprilOctober, open symbols to NovemberMarch. A Home loft at Andechs
near Munich, Germany; B home loft at Ospedaletto near Pisa, Italy. (Data from Fo et al. 1984)
dency to fly westward may have been stronger than the tendency to fly homeward. A decision on whether or not the pigeons vanishing bearings reveal
some knowledge about the direction of home at all can only be achieved by
summarizing a number of releases conducted at symmetrically distributed
sites (Fig. 3.4, diagram Home).
It is obvious that a loft-specific preferred compass direction (PCD), as resulting
from the data in Fig. 3.4 (upper right diagram), is very often one of the reasons
leading to a release-site bias (e.g. Schmidt-Koenig 1963a, 1970; Wallraff 1967,
1970a, 1978a, 1986, 1991a; Windsor 1975; Ioal 1995). At a given release site, pigeons tend to prefer a direction somewhere between PCD, which is specific for a
given loft site, and home. A PCD is not equally well expressed around all home
sites and varies in dependence on loft conditions and homing experience
(Fig. 3.9). There are other reasons for release-site biases as well (Sects. 3.4.3,
7.3.3, 7.6.3, 8.2) and it is hardly possible to separate the various potential influences contributing to making a mean bearing vector that characterizes a given
22
n Fig. 3.4. Initial orientation (mean vectors from vanishing bearings) of first-flight pigeons at 36
sites around Wrzburg. Arrows show mean vectors from all vanishing bearings per site
(n=2588; radius of small circles corresponds to a = 0.1. In the large diagrams, these vectors and
their directions are arranged according to their deviation from home and from north, respectively. Ellipses and numerical values (second-order direction and length) are analogous to those
in Fig. 3.3. The means from the total of 1,495 bearings are also given. Asterisks indicate level of
significance: ** P<0.01, *** P<0.001. (Data from Wallraff 1986, supplemented by additional data)
release site. Real navigational errors are most probably among the factors involved, but in most cases they cannot be clearly recognized (Sect. 3.7.1).
It should be emphasized that a simple arrow alone (as shown in Fig. 3.4) does
not illustrate the whole peculiarity of initial orientation at a given site. The
mean vector usually does not represent the mean of a circular normal distribution, but a computed direction which itself has not always been preferred. The
shorter arrows, in particular, mostly represent a multimodal directional pattern which is typical for the respective site (as an example see Fig. 8.6; for more
such patterns, see Fig. 8 in Wallraff 1959b).
23
3.1.2
The Problem of Motivation
A pigeon released at a remote site cannot be forced to fly as fast and directly as
possible homeward. Thus, the observed performances of the birds need not,
and mostly do not, immediately indicate the potentially achievable level of
performance. It is always difficult to distinguish between capability and motivation to make use of it.
Homing success of first-flight pigeons decreases considerably with increasing distance of displacement. From beyond 100 km, only a small minority returned to the loft at Wrzburg (Fig. 3.5A). Obviously, however, this decrease
was not due to a decreasing ability to determine the direction towards home. In
the same set of releases, initial homeward orientation even showed a slight tendency to improve over distance. Recoveries reveal that most of the non-returners were well on the way towards home, but stopped flying before they had
reached the loft (see below, Figs. 3.63.8). Thus, low homing rates over long distances appear to indicate limited persistence in flying and searching for home
rather than lowered navigational ability. With their usual homing velocity (e.g.
Fig. 2.5), inexperienced pigeons released more than 100 km distant have
hardly a chance to reach their loft during the day of release. Therefore, they
must look for shelter for the night. Many birds certainly find it in another pigeon loft, may perhaps stay there and finally lose their original homing drive.
Most of the first-flight pigeons we used were born in the same year, had not yet
bred in their loft and hence were probably not yet as firmly bound to this place
as older birds are after several years.
Figure 3.5B illustrates that there is a difference between individual pigeons.
Over shorter distances, the majority of the pigeons (64%) returned to the loft,
but on average even these homers were weakly homeward oriented at departure. Over short distances, prolonged searches including some trial and error
may finally lead to success. Over long distances, in contrast, only a minority
(21%) returned to the loft, but these were birds that were exceptionally welloriented from the beginning. Apparently, they were individuals with a particularly pronounced homing drive as well as a substantial navigational competence.
To home over a long distance, a bird must have both these qualities: motivation to return to its home site and capability to find the way towards it. If a bird
does not home, however, it is not immediately clear which of these qualities is
poorly developed. Is the bird unable or unwilling to home? Recoveries of
non-homers give some hints but no final answer. Pigeons that were reported
from sites not more than 2030 km away from the release site appeared randomly distributed around that site. Most birds that had covered longer distances, in contrast, were clearly on the way towards home (Fig. 3.6A). Thus,
roughly a quarter of the (fairly young) first-flight pigeons of the Wrzburg loft
did not even show any tendency to fly home. Either they were not interested in
homing and therefore ended their flight early without orienting it homeward,
24
n Fig. 3.5. Further results extracted from the experiments shown in Fig. 3.4, supplemented by
four releases from only about 7.5 km. A Return rates (solid line with filled dots) and mean homeward components of vanishing bearings (dashed line with open dots) as correlated with the distance of displacement. B Homeward components of vanishing bearings in three classes of distance from home, separately shown for successful homers (solid columns) and birds that never
returned (cross-hatched columns). Numbers give numbers of pigeons. Horizontal lines indicate
the mean of all birds pooled. (Data in this form unpublished)
or they were unable to determine the direction towards home and stopped flying early, because they did not know where to fly. We shall see later that a low
motivation to home appears to be the generally more appropriate explanation,
because pigeons whose homing ability was experimentally obstructed, nevertheless flew long distances away from the release site, albeit in arbitrary directions (Sect. 7.1.1).
Not only were the successful returners over long distances fairly well oriented already at departure (Figs. 3.5B and 3.6Bd), but also those pigeons that
were found far away from the release site, most of them approaching home but
without reaching it (Fig. 3.6Bc). The degree of angular variance of initial bear-
25
n Fig. 3.6. Recoveries and initial orientation of first-flight pigeons released at 14 sites 120, 150 and
180 km from the home site near Wrzburg (for locations of release see Fig. 3.4). A Absolute angular
deviations of recovery sites from the homeward direction as a function of beeline distance flown
from release site. Dots along the three lines show running medians 99% confidence limits. B Vanishing bearings classified according to subsequent recovery distances or homing success, respectively, of individual pigeons (homeward direction upwards). Peripheral dots refer to individual
bearings, central vectors to means per release site; ellipses indicate 95+99% confidence range of the
resulting second-order mean. Symbols refer to direction of release site from home as indicated.
Homeward components below the diagrams result as first-order means from pooled bearings and
as second-order means from release-site vectors, respectively. Sample sizes are given inside the circles: number of bearings/vectors (= included sites). (Data in this form unpublished)
ings seems to reflect not only external and internal circumstances controlling
the reachable level of navigational performance but also the birds impetus to
fly towards home. Pigeons with an apparently weaker homing drive showed
poorer initial orientation (Fig. 3.6Ba,Bb).
In releases over shorter distances, not only homing success but also homing
speed can be considered. Most of the pigeons homing fast from a given site
were well homeward oriented already at departure, whereas among the slow
26
homers and the never returned pigeons many had started towards wrong directions. On the other hand, however, initial flights towards home do not guarantee fast homing or arrival at home at all. As in the long-distance releases
(Fig. 3.6), part of apparent homeward orientation appears to be based on true
navigational performance, but another part merely on chance. Both temporal
variations between days of release and variability among individual pigeons
released on the same day contribute to the nevertheless overall positive correlation between initial orientation and homing performance that can be observed
at the individual release sites (Kiepenheuer et al.1993; Wallraff and Kiepenheuer
1994). No clear correlations, however, have been found between mean initial
bearings at different sites, as shown in Fig. 3.4, and homing performance.
The findings reported in this section make it very likely that, in general, results obtained with first-flight pigeons do not reflect the level of navigational
performance that in principle can be achieved by birds without any spatial experience far outside their very restricted home range. As the individual quality
of the birds cannot be determined before release, there is always a notable proportion of little-motivated birds in a sample used for an experiment. These
birds produce a bottom sediment of largely randomized poor data, which
cannot be separated from good data. Thus, the noise found in a data set as a
whole not only indicates poor navigational performance. Samples of experienced pigeons no longer contain such non-motivated failure birds which did
not return after the first displacements, but their unavoidable familiarity with
a larger area raises other difficulties (Sect. 3.2).
Our often limited knowledge of the pigeons motivation to home is not only a
problem in evaluations and interpretations of homing data obtained with normal untreated birds, but even more so in investigations using birds that are experimentally manipulated in a particular way. If their performances are reduced, it is necessary to ask whether the experimental interference reduced
their ability or their willingness to home. This problem will arise below in various contexts (e.g. Sects. 6.3.1, 7.17.3, 7.8.2).
3.1.3
Limited Significance of Exercise Flights at Home
Homing pigeons are usually living in great numbers (up to several hundreds)
in a loft in which they find shelter for the night, breeding niches, food, water,
etc., but which they can leave to fly around and perch outside on a roof or elsewhere. Spontaneously, and preferably in flocks, they fly in circles and loops in
the close vicinity of the loft, sometimes achieving considerable height above
ground. The spatial range of these spontaneous flights is not exactly known
[but should now be ascertainable by GPS tracking (Sect. 2.3; see Biro et al.
2002)]. According to my non-systematic observations at a number of different
lofts, the radius covered is usually within a range of 12 km (flocks rarely disappear from sight), but I cannot exclude that at some lofts or on some occasions greater distances (up to 5 km or so?) may be reached.
27
Clearly, these exercises habituate the young pigeons to skilful flying, which is
a precondition to home over long distances. In addition, flying around gives
them an opportunity to becoming familiar with an aerial view of their home
site and its closer vicinity. Theoretically, scanning a somewhat extended area
around home might also be necessary to learn spatial configurations (e.g. gradients) of environmental signals that are required to develop a navigational
map. Thus, testing of whether free flights at home are a precondition for longdistance navigation was one of the early experiments on pigeon homing (Kramer and von Saint Paul 1954).
Later on, many more experiments were conducted using pigeons that lived,
from fledging time onwards, in an aviary made of wire-mesh which allowed
straight flights over only 816 m (e.g. Kramer 1957, 1959a,b; Wallraff 1966a,
1970a,1979).Such aviary pigeons got their very first opportunity to fly in the free
air space when they were tossed into the air, at an age of 46 months, at a release
site far distant from home. It is not surprising that many of these unpractised fli-
n Fig. 3.7. Vanishing bearings (cir-
cular diagrams) and recoveries of

aviary pigeons displaced over 147
193 km from their home site at Wilhelmshaven (Wi) and near Giessen
(Gi), respectively, to a release site in
between, near Osnabrck (Os), and to
two opposing sites near Leck (Le) and
Stuttgart (St). Open symbols refer to
birds from Wilhelmshaven, filled
symbols and dashed lines to birds
from Giessen. (Modified from Wallraff 1970a)
28
n Fig. 3.8. Performances of first-flight pigeons that had been allowed to fly ad libitum around
their loft in Osnabrck (left) and others that had been confined all the time in an aviary in the
same garden (right). From top to bottom: vanishing bearings at the four release sites (distance
from home 8794 km); the same bearings pooled with respect to home and north (n=133 and
117); recoveries. Return rates are given as percentages of birds released. Inside circles is indicated
direction and length of mean vectors. (Modified from Wallraff 1970a)
3.2 Experience Gained by Homing Flights
29
ers landed soon afterwards nearby or disappeared low-flying behind trees or

hills or otherwise in the vicinity. Nevertheless, some pigeons continued their
flight sufficiently high to provide useful vanishing bearings. Also, many of the
initially landed birds were afterwards reported from far-distant recovery sites.
Most of these sites were closer to home than the release site (Fig. 3.7).
Compared with free-fliers released at the same site and mostly on the same
day, aviary pigeons were not worse in initial homeward orientation (Fig. 3.8).
Dramatic differences, however, were found in the return rates. Almost half of
the free-fliers returned, but only 1 out of 198 aviary pigeons. This failure was
apparently not due to lack of ability to navigate over long distances. Recoveries
of the non-homers show a more clear-cut approach to the home area in the aviary pigeons (Fig. 3.8). This outcome suggests that the latter birds were well able
to accomplish the true navigational task, but that their lack of experience of an
aerial view of the area prevented them from detecting the loft and the aviary itself between many similar looking houses and gardens which these birds had
never seen before. Nine aviary pigeons, but only one free-flier, were reported
from sites within the home town less than 4 km away from the loft. On the assumption that most of the free-fliers that reached the familiar town area were
able to localize the loft itself and hence were not reported from elsewhere, it is
understandable that the recoveries of the free-fliers are less closely clustered
around home. It is apparent that exercise flights at home are needed only to establish an enlarged target area around the loft by memorizing local landmarks,
but not to develop a navigation system enabling goal-oriented flights to that
target area from far-distant unfamiliar sites.
Even the deviations from a symmetrical pattern are similar in the two categories of pigeons. Initial bearings show some preference for west-northwesterly directions and the recoveries are accumulated in an area southwest of
home (Fig. 3.8). In other experiments, a more clearly expressed PCD has been
found also in aviary birds (Wallraff 1978a, 1979). Thus, the basic features of
long-distance navigation do not depend on free-flight conditions at the home
site. We shall see later, however, that aviary pigeons are able to orient homewards only on the condition that they are exposed in their cage to natural
winds (Sect. 7.5.1).
3.2
Experience Gained by Homing Flights
First-flight pigeons released somewhere abroad for the first time could have
developed their navigational abilities only at the home site itself and, if they
had not been confined in an aviary, in its close vicinity. In releases following
their very first homing flight, the birds potentially may, and actually do, profit
from experiences they have made within more extended areas. It is, therefore, a
serious difficulty in pigeon homing research that earlier experiments conducted with a given pigeon influence the results achieved with this bird in later
experiments. Moreover, as many pigeons do not return from the first release(s)
30
(Sect. 3.1.2), individuals with poor homing qualities may be sorted out, so that
the quality of the remaining sample is modified also by means of selection. The
consequences of experience and selection need to be known and considered in
order to design experiments properly and to draw appropriate conclusions
from them. A third parameter, age, usually proceeds in parallel, but its possible
role independently of the others cannot, with the commonly obtained data, be
isolated (Sect. 3.1).
3.2.1
Number and Range of Previous Homing Flights
Pigeons considerably experienced by many previous homing flights show,on average, better homeward-oriented initial bearings than first-flight birds, even if
they had not yet been released at the current test site. Owing to the great temporal variability (Sect. 3.3.2), comparisons are most reliable in experiments in
which both kinds of pigeons are released semi-simultaneously in alternating
succession (Fig. 3.9). Even then, differences between individual releases are very
variable, sometimes dramatic, sometimes non-existent, and sometimes the sign
is reversed. [Even if arranged optimally symmetrically, previous flights cannot
be spatio-temporally completely symmetric and may sometimes modify the
current bearings (Sect. 3.2.2).]
While initial homeward orientation is generally improved by experience, the
expression of a PCD (Sect. 3.1.1) is generally weakened (Fig. 3.9, B vs. D). It is
unknown whether the directional shift by about 35, as shown in Fig. 3.9, expresses a meaningful systematic change.
In these 44 simultaneous experiments with both inexperienced and experienced pigeons, the mean homeward component as well as the return rate were
higher in the veterans over the whole range of distances used (Fig. 3.10). While
there was no clear correlation with distance in initial orientation, homing success decreased drastically with distance in the first-flight birds (see also
Fig. 3.5), but only slightly in the veterans. Also, the difference between the two
groups in homing success was more consistent, which was better in the veterans in almost all experiments, whereas the initial bearings were better homeward oriented in only 70% of the releases.
Over shorter distances, also an increase of homing speed with an increasing
number of preceding flights has been observed. The results shown in Fig. 3.11
are representative in principle, but the improvement curve in detail depends
on pigeon strain and loft site (cf. Schmidt-Koenig 1963b). Compared with
other series of releases at distances of 2030 km around other lofts in Germany, the number of non-returners from the very first flight is remarkably low.
In other series it was in the range of 3050%, while in Italy it was found similarly low (e.g. Fo et al. 1984; Kiepenheuer et al. 1993; Wallraff 1994a).
The dashed curve and the bar graphs in Fig. 3.11 illustrate that losses of poor
individuals contributed little to the general improvement as compared with the
increasing homing speeds of the remaining birds. Pigeons that failed to return
31
n Fig. 3.9. Summaries of vanishing bearings obtained in 44 releases with two pigeon groups at
seven symmetric quartets of sites at distances of 30180 km from the home loft near Wrzburg.
A, B Inexperienced (first-flight) pigeons; C, D experienced pigeons after at least 12 previous
homing flights over varying distances from largely symmetrical directions. Most birds had never
been released before at the particular site used here or in its closer vicinity. Arrows give mean
vectors per release (radius = length 1). Second-order vectors from these 44 vectors (= centres of
95+99% confidence ellipses) as in Fig. 2.4 top part, second-order vectors from 44 first-order directions (irrespective of length) as in Fig. 2.4 bottom part, and first-order mean vectors from the
pooled bearings shown by peripheral dots are given numerically (direction and length; * P<0.05,
** P<0.01, *** P<0.001). Homeward components in 44 releases different between A and C with
P<0.001 (Wilcoxon matched pairs test). (Data from my files, in this form as a whole unpublished)
32
n Fig. 3.10. Mean homeward component of vanishing bearings (A) and return rate (B) as a function of distance in experienced pigeons (hatched columns) and inexperienced pigeons (black columns). The same material as included in Fig. 3.9. Numbers between the diagrams refer to number
of releases/number of (centrally symmetrical) sites
from later releases did not home significantly slower than others in earlier
flights and their initial homeward orientation was not, or only a little, poorer already in their first release (Wallraff 1959a, 1994a). Nevertheless, pigeons that fail
to return from their very first release appear to include constitutional losers
whose motivation to orient and fly homeward is weakly developed (Sect. 3.1.2).
Over long distances, such pigeons are sorted out, whereas over short distances
some of them may eventually reach the loft. Their success may then stimulate
their motivation to home from later releases.
It is hardly possible to decide to what degree improved initial orientation as
well as homing performance in experienced pigeons (Figs. 3.93.11) are due
(1) to improved knowledge of navigational signals in a more extended area, (2)
to improved practice in evaluating such signals, (3) to enhanced homing motivation owing to already experienced homing success, and/or, last but not least,
(4) to selective loss of inherently incapable or little-motivated individuals that
did not return from the first releases.
If not only short distances were included (as in Fig. 3.11) but also displacements over more than 100 km, only a small fraction of 1020% of the originally
available pigeons remained after some 10 releases (according to my experiences in Germany; see also Sonnberg and Schmidt-Koenig 1970). These remaining veterans were then very reliable in homing from almost everywhere
within a radius of at least 200 km. Their initial orientation, however, never
reached high accuracy (see Fig. 3.9C).
33
n Fig. 3.11. Homing performance (in units of classes as given in the insert) as a function of in-
creasing homing experience. Thin line with black dots starts with steps of exact number of
pre-releases and then summarizes broader intervals. Heavy line shows means three-fold mean
error of experience classes as indicated by Roman numerals; dashed line shows corresponding
means with non-returners omitted. Insert gives percentages of birds per performance class for
experience levels IV. Data from 18 release sites: 8% from five sites about 11 km distant, 70%
from eight sites at ca. 22 km, and 22% from five sites at ca. 44 km from the home loft in Wilhelmshaven, Germany. (Wallraff 1959a)
3.2.2
Directions of Previous Homing Flights
Spatio-temporally, experience gained during homing flights always includes
an element of asymmetry. Even within a training pattern using symmetrically
arranged release sites, an individual bird goes through a spatio-temporally unbalanced sequence. When asymmetrical experience is most obvious, as for instance in a pigeons second flight, its influence on initial orientation can also be
obvious. Figure 3.12 shows a very clear-cut example. While first-flight pigeons
departed in their loft-specific PCD, the mean bearings of simultaneously released second-flight birds agree almost exactly with the directions they had
flown in their preceding first homing trip.
Effects are not always so dramatic. Quantitatively, deflections on the bearings
in the second release caused by the direction flown previously vary between
100% and nil (Fig. 3.13). In most cases, they are somewhere in between, thus
more or less modifying the direction induced by the current local circumstances
34
n Fig. 3.12. Vanishing bearings of first-flight pigeons (open symbols) and second-flight pigeons
(filled symbols) at the same release site 161 km NNE of the home loft in Wilhelmshaven. Filled peripheral arrows and associated distances refer to the previous first flight of the respective birds.
Equally shaped symbols in different diagrams indicate simultaneous release on the same day.
(Wallraff 1967)
n Fig. 3.13. Ten second-flight releases were conducted, in which one group of pigeons had made
its first flight towards a compass direction approximately 90 left from the actual homeward direction (filled symbols) and another group towards approximately 90 right from actual home (open
symbols). Mean directions of the groups are ordered according to decreasing angular difference.
(Data from Wallraff 1974b)
35
alone. If a sample of birds as a whole is well oriented towards home, a strong influence of the previous release is less likely to occur than if current homeward
orientation is weak. This correlation may be considered trivial because the birds
cannot be well-oriented towards two diverging directions. Yet it raises the question for the causal relationship. Do the pigeons appear to be poorly homeward
oriented because they simply prefer to reproduce their previous homing direction, or do they decide on this formerly correct direction because they have difficulties in determining the currently correct direction towards home? In the first
case, variations between experiments would indicate varying states of motivation; in the second case, they would suggest varying environmental conditions
enabling homeward navigation. It is not immediately plausible why this particular kind of motivation should be variable, whereas varying availability of navigational signals is very likely (Sects. 7.3.2 and 7.6).
The assumption that the latter causal connection primarily controls the
amounts of directional after-effects is corroborated by results obtained at a site
at which usually no homeward orientation was visible at all. Roughly following
their loft-specific PCD (Sect. 3.1.1), by far most of the pigeons regularly departed there in directions leading away from home (Wallraff 1959a,b). So did a
group of pigeons in their first release (Fig. 3.14A). The mean bearing obtained
in their fifth release, after three releases from elsewhere, was quite similar
(Fig. 3.14B). However, in the meantime, the group had been divided into two
subsamples partly homing from different sites. Although the overall pattern of
bearings remained similar to the first release, the birds were differently distributed within this pattern according to their previous homing directions. In
other experiments using third-flight birds whose first and second flights had
been from opposite directions, significant deviations in favour of the compass
direction of the second flight were observed (Wallraff and Sinsch, unpubl.
n Fig. 3.14. First release (A) and fifth release (B) of the same pigeons at a site 23 km east of the
loft in Wilhelmshaven. The second release was from 22 km NW (group N, filled symbols) or SW
(group S, open symbols), respectively, the third release of all birds from 44 km W, and the
fourth release from about 10 km N (N, filled peripheral arrow) or S (S; upper arrow). In A, the
birds are distinguished according to their subsequent different experience; actually, they were all
completely inexperienced. (Wallraff 1974b)
36
data). Even if the previous flights were spatially symmetric, they cannot be spatio-temporally symmetric as well.
A casual observation by Wiltschko and Wiltschko (1987, Fig. 7; see also
Wiltschko 1991, Fig. 3) suggests the assumption that even directional self-training may sometimes occur. Part of a group of hitherto not yet displaced young pigeons made an unusually long spontaneous flight, remaining out of sight of the
loft area for 75 min, while another part remained at the loft. Released on the following day at 65 km distance, the two subsamples deviated similarly from home,
but in different directions, their mean bearings differing by 90. It is unknown,
however, whether this was actually an effect of directional self-training or of different map learning (as the authors assume) or of something else.Although such
long excursions away from the loft are certainly exceptions rather than the rule,
the case demonstrates that even samples of first-flight pigeons, whose spontaneous flights are not under permanent control, may be heterogeneously composed
according to possible effects of previous self-made homing experience.It is clear,
however, that loft-specific PCDs do not generally result from self-training by
asymmetric exercise flights around the loft, as they develop also in aviary pigeons (Fig. 3.8; Wallraff 1970a, 1978a, 1979, 2001).
In experiments using veteran pigeons that had been released many (e.g.
some 2040) times before, an immediate after-effect of the very last preceding
homing direction cannot usually be separated, provided that the pattern of experience was spatio-temporally largely symmetric (Schmidt-Koenig 1976).
However, with asymmetric training patterns comparing birds experienced in
homing from sites distributed within opposed semicircles, pronounced directional differences were found at the edge sites common to both semicircles and
also at more distant sites outside of the training range (Graue 1965).
Such effects of directional training, even if not intended by the experimenter,
need to be considered as possible sources of directional preferences which interfere with, and modify, the outputs of the navigational process referring to
the spatial relationship between current release site and home site (Sect. 3.7.1).
In a single release they cannot usually be recognized, because the participating
birds are usually not differently trained and because the amount of a possible
training effect cannot be predicted (cf. Fig. 3.13). On the whole, and particularly if the sequence of preceding releases is not roughly symmetrically balanced and/or the current conditions for homeward navigation are poor, an effect of directional training is one of the factors that potentially contribute to
the great variability of initial-orientation patterns.
3.2.3
Familiarity with the Release Site or Area
Within sequences of releases at sites in varying directions, an effect of general
homing experience (Sect. 3.2.1) is much more obvious than an effect of having
previously been released at the same site, albeit a slight improvement of homing
performance by local experience has been observed (Wallraff 1959a). Release-
37
n Fig. 3.15. Vanishing bearings in a sequence of six releases of the same pigeons at always the
same site 40 km from their home near Cambridge, England. Filled symbols Sun visible; open symbols overcast. Numbers give the resulting homeward components. (Data from Matthews 1963a)
site biases may decrease a little in a second or third release at the same site, but
mostly they remain typical for the site (e.g. Keeton 1973) or follow a shift based
on general rather than local experience (Wallraff 1959a). Some alteration observed in a second release at a given site need not always be due to previous
experience of location-specific signals. If only one of the two groups had been
released in the example of Fig. 3.14B, one might have concluded that local experience has reduced the release-site bias, although the deviation from the direction flown earlier was obviously due to directional training from other sites.
Improvement of initial homeward orientation is usually observed in successions of releases from always the same site (Matthews 1963a; Kowalski and
Wiltschko 1987). Considering some of the aspects discussed below (Sect. 5.3,
Chap. 8), the example in Fig. 3.15 suggests that the first release reflects action
of navigational map plus sun compass; in the second flight the sun compass
could not operate and local features were not yet sufficiently familiar; thereafter the pigeons became increasingly familiar with the landscape so that, finally,
in the sixth release, they could perfectly pilot home without needing the sun. In
such cases, when the sun is visible, the experimenter cannot distinguish between effects of familiarity with local (e.g. visual) signals, effects of directional
training and a possible combination of both (realizing that it is again where it
was last time, the bird repeats its last homing direction). Improvement may
mostly occur in the first repetition (Kowalski and Wiltschko 1987; Grter and
Wiltschko 1990), but in a sequence of 65 releases it required more than 30 repetitions until an original deviation from home of about 90 was reduced to a finally remaining mean deviation of 11 (Kowalski and Wiltschko 1987). A subsequent clock-shift (Sect. 5.1) revealed that, with such intense training, the
birds simply repeated the entrained compass course (Fller et al. 1983; see the
Appendix referring to Sect. 8.1.4).
Although general improvement of initial orientation due to limited local or
regional experience (and apart from immediate repetitions of analogous
flights) is not obvious in releases using non-manipulated pigeons, we should
not conclude that the birds are unable to recognize and evaluate environmental signals indicative of a given site or area with which they had previously
been familiarized. By means of specifically designed experiments it is possible
to show that pigeons do pay attention to such signals and can use them for
home-finding (Sects. 8.1.1 and 8.1.4).
38
3.3
Temporal Variability
Pigeon homing varies depending not only on the site of release (Sect. 3.1) and
on the individual birds previous homing experience (Sect. 3.2), but also on
time. The directions chosen at departure vary from day to day, to a different
amount at different sites (see Fig. 3.3), and also homing performance varies
considerably. Figure 3.16 illustrates that the greatest differences in homing
speed and return rate have been observed among experiments conducted in
summer and winter. Therefore, seasonal fluctuations shall first be inspected.
3.3.1
Annual Periodicity
Figure 3.16 refers to experiments in which the other variables, release site and
experience of pigeons, have been held constant. To cover the whole annual cycle with more data, it was necessary to compensate for these other variations
by some standardization procedures (Wallraff 1959b). Then it turned out that
homing performance exhibits a clear annual periodicity with a maximum in
late summer and a minimum in winter, which is closely correlated with ambient temperature (Fig. 3.17). At least two causes appear possible: some seasonal
influences may act either (1) on the pigeons physiological/motivational state
or (2) on the environmental conditions that are necessary for home-finding.
Several expectations linked with the first possibility have been tested: (1a)
The birds motivation to home might fluctuate with its hormonal state and
breeding cycle. The fact that the maxima of the two cycles are greatly out of
phase (Fig. 3.17B: April versus August) argues against this possibility. Further,
considerably increased breeding activity in winter, induced by an artificially
lengthened photoperiod in the loft, did not induce improved homing perfor-
n Fig. 3.16. Homing performance in 12 analogous experiments with first-flight pigeons released
at a site 22 km northwest of the home loft in Wilhelmshaven (days means homed after the day of
release; never means never returned). (Modified from Wallraff 1959b)
3.3 Temporal Variability
39
n Fig. 3.17. Homing performance of differently experienced pigeons from four fairly symmetrical sites at 22 km distance from home in Wilhelmshaven as a function of season. A Means per release in relative units, symbols referring to the four sites. B Monthly means resulting from A are
shown, the corresponding air temperature during the days of release (scale on the right, C) and
number of clutches in the loft as a percentage of total number per year (means of 4 years, ranging
from a minimum of 2.2% in January to a maximum of 16.3% in April). Dashed horizontal bar indicates the period during which performances were correlated with short-term changes of temperature. (Modified from Wallraff 1959b, 1960)
mance (Wallraff 1959b). (1b) Responses of the organism might be directly influenced by the ambient temperature. If so, interrelations would have to be
rather complex. Correlations between short-term deviations from the seasonal
mean temperature and homing performance were found from September to
March, but not during the other months (dashed bar in Fig. 3.17B). Thus, the
birds would have to be sensitive to short-term fluctuations at a fairly high temperature level in autumn and insensitive at a lower level in spring. Keeping pigeons for 14 h before release at a temperature between 20 and 30 C had no
effect on initial orientation and homing performance (Wallraff 1960).
40
n Fig. 3.18. Annual cycles of homeward component of vanishing bearings (A) and homing performance (B) in experiments over 1525 km at four loft sites, two in northern Germany
(Wilhelmshaven and Gttingen), one in southern Germany (near Munich) and one in Italy (near
Pisa). Monthly means (smoothed over 3 months), deviations from annual mean in percent of this
mean. Arrow-labeled intervals indicate periods during which all German values are below or
above the annual mean, respectively. Wilhelmshaven data are the same as in Fig. 3.17, the others
are from inexperienced birds (Munich and Pisa: same experiments as in Fig. 3.2). (Data from
Wallraff 1959b; Gronau and Schmidt-Koenig 1970; Fo et al. 1984)
Owing to these findings, it seems more likely that the correlation between
homing and temperature reflects an effect of temperature on decisive environmental conditions, e.g. availability or usability of navigational signals, rather
than a direct effect on the animals. Later on, when we deal with the involved
sources of information, this assumption will gain plausibility (Sect. 7.4.2).
3.3 Temporal Variability
41
Corresponding annual cycles in homing performance were also found at two

other loft sites in Germany (Fig. 3.18B). In Italy, some annual rhythmicity was
only weakly expressed; its phase was shifted 2 months forward against a simultaneous parallel series of experiments conducted in southern Germany (thin
curve in Fig. 3.18B).
Less coherent are the data that have been reported on initial orientation
(Fig. 3.18A). In the series of experiments around Wilhelmshaven, on which
Fig. 3.17 is based,no annual cycle could be detected at all.Very weak rhythmicity,
parallel to homing performance, was indicated in southern Germany and Italy.
The only pronounced annual cycle of initial homeward orientation, also in synchrony with homing performance, was found by Gronau and Schmidt-Koenig
(1970) at a release site 15 km north of home in Gttingen.It is not quite clear why
at the other German loft sites annual periodicity was only weak or lacking. Possibly the initial bearings were too much determined by directional preferences
that are independent of seasonally variable environmental cues indicating the
current position (Sect. 3.7.1).
3.3.2
Aperiodic Fluctuations
Other temporal fluctuations observed in pigeon homing did not involve periodicity. For instance, I did not find any consistent diurnal rhythm in extended
release experiments covering several hours (unpubl. data; see also Fig. 3.19
n Fig. 3.19. Two examples of intradiurnal fluctuations in initial orientation. Vanishing bearings
are summarized in circular diagrams and shown in their temporal sequence, 0 representing the
overall mean direction (arrow in the circle), positive values clockwise deviations from it. Grouping of consecutive bearings is highly significant (P<0.0001). Sixth and seventh release of the same
group of pigeons which had identical experience from four preceding homing flights. The home
loft was in southern Bavaria. (Wallraff 1971)
42
and Keeton 1974a). Aperiodic fluctuations, however, have been found in the
course of hours, from day to day and among different years.
Day-to-day fluctuations of initial orientation as well as homing performance
are very common. Examples are distributed over the whole literature on pigeon
homing (see, e.g., intraseasonal examples in Figs. 3.3, 3.16 and 3.17A). Differences among different days are often fairly small and indistinguishable from
noise typical for the data, but significant differences and clear exceptions to the
rule observed at a given site do also occur. As will be seen below, these fluctuations do not always, not only and perhaps not predominantly indicate varying
levels of homeward orientation, but appear to reflect also concomitant circumstances not directly involved in the home-finding process (Sect. 3.7.1). This
seems to hold true also for long-term differences in homing behaviour as observed among different years (e.g.Wallraff 1959b,1986; Kiepenheuer et al.1993).
Significant fluctuations of departure bearings beyond random noise have
even been observed at a time scale of several minutes (Fig. 3.19). The frequency range covered about 16 periods/h (Wallraff 1971). Such short-term
variations can, however, only be detected within long-lasting releases embracing a large homogeneous group of experienced pigeons sharing the same sequence of previous homing flights. In birds with no or little homing experience, whose bearings reflect less clearly pure homeward orientation, potential
small-scale, short-term fluctuations are probably masked by noise and homeindependent directional tendencies such as a PCD (Sect. 3.1.1).
3.4
Accuracy of Homeward Orientation
We have learned from the above graphs showing initial bearings, distributions
of recovery sites or homing times and return rates that pigeons, in particular
inexperienced pigeons, usually do not fly on a straight direct course home.
Mostly we observe a considerable angular dispersion or variability in homing
performance, respectively, among individuals. Even the mean course of a sample of birds deviates, in most cases, more or less from the beeline direction towards home. Some aspects of these inaccuracies (release-site biases, PCD, temporal variability) have already been described and discussed. A few more
aspects will be added in this section.
3.4.1
Distance of Displacement
Schmidt-Koenig (1966,1970) found,in very experienced pigeons,a clear correlation of the initial homeward component with distance from home around two
lofts, one in Durham, North Carolina (USA), and one in Frankfurt am Main
(Germany). Birds were well homeward-oriented at very short distances of less
than 20 km, poorly oriented in a range of about 2060 km, and reached again
fairly high levels in a range of 100200 km. Results of subsequent series of exper-
3.4 Accuracy of Homeward Orientation
43
iments around other lofts (and also around Frankfurt: Wiltschko 1992) were not
as consistent (see Fig. 30 in Wallraff 1974a and references therein), but some increase between about 50 and 200 km may prevail, possibly more in naive rather
than in experienced birds (see Figs. 3.5 and 3.10). At least within a radius of
about 25200 km and in fairly flat regions, there is no general decrease in homeward directedness with increasing distance from home (see also Sects. 3.5 and
7.4). It is impossible, however, to deduce a rule on the role of distance that is valid
for all geographic regions.
3.4.2
Geographical Variability
Pigeon homing functions at different levels of performance in different regions of the world. Figure 3.20A compares initial homeward components (and
lengths of PCD vectors) of first-flight pigeons released at four sites symmetri-
n Fig. 3.20. Initial orientation of first-flight pigeons at different latitudes. Black bars give second-order mean homeward components, white bars lengths of second-order mean compass vectors (directions as indicated). A Vanishing bearings at four symmetrical release sites 2031 km
distant around each of six home lofts. Small triangles Long-term annual mean of air temperature
in the respective region (scale above). B Corresponding data from two lofts showing means not
only of vanishing bearings (Van) but also of bearings observed 20 and 40 s after release (scale as
in A). (Fo et al. 1982, 1984)
44
cally distributed in 2031 km around six home sites at different latitudes

roughly along the same meridian. At the two southern lofts in Italy the birds
were clearly better homeward-oriented than at the four northern lofts in Germany. In an extra comparative series of experiments, pigeons from the same
German stock were held under similar conditions in Germany and Italy. In
each country, 12 simultaneous releases, evenly distributed over an annual cycle, were conducted at two opposite sites 2225 km distant from home. Not
only was initial homeward orientation significantly better in the southern area
(Fig. 3.20B; see also Fig. 3.3), but also so was homing performance.
It is readily possible to interpret these findings as a spatial analogue to the
temporal variations in the course of a year (Sect. 3.3.1). In either case, the level
of performance is correlated with ambient temperature. It should also be mentioned, however, that over longer distances Germany appears to be more advantageous for homing than Italy, probably due to other reasons (Sect. 7.4).
Regional differences among various countries and continents have also been
found in homing of more or less experienced pigeons (Fig. 7.4; Wiltschko et al.
1987d). As kind and degree of experience, keeping conditions, landscape characters and pigeon strains were also variable, however, the causes linked with
geography cannot always be clearly separated from possible other causes.
3.4.3
Distraction by Landscape Configurations
Pigeon homing experiments are usually conducted in structured landscapes
containing mountains, hills, valleys, lakes, forests, villages, towns etc. Do such
features influence the flight paths, i.e. do they deflect the birds from routes they
would have flown over a neutral blank surface?
Prominent topographical features such as high mountains, coastlines or large
lakes clearly act as barriers which the pigeons hesitate to overfly.Detours around
a mountain,along a valley or along a shore are obvious (Wagner 1968,1970,1972,
1974; Ioal et al. 1994; Bonadonna et al. 1997). Consequently, most of the release
sites used to investigate pigeon homing are located in non-spectacular surroundings which do not directly enforce a deflection. However, also these surroundings are usually not homogeneous in all directions. The birds might prefer
flying towards or away from one or another topographical constituent of the terrain into which they are released. In fact, it is sometimes intuitively apparent for
the observer that the released pigeons are attracted by a nearby village or even
by a single farmhouse or, respectively, by local pigeons sitting there or flying
around. Even if the released bird finally flies away, its vanishing bearing may be
influenced by the initial detour it has made.
Such observations and impressions cannot be evaluated as quantitative data.
At some sites, however, outstanding modes of bearing distributions in the directions of neighbouring villages strongly suggest attraction of these villages
(Sect. 8.2; Kiepenheuer 1993). Moreover, evidence for preferred flying towards
human settlements and some avoidance of flying over wooded areas resulted
3.4 Accuracy of Homeward Orientation
45
n Fig. 3.21. Influence of human settlements (filled symbols) and wooded areas (open symbols)
on vanishing bearings of pigeons released at 28 sites around their loft near Wrzburg. Ordinate
indicates the mean proportional difference between frequency of respective landmarks at a given
distance and frequency of pigeons vanishing in the direction of such landmarks. Plus means
more and minus fewer birds than expected in flights not influenced by topographical features.
Dotted area marks the approximate distance range within which the pigeons vanished from sight
after having overflown the shorter distances below that range. Symbols indicate level of significance for difference from zero: diamonds P<0.01; squares P<0.05; circles P>0.05. (Modified from
Wallraff 1994b)
from a statistical analysis of initial bearings at 28 sites (Fig. 3.21). These global
results are statistically highly significant (Wallraff 1994b), but do not allow a
reliable interpretation of the birds behaviour at each individual release site. A
more or less suggestive influence of topographical features on the initial bearings observed at a given site may still be a result of chance. However, the global
statistical results indicate that, in general, topographical features could have
influenced the initial bearings observed at a given site in such a way that they
do not directly reflect the directional output of the pigeons navigation system.
Responses to landscape features appear to depend on the surroundings to
which the pigeons have been habituated at their home site. Results as shown in
Fig. 3.21 have been gained with pigeons from a loft within an isolated small assembly of buildings in an open rural countryside, but not with those from another loft lying at some distance from human settlements and closer to a forest
(Wallraff 1994b). Similar differences according to loft sites have been found by
Hitchcock (1952) who observed in pigeons from one particular loft, a decided
preference ... for wooded rather than open country, i.e. just the reverse of the
preference shown in Fig. 3.21.
46
3.4.4
Stochastic Noise
Given the great angular dispersion of initial bearings and great variability of
homing speeds, as usually observed within one experiment conducted with a
sample of pigeons, we cannot expect that the homing paths of individual birds
are strongly determined. Circles and loops often flown within the first minutes
after release (see insert in Fig. 2.3) certainly contain random elements which
also affect the direction in which a pigeon eventually vanishes from sight. Inaccuracies of initial bearings, long homing times and, sometimes, fairly low return rates make it likely that a considerable amount of stochastic noise is included in all the collected homing data. These data do not immediately reveal,
however, to what degree the source of noise is inherent in the animals or in the
environment. Do the animals not do it better, although they were able to do it
better? Are the animals unable to evaluate the navigational signals provided by
the environment in an optimal way? Or are these signals themselves so noisy
that even an optimal mechanism cannot gain better performances?
It is clear that, at least in inexperienced pigeons, some source of noise is inherent in the animals. There are good birds and poor birds, and a noticeable deal
of the differences among individuals apparently concerns the motivation to
home rather than the navigational ability (Sect. 3.1.2). Comparisons among individuals of a homogeneous sample, in longer sequences of experiments, with
respect to their relative position within the group led to the following results
(Wallraff 1994a): Differences among habitual champions and failures were most
obvious in homing speeds after successive releases at the same site, i.e. in a kind
of performance that was determined by the motivational and physical state
rather than by the birds navigational capacity. Some differences were also found
in initial orientation at very familiar sites, suggesting that each bird tended to
follow its own traditional route. No consistent ranking of individual birds was
possible, however, in initial homeward orientation at varying unfamiliar sites.
This outcome suggests that it was primarily a matter of chance which bird vanished nearest to the homeward direction, or deviated most from it, in which release. The angular dispersion of bearings appeared to reflect stochastic noise
and is most reasonably compatible with the hypothesis that no one pigeon was
able to gain clear-cut information on the direction of home, because noise was
inherent already in the environmental signals providing positional information.
3.5
Spatial Range of Pigeon Homing
In sport races, pigeons return to their loft over distances of hundreds and
sometimes more than a thousand kilometres. Yet these races do not imply evidence that pigeons are able to determine their position relative to home if they
are so far away from it. As mentioned above (Sect. 1.2), the birds are usually
transported toward the same direction with distances increasing in the course
3.6 Multiple and Mobile Home Sites
47
of a season. Thus, they are directionally trained, fly in large flocks, and hence it
is not necessary for each bird to determine its position. In experimental releases using singly flying pigeons in Germany, initial homeward orientation,
homeward-oriented recoveries and actual homing have been shown over distances of some 300 km (e.g. Figs. 2.2, 2.7 and 3.4), and even 500 and 700 km appear to exceed the navigational range not everywhere (Wallraff 1981a, 1993).
On the other hand, without the opportunity to gain navigational information
during the outward journey, pigeons in Italy failed to orient homeward at distances between 100 and 150 km (Benvenuti et al. 1994). We shall deal with the
range problem more thoroughly when focusing on navigational cues used for
home-finding (Sect. 7.4.1).
Apparently, no generally valid upper distance limit for home-oriented navigation can be fixed. The range seems to be variable depending on geographical
conditions. Empirical data obtained in various regions are scarce. Using single
or only few release sites very far away from home, it is difficult to ascertain
whether directional preferences compatible with homeward orientation do actually indicate correct determination of the current position. I assume that the
range within which pigeon navigation continuously operates covers several
hundred kilometres in many regions, in some regions perhaps more than a
thousand kilometres. According to present knowledge (Sect. 7.4.1) it seems very
unlikely, however, that pigeons use a global grid of coordinates, as was considered a possibility in former years (e.g.Wallraff and Graue 1973; Wallraff 1974a).
3.6
Multiple and Mobile Home Sites
It is evident that various migratory birds have two home sites, a breeding site
and also a well-defined wintering site or area (e.g. Berthold 1996). Other birds
commute between their nesting site and a far-distant foraging area (e.g.
Fig. 10.6C; Mouritsen et al. 2003). Thus, a bird must be able to collect and memorize relevant data characterizing two or more geographic positions and to
choose one of these positions as a current target. Homing pigeons are rarely
confronted with this problem. It is difficult but possible to accustom them to a
second home site after they have been living over several months or years in a
given loft. If a pigeon fancier buys an adult pigeon from another fancier, he or
she has to confine the bird in an aviary over a prolonged period of time and,
nevertheless, even after years runs the risk that the bird returns to its old loft.
In principle, pigeons can also memorize two home sites which may be hundreds of kilometres away from each other. If released in between, they may return to one or the other loft (Wallraff 1974a; Baldaccini et al. 1976). However,
the system seems primarily adapted to learning a home site during an early
sensitive period (Sect. 7.5.1).
Lipp (1996) describes experiments in which pigeons were trained to be fed in
a feeding loft some 2530 km away from their home loft. When released at a
third site, aside from the line connecting the lofts and about 1520 km distant
48
from either one, it depended on their state of hunger whether they flew immediately to the one or to the other loft. As distances were fairly short and the pigeons familiarity with the area not quite clear, it is unknown whether the birds
found their way (primarily) by visual piloting or by true navigation (Sects. 1.3
and 12.2).
Such questions are also unsettled in the case of mobile lofts which in former
years were often used for military purposes. Pigeons were trained to varying positions of their loft and within 23 weeks (Lipp 1996) they learned to home to a
new position. Systematic experiments with mobile lofts under a scientific point
of view have rarely been made (for unsuccessful attempts to show a position-independent attraction of the loft itself and/or its inhabitants see Sheldrake 1994).
3.7
Conclusions and Perspectives
Release a pigeon somewhere at some distance and it flies straight home: no, pigeons are not as perfect in homing. Without previous training from the particular site of release, it is unlikely that the pigeon flies straight homewards and it
is uncertain whether it will return at all. Homing of pigeons is by far not as precise and reliable. Clearly, the birds have a remarkable capability of way-finding
even from far-distant unfamiliar areas, but they are not at all faultless navigators nor are they always willing to play their part in the experimenters game.
Their performances depend on a great number of internal and external variables and to a considerable degree they depend also on chance. Many years of
accumulating research experience were necessary in order to perceive, analyse
and evaluate these variables. Only when the peculiarities of the pigeons homing behaviour are adequately considered is it possible to produce and interpret
particular experimental interferences in a conclusive way.
3.7.1
Home-Related and Home-Independent Components
of Initial Orientation
It is clear that pigeons released in an unfamiliar area have some idea about the
direction they have to fly home, but also that their initial directions very often
deviate from a direct course towards home in a site-specific and temporally
fluctuating way (e.g. Figs. 2.2, 3.13.4, 3.8, 3.9). Such release biases and release-site biases can potentially be caused by different factors:
1. Navigational error. A priori, it appears very unlikely that a presumed avian
map, extrapolated from one site (home) to a large unfamiliar surrounding
region, represents the relevant spatial reality everywhere in a geometrically
correct way without any distortions and, in addition, that birds master the
geometry in a perfect way. Thus, it is more likely that a displaced bird
makes some site-specific error in determining the direction towards home
3.7 Conclusions and Perspectives
49
rather than that it is able to determine the accurate beeline course from everywhere. Even if referring to two consistently rectilinear coordinates, the
bird would probably produce release-site biases if the coordinates were not
perpendicular to each other (cf. Benhamou 2003).
2. Preferred compass direction (PCD) (Sect. 3.1.1). Very often the initial bearings
are polarized towards a loft-specific compass direction (e.g. Figs. 3.4, 3.8 and
3.9), so that the mean vector obtained at a given site is somewhere between
homeward direction and PCD. Theoretically, a PCD could be an outcome of
the navigational mechanism resulting from regional peculiarities of map
cues and/or from the birds specific strategy to make use of them (see, e.g., the
null-axis hypothesis proposed by Wallraff 1967, 1974a, 1980c, or the magnetic map hypothesis by Walker 1998). More recent findings (e.g. Wallraff
1991a, 2001; Wallraff and Kiepenheuer 1994; Figs. 7.2, 7.3 and 7.8) rather suggest that the PCD reflects a directional tendency that itself is not part or the
result of position determination, but nevertheless modifies initial orientation. Its origin and function are unclear, but similar directional tendencies
have been found in wild birds as well (nonsense orientation; see Sect. 10.1.2
for further discussion).
3. Deflection by landscape features (Sects. 3.4.3 and 8.2). In contrast to PCD influences, such deflections do not produce a regular regional pattern but add
local effects disturbing the regularity. It is difficult to determine the degree
to which topographically caused deflections (e.g. attraction by a nearby village or avoidance to overfly a wooded area) contribute to the release-site biases.
4. After-effect of previous homing flights (Sect. 3.2.2). Especially little-experienced pigeons retain some tendency towards a direction they had flown
while homing last time from another site. While this tendency itself depends on the individual birds history, the very variable degree of its expression may depend also on local conditions at the release site. On the whole,
the effect may interfere more with temporal fluctuations of navigational
signals rather than with spatial biases, but quantitatively its expression is
certainly also site-dependent.
Only the first of these biasing factors is a component of the navigational
home-finding process itself. The other three factors interfere with the pigeons
orientation behaviour independently of the spatial relationship between release site and home site. In a single release, or at a single release site, the contribution of each of these factors to the overall bias can hardly be quantified. Consequently, diagrams showing initial orientation at a given site neither reliably
reflect power and accuracy of homeward navigation nor reliably indicate
site-specific errors in position determination. Diagrams showing pooled bearings obtained at a number of symmetrical releases usually equalize release-site
biases so that the resulting mean vector points quite accurately homewards.
However, the biases are now integrated in a large angular scatter (e.g. Figs. 3.2
and 3.4). This broad dispersion does not realistically indicate the degree of in-
50
accuracy of homeward navigation, because it includes deflections from a

homeward course caused by disturbing influences from outside of the navigational mechanism.
3.7.2
Motivation, Selection and Homing Experience
Old experienced pigeons are usually better at homing than inexperienced
youngsters. Possible reasons for the differences are manifold; without particularly designed experiments, and thus in most actual cases, they cannot be
clearly separated. Again we can enumerate a number of contributing factors
without being able to rank them in a universally valid way:
1. Age (Sects. 3.1 and 3.2). Within the first 34 months of life, the basic steps of
ontogenetic development regarding site fidelity and navigational capabilities are largely completed. Because homing pigeons are rarely maintained
over a longer period without being used for homing experiments, and thus
to become experienced in homing, little can be said about a possible role of
further increasing age without concurrently increasing experience. A few
unpublished data suggest that site fidelity and motivation to home gain
weight in the second summer season when the birds start breeding.
2. Motivation (Sect. 3.1.2). If a bird fails to orient homeward, it is often difficult to distinguish between lacking motivation and lacking ability to home.
Samples of first-flight pigeons usually seem to include individuals with a
weak homing drive, so that the observed performance of a sample is below
the level of possible navigational ability of inexperienced pigeons.
3. Selection (Sect. 3.2.1). Improved performance in later homing flights is
partly due to positive selection of the better motivated and/or better oriented birds during the early flights. It seems, however, that this selection includes (also?) a great deal of chance. If greater distances (>100 km) are included, only a small percentage (some 1020%) of an initially used sample
of pigeons remains after 10 releases.
4. General homing experience (Sect. 3.2.1). Once a pigeon has homed several
times, and also from a greater distance, it is very reliable in homing even from
unfamiliar areas far away from any regions it has ever experienced before.
Also, initial orientation is usually better than in untrained birds, but it is uncertain to what degree improvement reflects improved navigational capabilities, improved motivation and/or selection of individual champions.
5. Local experience (Sect. 3.2.3). Clear improvement due to repeated releases
at the same site is usually apparent only after several such repetitions. However, pigeons apparently learn some local signals without immediately
showing behavioural effects (Chap. 8).
6. After-effect of previous homing flights (Sect. 3.2.2). This aspect has already
been mentioned in the previous section (Sect. 3.7.1) under point (4). No
homing flight can be directionally neutral.
51
Motivation, learning, improved motivation by learning, improved navigation by

learning (by learning what?), motivation-based selection, capability-based selection, chance-based selection: these are all factors involved in practical work
with pigeons which need to be considered in planning and interpreting homing
experiments. Weighting and contributions of these aspects have not yet been
analysed in much detail, because related analyses would require many specifically designed series of simultaneous releases of differently experienced pigeons
in varying combinations which to achieve necessitates huge amounts of statistical material. For an end in itself, such experiments have not exhaustively been
conducted.
3.7.3
Inaccuracy and Variability of Homing Orientation
Homeward orientation in unfamiliar areas is mostly not very precise and, in
addition, it is very variable in time and space, i.e. depending on the geographical region. Temporal changes have been observed within a day, among days,
among years and periodically among seasons. Levels of performance appear to
be correlated with climatic conditions: summer better than winter, Italy better
than Germany (at least over relatively short distances). Initial bearings and
homing speeds appear to vary stochastically among individual birds, suggesting that home-guiding environmental signals are not clearcut but contain considerable inherent noise.
Potential Input Signals Exploitable

for Home-Finding
Having seen the phenomena that we would like to understand, we can now ask
for explanations. The first question in any case of animal orientation concerns
the physical cues according to which the animals orient their activities. In
many cases, the answer is immediately obvious, but in the case of pigeon homing from far-distant unfamiliar areas it is not obvious at all. Our knowledge of
the physical world offers quite a limited number of potential signals. Experience has repeatedly shown, however, that cues are used that we did not consider potentially suitable in advance. The principal part of this book deals with
such a case. The following brief overview implies an anticipating valuation
based on empirical findings which justify selection, sequence and weights of
the sections following thereafter.
4.1
Directional References
The spatial relation between two points in a plane is characterized by direction
and distance. In our case, maintaining the direction from a release site to the
home site may suffice, because the bird can recognize its destination once it
has reached it. Theoretically, it need not be informed about the distance in advance, but it must know the direction. If direct sensory contact to the goal is
impossible, the respective direction can only be defined as an angle to something. In order to keep the angle invariable from the start position to the goal,
the signal of reference should not be taken from the landscape, but should be
as far distant as possible. Two such omnipresent signals are usually available
during daytime: the sun and the geomagnetic vector. Later, it will be shown
that pigeons actually use these references for home-finding. Primarily they apply a clock-controlled sun azimuth compass (Chap. 5). Under overcast skies,
when the sun compass is not applicable, a magnetic compass (Sect. 6.3.2) can
apparently be used as an alternative.
To be able to maintain a direction, however, is not sufficient for homing. The
bird must know which direction it has to maintain in order to reach its home,
i.e. it must know its directional position relative to the home site.
54
4 Potential Input Signals Exploitable for Home-Finding
4.2
Indicators of Position
4.2.1
Motion-Bound Signals
Homing is always a return movement to a site from which an animal has previously made an outward trip on its own or from which it has passively been
transported by man to a site of release. Homing may thus utilize information
collected during the outward journey. This information may be based on measurements of directional and linear motions, either by perception of self-motions or accelerations (inertial navigation; see Barlow 1964) or by utilizing
some external cues (e.g. the sun or the geomagnetic field and/or visual flow of
surrounding structures) or by applying a hybrid system making use of different kinds of input. By integration of these motion-bound signals, the animal
will always be aware of its position relative to the starting point in an egocentric system without paying attention to any external location-specific signals.
Such a method of path integration is widely used by animals, but usually requires an actively performed outward journey enabling recording of self-motions (e.g. Mittelstaedt and Mittelstaedt 1982; Wehner 1992; Etienne et al. 1998;
Etienne and Jeffery 2004). Pigeon homing experiments are always conducted
with passively displaced birds. It has been shown that, at least under this condition, pigeons do not require and certainly do not apply path integration to find
their way home (e.g. Wallraff 1980a, 2000b, 2001). Hypothesized exceptions
concerning very young pigeons only (Wiltschko and Wiltschko, e.g. 1985, 2000,
2003a) are unlikely to exist (Sect. 6.3.4).
4.2.2
Location-Bound Signals
The determination of an animals position relative to home may be based on
information deduced from location-dependent environmental cues. Such information may be collected at any stage of a homing tour, i.e. during the outward journey, at the point of turning back (after passive transport: the release
site) and during the return trip. The use of location-bound signals requires
that the animal has some knowledge about the spatial relationships among the
signals perceptible at different sites. In short metaphoric and anthropomorphic terms: the animal requires a map (Sect. 1.3). Two kinds of map are usually
distinguished (e.g. Wallraff 1974a, 1985, 1991a; Wiltschko and Wiltschko 1987,
1998, 2003a):
1. A topographical map (or mosaic map, familiar area map, possibly a cognitive map) may consist of any visual, olfactory or otherwise perceptible features whose spatial pattern, forming a landscape, has been learned by individual experience. Thus, this kind of map is restricted to an area with whose
4.2 Indicators of Position
55
distinct attributes the animal has become familiar at the home site, in the
course of exploration tours or, in the case of pigeons, by previous homing
flights (Fig. 4.1A, B).
2. A gradient map (or grid map) is applicable also in unfamiliar areas far away
from any previously visited localities. However, it requires the existence of
particularly structured environmental features (Fig. 4.1C, D). They must involve extended spatial gradients of any physical nature, at least two oriented
in different directions, so that each site in a two-dimensional plane is, in the
ideal case, characterized by a unique combination of quantitative values (coordinates). The gradient map is basically unlimited in extension (Fig. 4.1D),
n Fig. 4.1. A Mosaic of landmarks, symbolized by letters, surrounding a birds home site (central
dot). B The corresponding mosaic map (topographical map), which is limited in extent by the
birds range of experience. C Isolines (arbitrary units) of a fictitious gradient field. The line representing the scalar value observed at home is defined as 0, higher values are indicated by solid and
lower values by broken lines. D The birds corresponding gradient map as established by extrapolation of home-area conditions. Note that in C and D, for simplicity and clearness, gradients of
only one variable are shown. For complete site localization, at least two gradient fields are required intersecting at sufficiently large angles. The sections shown are thought to be of different
size; sides of the square in A and B may be at most a few hundred kilometres, those in C and D
1,000 km or more. (Wallraff 1985)
56
but its applicability may be spatially limited due to limited extensions of naturally occurring monotonic gradients (Fig. 4.1C: a bird displaced to the
northwestern or southeastern corner would fly away from home). Also, a
birds map may not include directional changes of gradients or other irregularities that may occur at some distance from home, so that the map does not
everywhere fully coincide with the natural spatial conditions.
Using a gradient map, the animal not only must learn the values marking its
home site, but also must have some information about the directions of the gradients, i.e. the directions towards which the respective scalar values increase or
decrease (cf. Fig. 5.5). This information could be acquired by scanning a field
around home large enough to measure differences against the home values. Alternatively, some signals indicating gradient directions might be available at the
home site itself.
In order to achieve the moderate levels of accuracy of homeward orientation,
as they were observed in pigeons (Sects. 3.4 and 3.7.3), it is not necessary to propose ideally monotonic gradients. By model calculations it has been shown that
a larger number of very noisy gradients can lead to similar levels of performance
as signals deduced from only two fairly reliable coordinates (Fig. 4.2; compare,
e.g., a homeward component of 0.5 achieved with either three gradients providing a probability of a correct upward/downward decision of 3:1=75% or seven
gradients with a proportion 2:1=67% or 20 gradients allowing a correct/false ratio of only 3:2=60%).
n Fig. 4.2. Precision of homeward orientation depending on the number of non-perfect gradients
at sites 100 km from home. Curves result from probabilistic right-or-wrong decisions based on scalar values of 220 arbitrary variables with spatial gradients in varying directions. The probability to
respond correctly to the separate quantities varies between 50% (=1:1 = random) and 75% (=3:1)
as indicated. Results of a model simulating homeward orientation on the basis of fictitious gradients providing information on different levels of reliability. (Modified from Wallraff 1989b)
4.2 Indicators of Position
57
Since pigeons are able to home from far-distant unfamiliar sites, it is obvious
that the most challenging problem is the physical basis of the proposed gradient
map. I shall now enumerate and briefly discuss those constituents of the environment that have been considered to potentially inform a displaced pigeon on
its current position relative to its home site. The sequence roughly reflects the
chronological order in which the factors entered the scientific scenery.
1. Geomagnetic field. It is one of the oldest (Viguier 1882) and still often favoured ideas that birds might use magnetic parameters to determine not
only directions but also positions. In fact, the earths magnetic field provides roughly northsouth-oriented gradients in total and vertical intensity
and in inclination. Empirical findings do not support the hypothesis that
pigeons might deduce positional information from these gradients or any
other geomagnetic signal [as mentioned in Sect. 4.1, however, the earths
magnetic field deserves attention as a directional reference (Chap. 6)].
2. Visual landscape. The simple assumption that pigeons might refer to familiar visual landmarks while returning to their loft is old (e.g. Schneider
1887), certainly correct, but still debated. Landscape features are suitable to
create a topographical map, but unsuitable to guide birds home from an
unfamiliar area whose appearance they have never seen before (Chap. 8).
3. Coriolis force. Yeagley (1947) suggested that the strength of this mechanical
force, which is caused by the earths rotation and varies with latitude, might
be used in combination with a magnetic gradient to form a bi-coordinate
grid. This idea has been out of discussion for many years (e.g. Matthews
1968) and shall not be considered further (Chap. 11).
4. Sun. The sun navigation hypothesis, put forth by Matthews (1953), was experimentally tested and heavily debated in the 1950s and 1960s. Theoretically, if a precise and stable chronometer is available, it is possible to deduce
geographical gradients (coordinates) from the suns orbit (cf. Pennycuick
1960). Experimental evidence, however, stands against the hypothesis that
pigeons make use of this possibility (Sect. 5.2). Also this idea will not be
considered further. However, the suns important role as a directional reference will be inspected more closely in Chapter 5.
5. Atmospheric chemosignals. Volatile trace substances in the atmosphere,
perceived by olfaction, were not among the candidate sources of navigational information until Papi et al. (1971) initiated, on the basis of related
experiments, three decades of intense research and debate on their role in
pigeon homing. Today, it seems that the atmosphere contains the crucial
signals that make homing from remote unfamiliar areas possible. Therefore, the longest chapter deals with olfactory navigation (Chap. 7).
6. Infrasounds. Atmospheric oscillations of air pressure in a frequency range
of some 0.110 Hz, originating at geomorphological structures such as
coasts or mountains, are known to spread over hundreds or thousands of
kilometres. Thus, infrasound waves of different lengths originating from
different sources might constitute a continent-wide grid of gradients or tri-
58
angulation posts, respectively (Hagstrum 2000, 2001). Pigeons have been

shown to be sensitive to infrasounds, provided that their inner ear (cochlea
and lagena) is intact (Kreithen and Quine 1979; Schermuly and Klinke
1990). It has also been shown, however, that bilateral removal of cochlea
and lagena did not impair homing capabilities (Wallraff 1972). In brief abstracts, Schps (1991) and Schps and Wiltschko (1994) report on perforations of the tympanic membrane which at some sites had a deteriorating
and others an improving effect on initial orientation. They conclude from
these data that infrasounds influence pigeon orientation. However, these
cursory reported findings as well as some casual observations reported by
Hagstrum (2000, 2001) are too scanty and non-specific to give infrasounds a
realistic chance of being utilized as indicators of position (see Wallraff
2001). It should also be kept in mind that in the various experiments with
olfactory deprivation (Sects. 7.1 and 7.2), undisturbed operation of the cochlea obviously did not help to find the way home.
7. Electromagnetic sferics. Comparable to the grid of mechanical infrasound
waves, a grid of electromagnetic pulses, originating from lightning discharges, covers the earths surface. As thunderstorm areas show a fairly regular spatio-temporal pattern, and low-frequency radio noise (atmospherics,
sferics) is detectable over long distances, determination of position might be
considered conceivable on the basis of triangulation using the directions, and
possibly other parameters, of such signals. Negative correlations between frequency of sferics and homing speeds or accuracy of initial orientation observed in pigeon races have been described (Dornfeldt 1977, 1996; SchmidtMatthiessen 1981). They do not indicate, however, that sferics are used as
sources of spatial information. It seems more likely that such observations,
together with correlations between speeds in pigeon races and sun-spot
numbers or magnetic activity (e.g. Schreiber and Rossi 1978; Dornfeldt 1996),
as far as significant at all, might indicate influences of varying environmental
(electrical, meteorological?) conditions on the birds physiological state (see
similar phenomena described for laboratory animals and humans, e.g.
Bersani 1999).
8. Gravity. Like the Coriolis force, the force of gravity has a northsouth gradient between poles and equator, but its slope is extremely gentle. Lednor and
Walcott (1984) did not find any indication that pigeons might make use of
this gradient.
In this brief overview, only two of the eight considered potential sources of positional information have been positively labelled for further consideration: the
visual landscape and atmospheric chemosignals. Because of its broad discussion
in the literature, a third factor, the geomagnetic field, will also be inspected in
some detail, although it has earned a negative label. Anyway, magnetism as well
as the sun need to be considered as indispensable directional references.
The Role of the Sun
Kramer (1950, 1952, 1953a) detected that birds can use the sun to determine
compass directions. As the suns position changes over time, its usability for
spatial orientation requires involvement of a clock. Hoffmann (1954) showed
that the birds internal clock can be shifted by a shift of the lightdark cycle
and that this temporal shift results in a corresponding angular shift of the
birds compass. Concurrently, Matthews (1953) developed his sun navigation
hypothesis which proposes that birds use the sun not only to determine compass directions but also to determine their position in relation to their home
site (Chap. 11). For this purpose, demands on chronometry would be much
higher. It was necessary, therefore, to investigate the effect of clock shift in the
context of homing.
5.1
Effects of Shifting the Birds Circadian Clock
Much is known today about so-called circadian rhythms which can be observed
in most organisms from unicellular algae to humans (e.g. Aschoff 1981; Dunlap
et al. 2004). Even in a temporally constant environment, many biological activities fluctuate with a period length of approximately 24 h (circa diem). Under
natural conditions they vary in phase with the daynight cycle. Shifting this cycle by artificial illumination results in a phase-shift of the circadian periodicity.
Schmidt-Koenig (1958, 1961) found that the initial bearings of pigeons that
had been living for several days in a lightdark cycle shifted by 6 h either forward or backward, deviated from those of non-shifted controls by roughly 90
counterclockwise or clockwise, respectively. The effect was similar to that in
stationary training experiments not connected with homing (Hoffmann 1954;
Schmidt-Koenig 1958). In the diagrams summarizing the results of these early
displacement experiments (e.g. Schmidt-Koenig 1965, 1979; Schmidt-Koenig
et al. 1991), the overall mean of the deflections was, however, considerably
smaller than predicted on the basis of pure sun-compass orientation, probably
due to some interfering concomitant circumstances (Sect. 8.1.4). Therefore,
Figs. 5.1 and 5.2 show the results of later experiments, obviously free of additionally interfering factors, in which the predictions were almost exactly met.
Figure 5.1 explains the rationale of clock-shift experiments by means of two
examples. The mean deflection in the whole series of releases (Fig. 5.2) corresponds to the mean angular difference between the suns real azimuth at the
60
5 The Role of the Sun
n Fig. 5.1. In two groups of pigeons the circadian clocks were shifted forward by keeping the
birds in an artificial lightdark regime advanced against the natural day by 6 h. The clock-shifted
birds (open circles) were released, alternately with unshifted controls (filled circles), at two sites
about 30 km north and south of home. Left-hand diagrams show theoretically expected flight directions on the assumption of perfect position determination and use of a sun-azimuth compass.
If the birds, at time T (here 10:00 A.M.), select an angle T relative to the sun ST, they fly in the direction towards home H. At time T, the shifted clock of the experimental birds shows T+6 h. At
that time (here 4:00 P.M.) the sun would be at position ST+6; H would be reached by selecting an
angle T+6. By keeping this angle relative to the actually visible sun ST, the birds would achieve
the course H. Under the given conditions (summer in southern Germany), H is approximately
120 left from H (like ST from ST+6=azimuth). Right-hand diagrams show actually observed vanishing bearings with their mean vectors. The experimental pigeons were as well oriented towards
H as the controls were towards H. (Wallraff 1988a)
time of a pigeons release and the expected azimuth according to the pigeons
phase-shifted time scale. When recalculated on the basis of these angular differences (azimuth), the bearings are as well homeward oriented as those of the
non-shifted control birds (Fig. 5.2, C versus A). These results strongly suggest
that pigeons apply a time-compensated sun-azimuth compass when they decide on a direction intended to guide them home.
Obviously, the birds refer only to the suns azimuth, i.e. to its angular position
along the horizon, and do not feel disturbed by differences in altitude between
the observed and the expected sun, even if these differences are considerably
large (Fig. 5.3). Angular scatter around the expected direction remains in the
order observed in the controls (cf. Fig. 5.2A) independently of whether or not
5.1 Effects of Shifting the Birds Circadian Clock
61
n Fig. 5.2. Individual vanishing bearings (peripheral dots) and mean vectors per release (arrows)
in 24 releases from pairwise opposite sites 2258 km from a loft at Andechs in Bavaria. B, C and E, F
Experimental pigeons with clocks phase-shifted 6 h forward; A, D control pigeons coming from an
artificial lightdark regime in phase with the natural day. AC Angular deviations from the direction towards home (H); DF angular deviations from the mean direction of control birds per release (M), so that vectors of A are superimposed upwards in D. Data of clock-shifted birds are
given twice, as directly observed (B, E) and recalculated by subtracting azimuth (individually calculated according to site, date and time of day) from each bearing (C, F). Mean azimuth of the 24 experiments (119) is shown in B and E. Numbers refer to the second-order mean vectors (direction
and length) derived from the 24 single-release vectors; they correspond to centres of the 95+99%
confidence ellipses as drawn in the diagrams. (Neuss and Wallraff 1988)
62
n Fig. 5.3. A Altitudes of the sun in the area and approximately at the date around which the re-
leases of Fig. 5.2 were conducted. Solid line Real observable sun; broken line expected sun according to the 6-h forward-shifted time scale of the clock-shifted pigeons (numbers in brackets at the
abscissa). B The 302 recalculated bearings as shown in Fig. 5.2C in relation to differences in expected and actually observed sun altitude. (After Neuss and Wallraff, unpubl. data)
the observed sun is at an altitude similar to that corresponding to the birds

subjective time scale. Even if the sun is some 30 or more higher or lower than
it should be at that time of the day, the pigeons respond in a normal, apparently
only azimuth-dependent way.
In the experiments shown in Fig. 5.2, the pigeons do not exhibit any uncertainty that might arise from the non-fit between the shifted sun compass and a
non-shifted magnetic compass. We shall see later that this is not always so
(Fig. 6.2) and that, under certain circumstances, it may happen that angular
deflections caused by clock shift are smaller than expected on a pure sun-azimuth basis (Sect. 8.1.4).
5.1 Effects of Shifting the Birds Circadian Clock
63
n Fig. 5.4. An equally sized phase shift of the circadian clock causes differently sized angular de-
flections. Four releases at different sites were conducted in summer near Auckland, New Zealand, at a latitude of 37S. A Untreated control pigeons; B pigeons clock-shifted 4 h forward and
released in the morning or in the afternoon with azimuth (cf. Fig. 5.2) varying between 35 and
41; C pigeons clock-shifted 4 h forward and released during the forenoon while azimuth varied
between 120 and 129. Vanishing bearings are drawn with respect to their deviations from home
(upwards), the releases (2038 km from home) distinguished by symbols, arrows giving second-order mean vectors. (Modified from Wiltschko et al. 2000)
The sun moves along its tilted orbit with a constant velocity of 15/h. The
change of sun azimuth along the horizon over the course of a day, however, is
non-linear and highly variable depending on latitude and season. The pigeons
are obviously aware of this non-linearity. Otherwise, the recalculated bearings
in Fig. 5.2C, based on azimuth differences varying between 88 and 136, would
not have led to such a good coincidence with the control diagram A. More directly, Wiltschko et al. (2000) compared the effects of an equally sized clock
shift during morning and evening hours with those during noon hours at a
lower (southern) latitude (Fig. 5.4). The angular deflections were clearly different in amount in accordance with the different speeds of sun-azimuth movement during the respective time of the day. Should the pigeons not consider
these varying speeds, steadily shifting initial bearings in the course of the day
during prolonged releases of untreated birds should occur, but these have not
been observed (Wallraff 1971, unpubl. data; Keeton 1974a). Continuous adjustment of the pigeons responses to the seasonal changes of the suns path appears sufficiently good, but is certainly not completely perfect (particularly
not at latitudes close to the equator; see Schmidt-Koenig et al. 1991). In the
southern hemisphere, the pigeons refer correctly to the suns counterclockwise
movement, so that a forward clock shift results in a clockwise deflection
(Fig. 5.4; Wiltschko et al. 1998).
64
5.2
Linkage Between Sun Compass and Map
Most importantly, the clock-shift experiments reveal that the sun is a substantial component of the pigeons homing mechanism, but that its function within
this mechanism is restricted to a directional reference, while the determination of positions requires additional means. If the birds would use the sun for
this latter purpose, clock shifts would not induce analogous rotational shifts
everywhere, but deflections varying with the birds position relative to home
(so that Fig. 5.2C would not mimic normal homeward orientation as in
Fig. 5.2A). Thus, the sun is not used for the entire navigational process, including position determination, as might be theoretically possible (Matthews 1953,
1955, 1968; Pennycuick 1960). The sun navigation hypothesis was discussed
decades ago in more detail and has been falsified also by means of several
other findings and arguments (e.g. Hoffmann 1965; Schmidt-Koenig 1965,
1979; Keeton 1974a; Wallraff 1974a).
As, in the context of homing, a compass can be usefully applied only if the animal knows which direction it has to select in order to reach its home, an appropriate linkage between the animals map and its compass must have been established during the long-term stay at the home site. Figure 5.5 illustrates the
n Fig. 5.5. Map and compass. A bird using a gradient map must know the compass directions to-
wards which the scalar values of some physical variables x and y regularly increase and decrease.
Using the moving sun for this purpose, it must know, for a given hour of the day, the angular relationship between the sun and the gradients. With respect to the geomagnetic field as a reference
(Nm), the angular relationship is temporally stable. H Home site; P current position of a bird. Azimuth positions of the sun during the course of a day are valid for 50N on 24 May and 20 July
5.2 Linkage Between Sun Compass and Map
65
requirements that must be met to make Kramers (1953b, 1957) map-and-compass concept functional. In a simplifying basic-principle manner, it is assumed
that a birds gradient map (Sect. 4.2.2) makes use of two arbitrary physical quantities, x and y, which form a grid of gradients usable as coordinates. [With some
modification, the scheme is applicable also to gradients of three or more ratios
(Sect. 7.6.1; Fig. 7.26).] At the home site, H, it measures the values xH and yH.
When displaced to site P, it measures xH+x and yH+y. This measurement
alone would not help home-finding. In addition, the animal needs to know towards which direction x and y increase and decrease. If x and y are not vectors
that themselves include a directional component (as, for instance, the geomagnetic vector), and if their gradients are too gentle to be measurable at P itself or
within its closest vicinity, the displaced bird can, at best, measure only scalar values of x and y and must bring some pre-established knowledge with it about the
compass alignment of the gradients. The bird must know that x=12.5 units
together with y=+7.5 units indicates that it has to fly an angle of 90 left of the
sun at noontime and away from the sun at 5 P.M.
If the alignment of the gradients is not constant everywhere on earth, it must
be learned in some way at the home site. Wiltschko et al. (1976, 1984) have in fact
shown that the directional linkage between map factors and sun compass is not
genetically fixed. Pigeons were kept over more than 2 months in a clock-shifted
state. In the overlapping light phases of the natural and the artificial lightdark
(LD) regime, they not only could fly freely at the home site but also were, in addition, intensely trained to home over distances of up to 35 km (probably, this
training was not necessary to reach the observed effect). Independently of
whether the birds had lived in this phase-shifted LD condition constantly since
fledging or were transferred from the natural day as experienced adults, the
n Fig. 5.6. A A normal clock-shift experiment (cf. Fig. 5.1) with unshifted controls (filled sym-
bols) and pigeons phase-shifted 6 h forwards (open symbols). B Open symbols refer to pigeons
that were living under sun at home and had been given exercise flights while their circadian
clocks were permanently phase-shifted 6 h backwards. If such birds were, a few days before release, brought into natural day conditions in a closed room, they behaved as if they were shifted
6 h forwards (filled symbols). (Modified from Wiltschko et al. 1976)
66
long-term clock-shifted pigeons flew towards the same directions as the nonshifted controls when released at a distant site (Fig. 5.6B, open symbols). After a
short-term re-shift into natural LD, however, without open-field exposure, they
behaved like short-term clock-shifted birds coming from natural LD (Fig. 3.6B,
filled symbols).Obviously,the pigeons labelled the observed positions of the sun
in relation to the spatial coordinates with different times of the day (6 instead of
12, 9 instead of 15, 12 instead of 18 h in Fig. 5.5; see also Fig. 1 in Wallraff 2004).
While staying within this system, everything functioned in a harmonic way, but
with the circadian clock synchronized with the natural day, the learned calibration no longer fitted.
Clock-shift experiments function perfectly with first-flight pigeons that
know only the immediate vicinity of the home site (e.g. Wiltschko et al. 1994).
Indirectly, the findings suggest that the pigeons are in some way able to recognize the directions of the assumed gradients (or some equivalent) without
leaving the home site, or its very close vicinity, and to memorize their angular
relationship to the suns daily path. Later, we will see that this calibration process works adequately only while the birds are exposed to natural wind conditions (Sect. 7.5.1). Angular calibration between the suns path and the geomagnetic field may take place in parallel (Wiltschko and Wiltschko 1990, 1998,
2003a), but this link-up between two compasses alone would not constitute a
home-finding mechanism. Decisive is the appropriate linkage between map
and (solar or magnetic) compass.
5.3
Orientation Under Overcast Skies
The question is very simple: what do the pigeons do when they are released
while the sun is obscured by clouds? The answer is less simple, as the related
data are notably inconsistent. Figure 5.7 gives three examples. In example A,
the pigeons departed homeward-oriented under sun but not under overcast.
The pigeons used had been allowed to fly ad libitum at the home site under all
conditions of weather. Pigeons kept in a similar way behaved similarly disoriented under overcast in example B. Another group of pigeons, however, released simultaneously, was homeward-oriented. They had been allowed to fly
at the home site only under complete overcast and had been confined in a
closed room while the sun was shining. The birds shown in example C, apparently homeward-oriented under overcast, could fly with and without the sun
around the loft, but were often forced to fly under bad weather conditions with
the sun invisible to humans. Under a complete cloud cover, clock shifts had no
deflecting effect. A fourth example is given below in Fig. 6.1C; it shows good
home orientation under overcast in pigeons that could fly ad libitum at home
without enforced flying under overcast. These experiments were conducted in
three different geographical regions characterized by different weather patterns.
The lacking effect of clock shift in Fig. 5.7C indicates that the pigeons determined their directions without referring to the sun compass. On the other
5.3 Orientation Under Overcast Skies
67
n Fig. 5.7. Examples of initial orientation under overcast conditions. Filled symbols Sun position
undeterminable (for humans); untreated pigeons with free-flight conditions at home under various weather conditions. Open symbols, in A, sun clearly visible, pigeons corresponding to those released under overcast skies; in B, sun position undeterminable, pigeons at home prevented from
seeing the sun and allowed to fly around only under overcast skies; in C, sun position undeterminable, pigeons were clock-shifted 6 h forward, otherwise corresponding to the black controls. A For
either condition, releases on six different days are summarized; home loft at Osnabrck, Germany
(Wallraff 1966b); B, C home loft at Ithaca, New York, USA; differently treated pigeons released simultaneously, one release from 66 km north (Wiltschko et al. 1987c) and one from 164 km east
(Keeton 1969)
hand, Schmidt-Koenig (1958) reported two releases in which clock-shifted pigeons were deflected under overcast conditions, which hindered the human
observer from localizing the sun. Also, in stationary training experiments
dealing with sun-compass orientation, he found deflections in corresponding
tests under overcast conditions, whereas Budzynski et al. (2000) report disorientation even without clock shift. The confusion is completed by a series of releases at familiar sites in which clock-shifted pigeons flew neither homeward
(as the controls did) nor deflected in the rotational sense according to the clock
shift, but appeared disoriented (Benvenuti et al. 1996).
The inconsistency of the results obtained under overcast skies is somewhat
puzzling. In part, it may be explainable by the fact that the conditions cannot
be clearly standardized. If a human observer feels unable to localize the sun, pigeons may mostly be unable to do so as well, but not necessarily in all cases. Extended flying experience under overcast skies seems to improve the birds
readiness to apply a non-solar, most likely a magnetic compass (Sects. 6.1.1
and 6.3.2). It is surprising that the pigeons do not always make use of this possibility when the sun is invisible (see also Keeton and Gobert 1970), even if they
had experienced overcast skies quite often at home and although very young
pigeons, according to Wiltschko and Wiltschko (1980, 1981) and Wiltschko et
al. (1981), use a magnetic compass before they learn to apply a sun compass. It
seems that the sun compass, once it is established, is so predominant over the
magnetic compass that the latter is not always immediately consulted when the
first fails to operate and that the pigeons do not, or do not always, feel disturbed when the shifted sun compass disagrees with the magnetic compass
(see Figs. 5.2 and 5.6, but also Fig. 6.2).
68
Puzzling as they are, the inconsistent results obtained under overcast skies do
not concern the substantial role of the sun compass in pigeon homing, but
merely the question of whether or not, or when and when not, it is replaced by
another (presumably magnetic) compass while the sun is invisible behind clouds.
5.4
It appears well documented and accepted by all researchers in the field that pigeons in their homing flights make use of a map-and-compass system in which
a time-compensated sun-azimuth compass is the primary mechanism providing the compass component. Deflections caused by clock shift make it clear
that positional (map) information and directional (compass) information
come from different environmental sources, which are linked to each other by
a learning process at the home site. Otherwise, i.e., if a bird could directly perceive, at each current position, the directions of gradients (x and y arrows in
Fig. 5.5 and not only scalar x and y values), a separate compass system would
not be necessary (see also Wallraff 1974a, 1991a, 2004).
The sun is a substantial part of the pigeons homing mechanism, but it does
not tell the animal where it is and in which compass direction it has to fly in order to reach home. Once the bird has deduced this direction using other
sources of information, it can pursue the appropriate course using the sun.
In this monograph, the sun compass itself (which has been investigated
also in many other animals) is not the focus of interest. It is merely seen here
as a component of the homing process. Many details about the avian sun
compass mechanism itself, including its operation near the equator, at high
latitudes and during night-time, have been reported and discussed by
Hoffmann (1965), Schmidt-Koenig (1979), Wallraff (1981b), Schmidt-Koenig
et al. (1991), Wiltschko and Wiltschko (1999a) and others. Its ontogenetic development in pigeons and included learning processes within and apart from
homing tasks have been investigated by Wiltschko and Wiltschko (1980,
1981, 1990, 1998) and Budzynski et al. (2000).
For future research, particularly in the context of homing, I see only two
more peripheral aspects. First, orientation under overcast skies requires clarification (Sect. 5.3), which is primarily important, however, with respect to the
operation of non-solar compass orientation (Sect. 6.3.2). Second, releases of
clock-shifted pigeons may further be used as tools to elucidate the role of the
visual landscape in pigeon homing (Sect. 8.1.4).
The Role of the Geomagnetic Field
Theoretically, the magnetic field of the earth could be used, like the sun, in a
two-fold manner: for compass orientation and for the determination of position. Due to its northsouth gradient in total intensity, vertical intensity and inclination (dip angle), it could provide information on latitude. The necessary
second coordinate would be less easily accessible, but could be derived, in principle, either also from the geomagnetic field itself (at least in some regions) or
from a combination with other parameters such as the Coriolis force or the sun
(see, e.g., Yeagley 1947; Walcott 1982, 1991; Gould 1985, 1998; Wiltschko and
Wiltschko 1995; Walker 1998, 1999; Wallraff 1999; Walker et al. 2002). Thus, any
experiment concerned with a possible role of magnetism in pigeon orientation
focuses on four alternatives: is a compass affected,a map,both,or none of them?
Since a sense organ for magnetoreception so far has not clearly been identified (Sect. 6.3.5), the classical way of examining the importance of a given sensory input by interfering with the related sense organ or its neural connections
cannot be used (or would include a test on whether magnetoreception might be
affected; e.g. Beason et al. 1997). Thus, experiments are restricted to artificial alterations of magnetic fields to which the birds are exposed and to correlations of
the birds behaviour with naturally occurring magnetic irregularities.
6.1
Effects of Artificial Magnetic Fields
6.1.1
Magnets or Coils During Flight
Various experiments have been conducted in which pigeons were released
with permanent magnets attached to their wings, heads and/or backs (Yeagley
1947, 1951; Gordon 1948; Matthews 1951; van Riper and Kalmbach 1952;
Keeton 1971, 1972; Ioal 1984, 2000; Wallraff et al. 1986a; Moore 1988; for more
references see Keeton 1971, ref. 7) or equipped with a pair of Helmholtz coils
around their heads (Walcott and Green 1974; Walcott 1977; Visalberghi and
Alleva 1979; Lednor and Walcott 1983; Papi and Ioal 1986). Taken together,
these experiments seem to provide evidence that under certain conditions
magnets can confuse homing pigeons (Keeton 1971). This cautious sentence
implies that confusion does not occur consistently nor under clearly defined
conditions.
70
6 The Role of the Geomagnetic Field
n Fig. 6.1. Initial orientation of pigeons under sun (A, B) and under overcast (C, D), respec-
tively, with magnets (B, D) or brass bars (A, C), respectively, attached to their heads and wings. A,
B Simultaneous releases of inexperienced experimental and control birds at four symmetrical
sites 2027 km from their loft in central Italy. C, D The same birds, after having completed flights
from all four sites used in A and B, now released at four unfamiliar sites 4043 km distant from
home. (Modified from Ioal 1984)
Under sunny conditions, the pigeons appeared only exceptionally confused

by magnets or coils, albeit some moderate increase in angular dispersion of
initial bearings was frequently observed (see Fig. 8 and Table 3 in Wallraff
1983). A typical example is shown in Fig. 6.1A, B. Some of the exceptional
larger effects concerned very young first-flight pigeons (Keeton 1971, 1972) or
pigeons that had never seen the sun during the given time of day (Wiltschko et
al. 1981). The few other exceptions remain unexplained (Keeton 1971, 1972).
Today, it seems possible that these results belong to the category of stress-induced effects discussed below (Sect. 6.3.1). Even in the exceptional cases, only
initial orientation was affected. As a rule, i.e. with a few minor unclear exceptions, homing rates and homing speeds were not reduced by experimental alterations of the magnetic field to which the pigeons were exposed, even if this
field fluctuated with the wing beats or oscillated at higher frequencies or was
disturbed already during the outward tour from home to release. Thus, experimental evidence is against the hypothesis that pigeons might require the geomagnetic field to determine their position with respect to the home site. Apparently, magnetic cues do not even play a subordinate, mostly redundant role
6.1 Effects of Artificial Magnetic Fields
71
which might become visible when the predominating olfactory cues are excluded (Wallraff et al. 1986a; Sects. 7.1 and 7.2).
Under overcast skies, in contrast, disorientation or false orientation caused
by magnets or coils seems to be the rule, but cases of no-effect were also observed. These cases are mainly assembled in a publication by Moore (1988) in
which release protocols left by the late W.T. Keeton were evaluated. The overall
result did not confirm the earlier findings published by Keeton (1971, 1972)
himself. This may raise doubts about the weight of these earlier findings, but it
cannot be taken for granted that the conditions were always fully comparable
(see Appendix).
Firm conclusion can only be drawn if (1) control pigeons are well oriented
under overcast and (2) with sufficient certainty the release area is unfamiliar
to the birds. Both requirements were most reliably met in the experiments
shown in Fig. 6.1C, D. The contrast to the bearings observed under sun (D versus B) suggests that compass orientation was concerned and not the determination of position, which otherwise should have led to disturbed behaviour
also under sun. In a later series of releases under overcast, Ioal (2000) obtained results analogous to those in Fig. 6.1C, D, but with the difference that the
controls were considerably less clearly homeward-oriented [thus adding to the
inconsistent results under overcast (Sect. 5.3)].
Surprisingly, even in those experiments in which the attached magnets clearly
disturbed initial orientation (Fig. 6.1D), they did not noteworthily affect the
homing speeds, although the cloud cover was spatio-temporally quite persistent
and landmark orientation hardly possible (Ioal 1984). An explanation for this
finding is lacking (eventual appropriate evaluation of magnetic signals which
still varied depending on the direction of flight; or eventual localization of the
sun through the clouds?). It is, therefore, not clear whether these results really
prove the use of a magnetic compass or whether they also reflect a stress-connected effect of magnetic oscillations as discussed below (Sect. 6.3.1).
Such an effect could hardly have been involved in a series of experiments suggesting the use of a magnetic compass most clearly. Wiltschko and Wiltschko
(2001) combined bar magnets attached to the pigeons backs with clock shifts
(cf. Sect. 5.1). In each of the 12 releases, the birds carrying magnets showed a
larger angular deflection than those carrying merely brass bars (Fig. 6.2). The
median deflection of the latter group reached only 60% of the expected angle
(=azimuth; see Fig. 5.2), whereas the birds with magnets reached 89% (but were
still significantly below 100%; P<0.01). These findings indicate that the pigeons
do not always neglect the divergence between shifted sun compass and nonshifted magnetic compass, as they apparently did in the experiments described
above (Figs. 5.2 and 5.6). Obviously, the birds without magnets made some compromise in the releases shown here, but not in those shown there. An attempt to
interpret this discrepancy will be postponed until the role of a third potentially
interfering factor, the visual landscape, is discussed (Sect. 8.1.4). In the present
context, it is important to note three points: first, as the magnet birds were well
oriented towards a reasonable direction (and with even smaller angular disper-
72
n Fig. 6.2. Clock-shift experiments with pigeons carrying bar magnets (filled symbols) or brass
bars (open symbols) on their backs. On the abscissa is the deviation of the mean vector of vanishing bearings per treatment group and release from the mean direction of a simultaneously released group of non-shifted control birds. Numbers refer to vector length a. The ordinate refers
to 12 individual releases at 7 different sites 3882 km (most about 40 km) from their loft in Frankfurt am Main, Germany. Clock shifts were 6 h forward (triangles) or backward (squares). Approximate ranges of azimuth (cf. Figs. 5.1 and 5.2) are indicated by hatched areas. Most of the pigeons were old veterans with considerable homing experience from various sites in the
surrounding area. (Data from Wiltschko and Wiltschko 2001)
sion than the clock-shifted controls), the magnets obviously did not induce
non-specific general disturbance, but, on the contrary, reduced an interfering influence; second, determination of position was not affected, although, third, the
bar magnets glued onto the pigeons backs were apparently sufficient to set the
magnetic compass out of function [which requires less delicate measurement
than a potential map receptor (Sect. 6.3.3)].
Another kind of experiment, with analogous results in two countries (Walcott and Green 1974; Visalberghi and Alleva 1979), suggests even specifically
the use of a certain kind of magnetic compass. Pigeons were equipped with a
pair of Helmholtz coils around their heads which induced a fairly vertical magnetic field of about 0.6 Gauss. Under sun, little effect was observed. Under overcast, again little effect was observed when the north-seeking pole of the artificial field pointed downward, thus at least doubling the natural vertical
component, but leaving the magnetic vector axis northward-downward. However, with the polarity reversed (north-up), the pigeons flew in roughly reversed directions (Fig. 6.3). In this case, the resulting magnetic field in the
birds heads was considerably attenuated and its axis pointed probably upward
73
n Fig. 6.3. Vanishing bearings of pigeons with coils around their head under overcast in North
America (A, B) and Italy (C, D). The north-seeking pole of the artificially produced magnetic
field pointed downward in A and C and upward in B and D. Vectors with symbols indicate means
of individual releases; in A and B, four releases at a site 68 km west and one (filled triangle) 92 km
west of home; in C and D, at three sites 2345 km east of home. Heavy arrows indicate mean vectors from the pooled bearings as drawn at the periphery (in A and B, without distinction according to different releases). (Data from Walcott and Green 1974; Visalberghi and Alleva 1979)
in northerly directions instead of downward. Under better controlled laboratory conditions, a reversed vertical component induces a reversed directional
preference in caged migratory birds, indicating that they use a magnetic inclination compass defining north as axis downward and ignoring polarity of the
magnetic vector (Wiltschko and Wiltschko 1972, 1995; Fig. 6.6). An interpretation of Fig. 6.3 along this line is, at least, suggestive.
6.1.2
Magnetic Fields Before Release
A considerable number of experiments have been conducted in which magnetic pre-release treatments affected, or failed to affect, initial orientation of
pigeons. Presently, it seems that any influences, if occurring at all, concerned
the pigeons readiness to respond in the usual way rather than their navigational mechanism. Nevertheless, in order to substantiate this conclusion, it is
necessary to briefly review also these investigations. It is important to note that
all the observed effects were restricted to initial orientation, while homing performances remained unaffected or were not recorded. Thus, there is no indica-
74
tion that any of the magnetic treatments might have interfered with an indispensable homing mechanism using magnetic information collected during
the outward journey (Sects. 4.2.1 and 6.3.4).
1. Transport in a static magnetic field varying at irregular intervals. Considering that pigeons might not use a map system based on location-bound signals
but instead a path-integration system making use of a magnetic compass
(Sect. 4.2.1), available inputs for such a compass were disturbed by placing
the pigeons, during the outward journey, in an artificial magnetic field. Two
pairs of Helmholtz coils produced a roughly horizontal, largely inhomogeneous field of about 1 or 1.4 Gauss in eight possible directions with respect
to the car. The directions were randomly altered at intervals varying between
0.5 and 9 min. In addition, the container with the pigeons was ventilated with
synthetic air and rotated at fairly high speeds (540/s; sense of rotation irregularly varying) within the artificial plus the natural magnetic field until arrival at the release site. The birds were released on the following day. In ten
experiments performed in Germany (distances 15300 km), no differences
were observed in initial orientation, homing performance and distribution of
recoveries between pigeons treated that way and untreated controls (Wallraff
1980a). Some slight differences (but no breakdown of homeward orientation
and homing) observed in Italy (Wallraff et al. 1980) need not be explained on
the basis of lacking magnetic en-route information, but can be more easily
understood on an olfactory basis (Sect. 7.4.1).
2. Transport in a magnetic field indicating reversed directions. Specific false
magnetic fields during the outward journey were applied in two series of experiments in which the pigeons were released shortly after transportation.
In a first approach, using Helmholtz coils inside a car, the horizontal component of the magnetic field was roughly reversed while a fairly straight
route towards a release site either in the north or in the south was driven
(Wiltschko et al. 1978; Wiltschko and Wiltschko 1978a, 1985). In a second
approach, Kiepenheuer (1978a,b) reversed not the horizontal but the vertical magnetic component. Provided that pigeons, like other birds, possess an
inclination compass, a reversal of the dip angle (downward towards south
instead of north) causes reversed directional information too (see Fig. 6.6).
This method has the advantage that the false information remains stable independently of the direction in which the vehicle is driving.
Neither method of intended compass reversal during transport caused a
reversal of the pigeons initial bearings, as might be expected if a path-integration system (route reversal) (Sect. 4.2.1) had been at work. Only some
deterioration of initial orientation was found (initial disorientation with the
first method in some releases, mostly restricted to very young pigeons;
slight angular deflections with the second method). No deterioration in
homing speed or homing rate was observed. Thus, independent of the causation of the sometimes occurring initial effects, there is no indication that
the ability to find the way home was affected.
75
3. Oscillating magnetic field before, during and/or after transport. Many experiments have been conducted with pigeons that had been placed, before release, in the centre of three pairs of Helmholtz coils producing a semi-chaotically oscillating magnetic field (Benvenuti et al. 1982; Papi et al. 1983; Ioal
and Guidarini 1985; Ioal and Teyssdre 1989). In most of these experiments,
the intensity produced by each pair of coils varied sinusoidally, with a maximum of 0.6 Gauss and with a period of 15, 23 and 29 s per pair. Independently
of whether the treatment was given for a few hours at the loft site before transport, during transport or at the release site thereafter, the treated pigeons
were in most cases considerably weaker in initial homeward orientation than
the untreated controls. They either behaved disorientedly or switched to the
loft-specific preferred compass direction (PCD) (Sects. 3.1.1 and 3.7.1; example: Fig. 6.4A, B). Some differences in efficiency were found depending on the
wave form of the variations (sinusoidal, rectangular, triangular) and on the
frequency. Homing performance was never significantly affected by any kind
of application of an oscillating magnetic field, not even in experiments in
which this field, produced by coils attached to the pigeons heads, was acti-
n Fig. 6.4. Effects of magnetic oscillations or complete darkness during transport on subsequent
initial orientation. Releases at four (AC) and three (DF) sites 2331 km roughly east of their
loft near Pisa, Italy (homeward directions 245320). B, C Transport in an irregularly oscillating
magnetic field; E, F transport in a light-proof dark box. In C and F, the birds were injected, before
transport, with the tranquillizer promazine. In each release three differently treated groups participated (AC and DF, respectively, releases distinguished by symbols). Second-order mean
vectors are shown as heavy arrows. (Modified from Luschi et al. 1996)
76
vated not only during the outward journey and at the release site (where initial bearings were made worse), but also during the flight home (Papi and
Ioal 1986).
Strikingly, pigeons impeded in smelling environmental odours during
pre-release exposure to an oscillating magnetic field did not show the otherwise typical worsening of initial homeward orientation (Wallraff et al. 1986b;
Papi and Ioal 1988). Moreover, sensitivity to this kind of treatment was
found only in Italian pigeons, but not in birds originating from some German
stocks, even if they were raised, from fledging time onwards, in Italy and then
tested in Italy together with Italian-stock birds (Benvenuti and Ioal 1988).
Also these findings do not suggest that the observed responses to magnetic
fields reflect disturbance of a mechanism that is generally applied by pigeons
to find their way home. There is no indication that this mechanism might be
basically different in pigeons of different strains or from different countries
(Kiepenheuer et al. 1979; Benvenuti et al. 1990a).
However, differences seem to exist, and are more likely to exist, in the birds
sensitivity thresholds with respect to potential stress factors. Handling in the
course of a release experiment, such as crowded confinement of many pigeons in a crate or transporting them in darkness, appears to involve such
factors (Del Seppia et al. 1996). To some degree, the birds opioid system is
able to compensate stressing influences, but specific opiate antagonist drugs
as well as magnetic variations inhibit this compensation and thus obviously
prevent the generally excited birds from immediately focusing their interest
in flying home (Papi and Luschi 1990; Papi et al. 1992; Papi 1995; Luschi et al.
1999; see also Kavaliers and Ossenkop 1986; Del Seppia et al. 1995). Accordingly, if the stress level was reduced by injection of a tranquillizer, the otherwise drastic effects induced by either magnetic oscillations or transport in
darkness (Sect. 6.3.1) were no longer observed (Fig. 6.4). At least from a certain age onward, pigeons of the tested German strains may be less easily
stressed and therefore need no compensation. As far as was tested, even the
pigeons initially showing disturbed orientation behaviour following magnetic treatment were not impeded in finding their way home. For further discussion see Section 6.3.1 below.
4. Near-zero magnetic field during or after transport. When pigeons were transported to the release site in an iron container in which the magnetic field
strength was reduced to 5103 of its natural value, their initial bearings were
not oriented homewards, while those of pigeons transported simultaneously
in otherwise equivalent containers made of aluminium clearly pointed towards home (Papi et al. 1978b, 1980b; Benvenuti et al. 1982). Similar effects
were achieved when the birds were exposed to a near-zero magnetic field produced by three pairs of Helmholtz coils over 2.5 h at the release site (Ioal and
Teyssdre 1989). Homing performances were not reduced by either treatment. There are indications that these findings also belong to the stress complex (Del Seppia et al. 2000).
6.2 Geomagnetic Irregularities
77
5. Exposure to strong magnetic fields before transport. In various head tissues of

pigeons and other birds, magnetite crystals or other ferromagnetic materials
have been found (for review see Kirschvink 1989; Able 1994; Wiltschko and
Wiltschko 1995; Kirschvink et al. 2001). Considering that such magnetic domains might possibly be used for purposes of orientation (Sect. 6.3.5), attempts were made to change the alignment of the material or of its magnetization by exposing the pigeons heads to strong magnetic fields before
displacement. In one of three investigations, no influence of such treatments
was found (Walcott et al. 1988), but in the other two at least some effect on
initial orientation was indicated (Kiepenheuer et al. 1986; Beason et al. 1997).
The effects were not dramatic; homeward orientation was not abolished.
Homing speeds were not always recorded, and, if they were, they were not affected. Homing rates were apparently not reduced. The causal path of interference is unknown. If map information would have been affected (assumption by Beason et al. 1997), it could have played a marginally redundant role
in the homing process at best.
6. Long-term magnetic fields at the home site. One publication (Wiltschko et
al. 1983) did not report on short-term magnetic treatment in immediate
connection with a certain release, but on long-term exposure to an artificial
field at the home site. The experiments aimed to test whether the sun compass developing in adolescent pigeons might be calibrated by their magnetic compass. Probably because the expected compasscompass calibration was intermingled with the calibration process linking compass
elements with map elements (Sects. 5.2 and 7.9.2), the authors were unable
to interpret observed deflections of initial bearings in a comprehensible
way and concluded their study by stating complexity (see Appendix).
6.2
Geomagnetic Irregularities
6.2.1
Spatial Magnetic Anomalies
Peculiar or seemingly peculiar orientation behaviour of pigeons released at
magnetic anomalies has repeatedly been described (for reviews and references
see Wallraff 1983; Walcott 1991; Wiltschko and Wiltschko 1995). Due to the
anyway very variable patterns of initial orientation (Chap. 3), it is difficult to
verify, in each particular case, whether the observed peculiarity (release-site
bias, disorientation) has anything to do with the magnetic condition at a given
site. At best, some suggestive correlation can be stated which does not allow
conclusions about causal connections. Moreover, Walcott (1992) has shown
that an anomaly may influence pigeons from one loft, but not those from another loft, and that an anomaly may be influential in some years, but not in
another year. Similar differences in orientation can be observed also at mag-
78
netically non-peculiar release sites with pigeons from different lofts or in different years (e.g. Wallraff 1959b, 1970a, 1986, 1993; Schmidt-Koenig 1963a). I
do not see that the studies on possible effects of magnetic anomalies suggest
the use of a magnetic map (for a more detailed discussion see Wallraff 1983).
In a recent thorough investigation next to and within an area of magnetic
anomaly in Germany, Wiltschko and Wiltschko (2003b) have not found any indication of a specifically disturbed initial orientation.
6.2.2
Temporal Magnetic Fluctuations
Correlations have been reported between solar activity (sun spots) and performances in pigeon races (Yeagley 1951; Schreiber and Rossi 1978; Dornfeldt
1996). Also, correlations were found in some cases, but not in others, between
geomagnetic activity (K index) and angular variations between daily means of
initial bearings of pigeons that were released many times in succession at always
the same site (Keeton et al. 1974; Kowalski et al. 1988). At some sites, mean deviations of bearings from the home direction increased with increasing magnetic
activity, while at other sites they decreased; at some sites, angles changed clockwise, at others counterclockwise. It is not clear what these correlations indicate.
It is very unlikely that they indicate variations in geomagnetic parameters indicating position, as the pigeons always released at the same site did not need to
determine their position on a navigational basis again and again. They just
needed to fly an entrained compass course which they most likely determined as
an angle to the sun (Fller et al. 1983; Sect. 3.2.3). Southern (1978) found analogous correlations with magnetic activity in compass orientation of gull chicks
inside a small arena and apart from any context with homing (for discussion see
also Wallraff 1983). Investigations in this field have more extensively been reviewed by Wiltschko and Wiltschko (1995), but also they do not come to a conclusion on the essence of the sometimes observed correlations between temporal variations in pigeon orientation and geomagnetic parameters.
6.3
Experiments aimed at clarifying the possible role of geomagnetic signals in
the home-finding process produced results that, at first glance, appeared confusing rather than elucidating. On the one hand, they showed that homing itself remained unaffected by various magnetic treatments and conditions. On
the other hand, initial orientation appeared often drastically disturbed
(Sects. 6.1.2 and 6.2). Thus, pigeons proved in some way sensitive to magnetic
stimuli, but apparently did not need them to find the way home. In the following sections I attempt to make the case understandable. An additional section
then deals briefly with the still not clearly recognized sensory mechanism
making magnetoreception possible.
79
6.3.1
The Interference with the Stress Issue
Disturbed initial orientation caused by artificial magnetic conditions or correlated with natural ones, even if not occurring under all circumstances, seemed
to indicate that magnetism is in some way involved in the navigational process.
As the magnetic field of the earth was the most favoured candidate basis of
home-finding, it is understandable that, over many years, nobody considered
the possibility that modified magnetic parameters might interfere with pigeon
orientation as an unspecific cause of disturbance. Finally, it was the merit of
Papi and colleagues to present evidence that some of the observed phenomena
can be explained on a motivational rather than a navigational basis (Papi and
Luschi 1990; Papi et al. 1992; Del Seppia et al. 1995, 1996; Papi 1995; Luschi et al.
1996, 1999). Artificial magnetic fields can interfere with the animals opioid
system and thereby inhibit the compensation of stress (e.g. Kavaliers and
Ossenkopp 1986, 1991) which is, to a smaller or larger degree, always induced
n Fig. 6.5. Stress factors, opioid system and experimental treatments: their influence on initial
homeward orientation according to Luschi et al. (1999). A Stress levels occurring in each homing
experiment are usually compensated by the birds opioid system. B This compensation can be
blocked by artificial magnetic oscillations or by an opiate antagonist drug such as naloxone. C
Particularly strong stress factors (e.g. transport in darkness) cannot be fully compensated. In
cases B and C, the pigeons are distracted from focusing their attention to directing their flight
homewards immediately after release
80
by the handling procedures during an experiment. Pigeons excited above a

certain threshold tend to fly away immediately after release in arbitrary directions or according to their PCD (Sect. 6.1.2, Fig. 6.4). After calming down a few
minutes later, they direct their course homeward, as non-stressed birds do immediately, and return at normal speeds. Thus, various pre-release treatments,
among them not only oscillating magnetic fields but also apparent stress inducers such as darkness or immobilization, affect the birds behavioural motivation, but do not impede their ability to perceive navigational signals. The different ways in which magnetic and other treatments are thought to induce
transitionally disturbed orientation are illustrated in Fig. 6.5. Threshold levels
for stress-induced disturbance of initial orientation are probably variable depending on strain, age and preceding handling experience of the pigeons and
possibly also depending on the difficulty of orienting homeward in a given situation (cf. Wallraff 2000b).
Although these kinds of magnetic influences have nothing to do with the
home-finding mechanism, it was necessary to deal with them. Any magnetic
effect that has been or will be observed should now be tested for possibly being
an artefact with respect to navigation. Even the correlations with spatial and
temporal geomagnetic irregularities (Sect. 6.2) should not be viewed separately from this possibility (cf. Ossenkopp et al. 1983; Ghione et al. 1998). For
the pigeons, they are all connected with an exposure to temporal magnetic
variations before or during their initial flight, as also flying over a magnetic
anomaly usually results in temporal magnetic noise. Thus, it cannot be excluded that also these observations, often restricted to small variations in initial orientation, fit into the complex of stress-induced phenomena.
6.3.2
The Magnetic Compass Issue
Despite the difficulties of interpreting magnetic effects in an appropriate way,
there is hardly any doubt that pigeons can use a magnetic compass as an alternative to the sun compass, if the latter is unable to operate when the sun is obscured by dense cloud cover (Figs. 6.1 and 6.3). In fact, it would be surprising if
homing pigeons did not use a magnetic compass, which is clearly applied by
many migratory birds (e.g. Able 1994; Wiltschko and Wiltschko 1995, 1999a).
Hints on its conceivable mechanism are given below (Sect. 6.3.5).
As long as the sun is visible, the magnetic compass obviously plays, at least in
fully mature pigeons, a remarkably subordinate role. If the sun compass is rotated by clock shift, the pigeons often appear unbothered by its non-fit with the
magnetic compass (Figs. 5.2 and 5.6). Probably it requires additional circumstances to make directional conflict apparent (Fig. 6.2; Sect. 8.1.4).
81
6.3.3
The Magnetic Map Issue
There is no indication that pigeons might have not only a magnetic compass,
but also something like a magnetic map, i.e. that they deduce not only directional but also positional information from the geomagnetic field. As reported
above, all attempts to disrupt their navigation mechanism by disturbing magnetic inputs failed to prevent the birds from finding their way back to their
home loft. Usually, even their homing speeds were not lowered by magnetic
treatment of whatever kind.
Nevertheless, much has been speculated about magnetic maps, and such
speculations are still going on (Gould 1998, 2004; Walker 1998, 1999). Walker et
al. (2002) extensively describe theoretically constructed navigation mechanisms based on geomagnetism. Although they refer also to pigeons, they ignore experimental research on pigeon homing carried out over the past
25 years almost completely. Finally, they would like to stress that up to now
there is no experimental evidence directly supporting any of the models of position determination discussed in their article. Such speculation, even if intellectually brilliant and updated from time to time over more than 100 years
(Chap. 11), has little relevance to real pigeon navigation and little heuristic
value if it is incompatible with empirical findings. Other authors who tend(ed)
to favour the idea of a magnetic map either came to the conclusion that it is
hard to believe that pigeons use a magnetic map (Walcott 1991) or, at least,
conceded that magnetic cues are not essential for homing (Wiltschko and
Wiltschko 1995), or could not detect any disturbing effect of a magnetic anomaly within which magnetic position determination should have been disturbed (Wiltschko and Wiltschko 2003b). It should be added that available evidence even argues against the assumption that the geomagnetic field might
serve as a (usually redundant) backup source of positional information. If
other such sources (olfactory cues) are eliminated, pigeons are obviously unable to replace them by magnetic signals (Sects. 7.1 and 7.2).
The case of magnetic navigation appears similar to that of sun navigation.
Either of these navigation systems is theoretically possible in principle, most
clearly as regards positions along the northsouth axis, along which magnetic
inclination and total intensity gradually decrease from the poles to the equator,
whereas the sun culminates at increasing altitudes. However, physically and
physiologically problematic details (e.g. gradients involving merely minute
quantitative differences within the distance range of pigeon homing, spatial
and temporal magnetic noise exceeding the range of necessary precision), together with the generally problematic second coordinate, may not have allowed the evolution of suitable navigation mechanisms based on these global
environmental frameworks (for difficulties with geomagnetism, see Lednor
1982; Walcott 1991; kesson and Alerstam 1998; Wallraff 1999; Reilly 2002).
While it is apparent that pigeons, within the usually tested distance range of
some 50, 100 or 300 km, do not use magnetic signals to determine positions,
82
there are some indications that birds migrating over longer distances are able
to respond to magnetic variables changing with latitude (Beck and Wiltschko
1988; Fransson et al. 2001; Fischer et al. 2003). Determination of only a line,
however, i.e. unicoordinate navigation, would not be sufficient to home to a
given position and thus would not imply the properties characterizing a map.
Also, such latitude measurements are possibly coarse; it is unknown whether
they allow the birds to recognize northsouth differences of only 50 or 100 km.
These negative conclusions concerning a magnetic map are valid for pigeons
and probably for (all?) other birds as well. Some other animals, namely hatchling
sea turtles as well as newts and lobsters, have been shown to respond to magnetic inclination and/or intensity and in some experiments they appeared to determine positions either completely or partly on a magnetic basis (Fischer et al.
2001; Lohmann et al. 2001; Boles and Lohmann 2003). As distances of displacement were fairly short (1245 km), it seems likely that the animals responded to
regional geomagnetic profiles which often include gradients whose steepness
exceeds the very smooth global northsouth slope by some orders of magnitude.
Adult sea turtles, however, were homeward oriented after magnetically simulated displacements over distances of 340 km north and south, respectively
(Lohmann et al. 2004).
6.3.4
The Path Integration Issue
Very theoretically, it might be considered possible that pigeons perform path integration while being passively transported to the release site (Sect. 4.2.1). Integrating their linear and directional motions (as perceived, for instance, by measurement of inertial forces and by applying a magnetic compass), they might be
able to reconstruct the direction towards home from any position they have
reached during the outward journey (route reversalaccording to Wiltschko and
Wiltschko, e.g. 1998). Experiments exposing pigeons to irregularly varying artificial magnetic fields and high-speed rotations during transport rendered this
idea non-realistic at least for homing after passive displacement (Wallraff 1980a;
Wallraff et al. 1980; see also Walcott and Schmidt-Koenig 1973; Sect. 6.1.2). The
hypothesis that a path integration mechanism might be ephemerally used during only a few early weeks in the pigeons lifetime (Wiltschko and Wiltschko, e.g.
1985, 1998, 2000, 2003a) is neither plausible nor sufficiently supported by experimental evidence (Wallraff and Sinsch 1988; Wallraff 2000b, 2001).
6.3.5
The Magnetic Sense Issue
Although it is apparent that many birds and other animals determine compass
directions by relying on the earths magnetic field, it is not yet apparent how
they perceive magnetic signals (e.g. Able 1994; Wiltschko and Wiltschko 1995;
Lohmann and Johnsen 2000). In recent years, experimental and theoretical re-
83
search has predominantly been focused on two potential principles of

magnetoreception: (1) a sensory system based on single-domain ferromagnetic crystals of magnetite (Fe3O4) as found in various tissues of various animals (for review see Kirschvink et al. 2001); and (2) a sensory system based on
light-induced radical pair processes in macromolecular photopigments orderly
arrayed in the retina (for review see Ritz et al. 2002; Wiltschko and Wiltschko
2002).
Most concrete evidence points to a mechanism based on superparamagnetic
nanocrystals of magnetite that have been found in trigeminal nerve terminals in
the upper beak of homing pigeons (Fleissner et al. 2003; see also Williams and
Wild 2001). The crystals are arranged in clusters embedded in a fibrous cup adhered to the cell membrane by fine fiber strands. In addition, non-crystalline
platelets of iron phosphate have been found along a fibrous core of the terminal.
The anatomical structures suggest that these nerve endings could be able to detect small intensity changes in the magnetic field. Now we can hope that it will be
possible to establish relationships between these anatomical structures and
physiological and/or behavioural functions.
The other proposed mechanism, suggesting a link between magnetoreception and photoreception and profiting from the spherical structure of the
eyes, originates from theoretical biophysical considerations (Leask 1977; Ritz
et al. 2000) and empirical behavioural observations. The latter concern experiments with caged migratory birds (and similarly also with amphibians) showing differently oriented preferred directions or disorientation, respectively,
when tested under monochromatic lights of different wavelengths and intensities (e.g. Muheim et al. 2002; Wiltschko and Wiltschko 2002; Wiltschko et al.
2002, 2003a, 2004a,b), when tested with the left or right eye occluded (Wiltschko et al. 2002, 2003b), or when tested in differently aligned weak magnetic oscillations (Ritz et al. 2004). The manifold results obtained under manifold conditions are interpreted as pieces of a puzzle whose eventual correct placement
might lead to an understanding of a light-dependent radical-pair mechanism
of magnetoreception. However, the experiments do not yet confirm definitely
that the various conditions interfered peripherally with the primary process of
magnetoreception and not more centrally with the observed behavioural activities making use of magnetoreception.
Different animals apparently apply different methods to determine compass
directions (Wiltschko and Wiltschko 1995). While some mammals and fishes
apply a polar compass with north determined by its polarity (as our compass
needle works), all thus far investigated birds, reptiles and amphibians apply an
inclination compass which deduces the direction to the nearest magnetic pole
from the inclined magnetic field vector pointing downwards irrespective of its
polarity (Fig. 6.6). It is very likely, and supported by the findings shown in
Fig. 6.3, that pigeons also have an inclination compass. These differences among
different taxa indicate that receptors sensing the magnetic field are not universally homologous in all animals.
84
n Fig. 6.6. Schema of orientation by means of a magnetic inclination compass at median north-
ern latitudes in a normal magnetic field (a) and in manipulated fields (bd). Arrows above and
below show directions chosen by birds intending to fly southwards or northwards, respectively;
correct direction is indicated by filled arrowhead. Compass needles (north-seeking pointer dotted) are thought to be free to rotate around a horizontal axis in a vertical northsouth plane. Arrows indicate the horizontal component of the magnetic field vector. Birds fly correctly in the intended direction if the magnetic field is either natural (a) or completely reversed (c); they fly in
the opposite direction if only either the vertical (b) or the horizontal component (d) is reversed.
(According to Wiltschko and Wiltschko 1972; graph from Wallraff 1984)
As regards birds, there is a tendency to assume that the two principles of

magnetoreception mentioned above are not theoretical alternatives, among
which a decision could not yet be made, but that they both co-exist in two receptor systems operating in parallel and having different functions (e.g.
Beason and Semm 1991; Wiltschko and Wiltschko 1995, 2002; Ritz et al. 2004).
Light-mediated perception is thought to constitute the inclination compass,
whereas magnetite-mediated perception is thought to be used as a map basis.
Apart from the fact that birds, at least pigeons, hardly have a magnetic map
(Sect. 6.3.3), there is no a priori reason to propose two independent mechanisms. The spherical geometry of the eye appears suitable to measure both
horizontal and vertical angles. As the compass is known to use magnetic inclination, it could easily be supposed that not only the sign but also the magnitude of the dip angle can be determined. On the other hand, it is unlikely that a
receptor being able to measure total intensity or inclination can operate without recording different values depending on the alignment of the body or head
in the horizontal plane and hence without recording compass directions as
well. Hatchlings of sea turtles are known to use a light-independent inclination
compass (Light et al. 1993; Lohmann and Lohmann 1993). Further, they appear
to use magnetic inclination and/or intensity as releasers varying an intended
compass course (Lohmann et al. 2001). At least in these animals there is no indication that two independent magnetoreceptors might be at work fulfilling
different functions. The above-mentioned putative magnetoreceptor detected
by Fleissner et al. (2003), if it actually measures small changes of intensity, might
be adapted to record such changes following body turns in the horizontal plane.
It might then act as a probe deriving directional information from varying in-
85
tensities (cf. Wallraff 1978b, 1991a). In this case, the rotational pattern of recorded intensities, or changes in intensity, could additionally roughly indicate
latitude (a chance possibly taken by long-distance migrants: Sect. 6.3.3).
The problem of magnetic sensing is still unsolved, but it may be on a good
way of becoming solved in the forthcoming years.
6.3.6
Overall Conclusions and Outlook
Empirical evidence supports the conclusion that pigeons, like other birds, can
use the geomagnetic field to determine compass directions. Hierarchically,
however, the magnetic compass ranks lower than the sun compass, which is
obviously preferred as long as the sun is visible.
Empirical evidence invalidates both the idea of a magnetic map and the idea
of path integration using magnetic input. Thus, it would hardly be promising
to direct continued research on the question of whether or not passively displaced pigeons can utilize the geomagnetic field to determine their current position with respect to the home site.
Nevertheless, some further efforts could be directed towards less crucial
questions. It would be desirable, for instance, to achieve a better clarification of
the role of the magnetic compass under particular circumstances. Why is it apparently sometimes used under overcast and sometimes not (e.g. Fig. 5.7)? Under which circumstances does it or does it not compete with a deflected sun
compass (Sects. 5.1, 6.1.1 and 8.1.3)? Actually very important, of course, are
further efforts to understand the sensory physiology of magnetoreception, but
this is a problem outside of the scope of this book.
Also, the stress complex, as associated with magnetic fields and other factors,
has not yet been exhaustively investigated (e.g. sensitivity thresholds depending on general condition and particular treatment, on age, strain, previous experience etc., as correlated with molecular processes in the opioid system). Research in this field may help us to understand certain behavioural responses in
orientation experiments, but it seems unlikely that it will contribute to the elucidation of the birds navigational mechanism.
The Role of the Chemical Atmosphere
Until 1971, olfaction was not among the candidate senses that had been considered potentially useful for pigeons to find out where they are in relation to
home. Although I was closest to this threshold when showing by various
shieldings around aviaries (Sect. 7.5.1) that some atmospheric factors appear
to be important for homing (Wallraff 1966a, 1970b), I did not draw the conclusion that these factors might be chemical substances. It was evident that a pigeon loft cannot act as an odour source like a female moth or a certain branch
of a river system towards which salmon migrate upstream for spawning. Pigeons return not only against the wind, which might carry some specific home
odour, but also at least equally well with tailwinds or lateral winds. A largescale olfactory map system appeared inconceivable due to the instability of
the soft atmosphere with its frequently varying wind directions. However,
since the search for conceivable navigational cues had reached a deadlock, it
was reasonable to test every sensory input for its possible involvement in homing without feeling inhibited by theoretical prejudice about its suitability. The
initial step in this direction by Papi et al. (1971) led finally to success and to
the consequence that the longest and most substantial chapter of this book
deals with olfactory navigation (for earlier reviews, see Papi 1976, 1982, 1986,
1989, 1991, 1995, 2001; Wallraff 1983, 1988a, 1990a,b, 1996, 2001, 2003, 2004;
Benvenuti et al. 1992b, 1998; Able 1996; Roper 1999).
7.1
Effects of Olfactory Deprivation
The most straightforward way to test the importance of a sensory modality in a
given behavioural context is the functional elimination of the related sense organ or its neural connection with the brain. For this purpose, it is necessary to
invade the animal. Depending on whether or not the applied method affects the
animals behaviour, two different sources of possibly erroneous conclusions
must be considered: (1) if the critical behavioural output is not, or not completely, eliminated or altered in the expected way, it must be carefully checked
whether the sensory input was really entirely blocked during the whole decisive
period of time; and (2) if the critical behavioural output is affected, it is often difficult to ascertain whether the effect is specific only for the currently interesting
function and not the consequence of an interference on a lower, more general
level, which affects this function among others. (If a bird does not home while
88
7 The Role of the Chemical Atmosphere
blindfolded, for instance, we should not automatically conclude that its homefinding under normal conditions is based on visual inputs. Blindness blocks almost any normal behaviour and hence homing as well.)
7.1.1
Olfactory Nerve Section
The most reliable method of making pigeons definitely anosmic is bilateral
sectioning of the olfactory nerve. This simple surgery lasts about 10 min, and if
done properly, no nerve reconstitution occurs, so that the achieved anosmia is
permanent (Wallraff 1988b). There are no directly observable side effects in
everyday behaviour such as walking, flying, perching, courtship, breeding,
feeding etc. (cf. Papi et al. 1972). Nevertheless, it is an invasive method and in
laboratory experiments some influences on non-olfactory behaviour have
been found (Wenzel and Salzman 1968; Wenzel et al. 1969; Wenzel and Rausch
1977). In their second publication on the role of olfaction in homing, Papi et al.
(1972) took this problem of possible side effects into account by combining
unilateral nerve cutting with unilateral plugging of a nostril. In one group of
pigeons the two treatments were applied contralaterally, thereby producing
impairment of olfaction. In a control group the double treatment was given
ipsilaterally, so that olfaction on one side remained unimpaired. Thus, interfer-
n Fig. 7.1. Results of releases at five sites 7899 km distant from a home loft near Florence, Italy.
All birds with one bisected olfactory nerve and one occluded nostril. Filled symbols Both treatments on the same side (controls); open symbols treatments on different sides (experimental
birds). A Vanishing bearings at the five sites. B The same data pooled with homeward direction
pointing upwards. C Cumulative percentage curves of homing performance. The pigeons were
moderately experienced in homing over distances up to 30 km. (Modified from Papi et al. 1980a)
7.1 Effects of Olfactory Deprivation
89
n Fig. 7.2. Initial orientation and homing success of inexperienced and experienced untreated
control pigeons (filled symbols) and bilaterally nerve-sectioned anosmic birds (open symbols). A
Each peripheral triangle represents the mean vanishing bearing of a sample of 1553 first-flight
pigeons released at one of 24 sites symmetrically distributed at distances 30300 km from their
home loft near Wrzburg. The same data are arranged with respect to home and to north; arrows
give resulting second-order mean vectors. B Corresponding mean bearings from 717 very experienced pigeons released at four and four symmetrical sites 30 and 150 km distant from home.
Numbers in A and B give mean vectors and statistical significance (cf. Fig. 3.9). C, D Homing success of first-flight and experienced pigeons simultaneously released at the latter two quartets of
sites (numbers = n pigeons released). E Return rate of anosmic pigeons (ANO) as a percentage of
the return rate of simultaneously released control birds (OLF). Large symbols and solid lines Same
data as in C and D; small symbols and dashed line all results with inexperienced pigeons within the
range of 30150 km from home. (Data from Wallraff 1980b, 1981a; Wallraff et al. 1986a, 1989)
ence with the nervous system as well as interference with breathing and mechanical irritation occurred in both groups alike. The differences in treatment
and not the similarities determined the outcome of the experiment: Only the
control birds were homeward-oriented at departure (see Fig. 7.1, which shows
such experiments of a later series). The stronger olfactory deprivation resulted
also in significantly reduced homing performance. The observation that a
noteworthy portion of contralaterally treated birds reached home at all can be
plausibly explained by the fact that nose plugging does not eliminate but
merely reduces olfactory sensitivity (Sect. 7.1.4).
Initial bearings of bilaterally nerve-sectioned first-flight pigeons released at
many sites in Germany at distances ranging from 30 to 300 km are summa-
90
rized in Fig. 7.2A. Unlike the control birds which show, as usual (Fig. 3.4), a
preference for both the homeward direction as well as a preferred compass direction (PCD), the anosmic pigeons appear disoriented with respect to home,
whereas their PCD is overexpressed. Even more dramatic is the difference in
old experienced pigeons which had previously homed many times from various directions and distances, but not from the sites used now (Fig. 7.2B). Untreated control birds were well homeward oriented without exhibiting a PCD,
whereas the anosmic birds did not indicate any homeward orientation, but revealed that the veteran birds had not definitely lost their original PCD. Also, the
surgical intrusion had obviously not eliminated ability and motivation for all
kinds of controlled spatial orientation.
Bilaterally nerve-sectioned pigeons completely failed to return when released far distant from home (Fig. 7.2CE). Only over short distances did several anosmic birds home. From 30 km, the rate was much smaller in first-flight
pigeons compared with extensively experienced birds which certainly were
largely familiar on a visual basis with the whole area in this range (Sect. 8.1).
What did the non-homing anosmic pigeons do? Some of them covered considerably long distances without, however, showing any relation to the home
site (Fig. 7.3B). Most of the inexperienced control birds, in contrast, even if not
reaching home, shortened their distance from home (Fig. 7.3A; for more details about such recoveries, see Wallraff 1989a). Comparable data from untreated experienced pigeons are not available, because almost all of them were
successful in homing (see Fig. 7.2D). Even more contrasting, therefore, is the
behaviour of the anosmic experienced pigeons (Fig. 7.3C). Without approaching the loft, many of them covered remarkably long distances; half of those released 150 km from home were found more than that distance away from the
release site. Possibly these veteran birds were previously selected for, and
trained by, persistence in searching for home.
Even those of the untreated first-fliers that did not reach their home loft were
found clearly homeward-oriented (Fig. 7.3A,D,G). The farther away from home
they were released, the longer were the median distances they subsequently flew
(Fig. 7.3F).This does not necessarily mean that they were aware of their distance
from home, but if released far away, they still felt that home was ahead after a
long flight toward it and thus they continued flying in that direction. The anosmic pigeons, in contrast, were obviously disoriented everywhere (Fig. 7.3B,
D,G) and hence had no indicator stimulating them to continue flying in a certain direction. Thus, their distances of recovery were independent of distances
of displacement (Fig. 7.3F). As shown in Fig. 7.3E, they maintained their
(north-)westward PCD not only at departure (cf. Fig. 7.2A), but also while flying longer distances.
In conclusion, bilateral olfactory nerve section definitely prevents any homeward orientation in unfamiliar areas without generally preventing spatial orientation and without abolishing the pigeons motivation to fly considerably long
distances. Reduced homing success remains only in a limited range around
home, its size depending on the degree of preceding homing experience, which
91
n Fig. 7.3. A, B Recoveries of inexperienced pigeons released at 12 sites about 150, 180 and
300 km distant from their home loft near Wrzburg (centre H); A untreated control birds; B
nerve-sectioned anosmic birds; minimum distance from release site 20 km. Numbers in boxes
(referring to the nearest release sites) indicate number of pigeons released/returned. C Recoveries of very experienced anosmic pigeons released 30 and 150 km from home at Wrzburg. D, E
Mean directions of recoveries (>20 km from release) of inexperienced pigeons after release at 24
sites 30300 km from home (same releases as in Fig. 7.2A). Each peripheral triangle represents a
mean from 223 birds, each arrow a resulting second-order mean vector. Filled symbols Controls
(OLF); open symbols anosmic birds (ANO). The same data are differently arranged in D and E.
F Median distances of recoveries from the release site as a function of the distance of displacement. Connected points are from first-flight pigeons released at the 24 sites plus one site 500 km
north; the square exp. refers to experienced anosmic birds released 150 km distant (median of
simultaneously released first-flight birds almost identical with median of the total). G Running
medians of absolute angular deviations of recoveries of inexperienced pigeons from the direction towards home as a function of the distance from the release site (cf. Fig. 3.6). Note that the
likelihood of getting pigeons reported was higher from inside the borders of western Germany
than from outside; this may have caused the ANO average to lie below the random value 90.
(Data from Wallraff 1980b, 1981a, 1989a; Wallraff et al. 1986a, 1989)
92
could have made landscape features familiar (Sect. 8.1). Experiments combining
unilateral nerve section with nose plugging in both experimental and control
birds (Fig. 7.1) make causation of the results by non-olfactory side effects unlikely.
7.1.2
Inactivation of the Olfactory Epithelium by Zinc Sulphate
Intranasal irrigation with a solution of zinc sulphate (ZnSO4) causes selective
degeneration of the olfactory epithelium (e.g. Matulionis 1975, 1976; Cancalon
1982). As in other animals, the sense of smell of pigeons is strongly impaired or
eliminated over a period of several days (Benvenuti et al. 1992a; Schlund 1992).
Effects of treatment with ZnSO4 on homing were analogous to the effects of olfactory nerve section. Figure 7.4 shows results obtained with this technique in
seven regions of the earth (see also Benvenuti et al. 1998). Performances of untreated control birds varied considerably among the different areas, but anosmic
pigeons failed to orient homeward at departure everywhere and their return rate
n Fig. 7.4. Experiments using zinc sulphate for olfactory deprivation. Mean homeward compo-
nent (A) and return rate (B) per release of samples of pigeons from different, mostly symmetrical
directions and distances (the latter given below). Seven home sites in four continents. Means per
home site indicated by horizontal lines, overall means by arrows on the right. Filled circles and
solid lines Control birds; open circles and dashed lines pigeons whose olfactory epithelia were irrigated with a solution of ZnSO4. The pigeons had not yet been released at the respective sites;
their general homing experience (and hence their potential familiarity with the area) was heterogeneous (sources: Pisa: Benvenuti et al. 1992a; Benvenuti and Gagliardo 1996; Casablanca:
Gagliardo et al. 2000; Arizona: Bingman et al. 1998a; Georgia: Bingman and Benvenuti 1996; So
Paulo: Ranvaud et al. 2001; Tbingen: Schlund 1992; Oxford: Guilford et al. 1998)
93
was everywhere clearly lowered. Homing speeds were even more drastically affected than homing rates in Fig. 7.4, which include very slow birds returning several days after release.At least over the shorter distances,some pigeons may have
been more or less familiar with the area from previous homing flights (though
they had not yet been released at the exact current site). Also, anosmia was probably not always perfect and may have ceased after a few days, so that some of the
treated birds were able to return with some retardation (information about
homing speeds has been communicated in the literature in different ways, making standardized graphic presentation impossible).
Figure 7.4 shows the clearest effect of olfactory deprivation in Italy, where it
was originally detected (Papi et al. 1971, 1972). Less clear effects reported from
other countries (e.g. Wiltschko et al. 1987d) led frequently to the opinion that
atmospheric odours may play a major role in Italy, whereas homing in many
other regions relies primarily on other cues. However, the geographical differences do not concern the degree of involvement of olfactory navigation, but its
efficiency. Without olfactory inputs, homeward orientation in reliably unfamiliar areas appears to be impossible everywhere on the earth. Yet these inputs
appear to contain variably reliable positional information (Sect. 7.4.1).
Benvenuti and Gagliardo (1996) combined unilateral nose plugging with
unilateral ZnSO4 irrigation in the same way as described above for the combination with nerve section. Differences between ipsilaterally and contralaterally treated birds in initial orientation as well as in homing performance
were very similar to those shown in Fig. 7.1. Thus, also the effect of ZnSO4 on
pigeon homing can hardly be explained solely by non-olfactory side effects.
7.1.3
Nasal Anaesthesia
Local anaesthesia of the olfactory epithelium (first applied by Schmidt-Koenig
and Phillips 1978) is a frequently used method to exclude or impair the sense
of smell for a restricted period of time. Usually a few minutes before release of
a pigeon, a commercially dosed puff of a spray containing Xylocain (sometimes Gingicain) is applied into each nostril. The effect on olfactory sensitivity
is, on average, considerably greater than that of occlusion of the nostrils
(Sect. 7.1.4), but complete anosmia is not always ensured. Application cannot
be fully standardized, so that in practice some range of uncontrolled variability remains (Wallraff 1988b). It is unknown whether in experiments conducted
by different persons the average level of sensory deprivation might have been
somewhat different. The duration of reduced sensitivity varies over a range of
about 12 h (Wallraff 1988b). Non-olfactory side effects have been described
(Dornfeldt and Bilo 1990; Schlund 1990), but appropriate control experiments
ensure that they alone cannot be responsible for the deficits found in initial
homeward orientation and homing (Sects. 7.2 and 7.3.1).
Due to the limited duration of influence, nasal anaesthesia consistently protracts but does not prevent homing (e.g. Ioal 1983; Wallraff et al. 1984; Wiltsch-
94
n Fig. 7.5. A: Homing times (min/km beeline distance) of pigeons released under nasal anaes-
thesia (open circles) and of unimpaired control pigeons (filled circles). Symbols indicate medians
of single releases from different symmetrically distributed sites. B Differences between the medians of experimental and control birds per release, also including second-order means and 95%
confidence intervals of the two distance classes. Dotted line at 15 km indicates the upper limit of
preceding training flights. (Data from Wiltschko et al. 1987d)
ko et al.1986,1987ac).Figure 7.5 gives an example (obtained in a third region of

North America, not shown in Fig. 7.4), which additionally indicates that delayed
homing is not merely an effect of general disturbance. If this were the case, decreasing effects with increasing distances of release would be expected, because
the time of drug action would constitute a decreasing portion of the homing
time. However, just the contrary happened; pigeons departing under nasal anaesthesia homed significantly more slowly only over the longer distances. This
result is best compatible with the assumption that olfactory signals were necessary only in the more distant areas, while previous releases have been familiarized with the closer surroundings of the loft also on a visual basis (Sect. 8.1).
Effects of nasal anaesthesia on initial orientation cannot be tested without
considering the birds exposure to environmental odours prior to application
of the drug immediately before release. It will be shown below that these prerelease conditions are, or can be, more influential than the effect of Xylocain
during the first few minutes of flight (Fig. 7.9). Various experiments using nasal anaesthesia will be discussed later in various contexts.
7.1.4
Occlusion of Nostrils
The simplest way to impede olfaction is the occlusion of the pigeons nares either by inserted plugs or by adhesive tape firmly glued around the upper mandible, thereby pressing the ceres on the openings. It is obvious that this method
has a strong side effect, as it blocks the duct through which the birds normally
breathe. Therefore, occlusion of nostrils alone has rarely been used in homing
95
n Fig. 7.6. Cardiac acceleration depending on olfactory stimulus intensity (vapour saturation, in
log scale). Each symbol represents the mean of 40 tests (2 with each of 20 different pigeons). Circles Nostrils open; squares nostrils occluded. Filled symbols, solid regression line Test substance
amyl acetate; open symbols, dashed line test substance -pinene. Thin dashed lines mark stimulus intensities of corresponding response levels. (Modified from Wallraff 1988b)
experiments (Snyder and Cheney 1975). However, it has been used unilaterally
in combination with other methods (Sects. 7.1.1 and 7.1.2). Moreover, the
method has sometimes been used during transportation to a release site and
during some time of waiting there before release (Sect. 7.8.3).
Bilateral occlusion of the nares clearly reduces olfactory sensitivity, but it
does not make the pigeons anosmic. It has not been tested but appears most
likely that odorous molecules still reach the olfactory epithelia through the
choanae. In laboratory tests, a four-fold higher concentration of odorant substances was necessary to reach the same level of cardiac acceleration with
sealed nostrils as was achieved with open nostrils (Fig. 7.6). Reduction of sensitivity to approximately 25% is not very much in view of a range of olfactory
sensitivity covering several decimal powers. Although the concentrations of
substances in the free atmosphere are only a small fraction of those used in the
experiments, there is no reason to assume that relative reduction of sensitivity
at such lower levels is different from that shown in Fig. 7.6. Since the curves obtained with occluded nostrils retained the original slopes, relative quantification was apparently not affected. Thus, the perceived ratio spectrum of a number of compounds can be expected to remain largely unchanged while the
levels of all stimuli are reduced in parallel.
If homeward orientation depends on olfactory inputs, some reduction of olfactory sensitivity may or may not impair orientation. As concentrations of atmospheric trace gases are regionally and temporally variable (Sect. 7.6.2), decisive compounds may sometimes still reach above-threshold levels and
sometimes not. Inconsistent results are then likely to occur (Sect. 7.8.3).
96
7.1.5
Insertion of Nasal Tubes
Plastic tubes inserted through the nostrils prevent or impair smelling without
preventing breathing through the nares. The method is a bit complicated, may
cause some general discomfort and possibly it does not always reliably produce complete anosmia. On the whole, the results obtained with this technique
are consistent with those obtained in experiments using other methods of olfactory deprivation (Hartwick et al. 1977; Keeton et al. 1977; Papi et al. 1978a;
for discussion see also Wallraff 1980b). Again, clear deterioration in initial orientation and homing performance was observed only after release at unfamiliar sites, while there was little influence in experiments using familiar release
sites.
7.2
Dependence on the Air the Pigeons Breathe
7.2.1
Removal of Trace Gases by Filtration
The flow of olfactory information can be interrupted not only by inactivation
of the receptor or the neural pathway, but also by removal of potential signals
from the air the pigeons breathe. Such a removal is hardly possible while the
pigeons are in flight, but it is possible during displacement and at the release
site prior to release. For this purpose, pigeons were kept in an airtight container ventilated either with artificial air from a bottle or, more often, with natural air sucked through a filter consisting of a large volume of activated charcoal which adsorbs the somewhat larger volatile trace substances dispersed in
the ambient air. Control pigeons were transported, on top of a car, in a similar
container without a filter but with small inlet holes producing air resistance,
and hence reduced pressure in the bird chamber, similar to that caused by the
filter (Fig. 7.7). In order to prevent unimpeded smelling during the take-off, the
olfactory epithelia of each pigeon were anaesthetized immediately after the
bird was taken out of the container. In the critical tests, Xylocain was sprayed
into the nostrils of both the filter birds and the non-filter control birds (for details of the method, see Wallraff and Fo 1981).
Figure 7.8 summarizes the initial orientation data of the 46 experiments conducted under these conditions with pigeons from four lofts, two in Germany
and two in Italy. Corresponding releases with equally experienced birds were
conducted pairwise from similar distances (20170 km) in opposite directions
so that, in summary, home-independent directional tendencies could not have
simulated homeward orientation. A clear majority of the pigeons that were allowed to smell natural air before release vanished from sight on largely
home-related courses, whereas the pigeons that had inhaled filtered air did not
show any preference for directions towards home. Nevertheless, the latter birds
7.2 Dependence on the Air the Pigeons Breathe
97
n Fig. 7.7. A Airtight containers placed on the roof of a van. The lattice cover usually damping
sun irradiation is removed from one container, so that pigeons and the vacuum cleaner in the
back compartment are visible. The other fully visible container is equipped with a charcoal filter
at its air inlet (for more details, see Wallraff and Fo 1981). BE Chromatograms of atmospheric
air sampled over 2 h on 2 days (B, C and D, E) in October in southern Bavaria (for methods of air
sampling and analysing see Sect. 7.6.2). B, D Normal natural air; C, E air sampled simultaneously
in a container equipped with a charcoal filter as shown in A. C The container was empty. E Eight
pigeons were sitting for 1 h before sampling in the front compartment, while airborne gases were
collected in the middle compartment (cf. A). Abscissa Retention times in gas-chromatographic
analyses; ordinate relative units (identical for each graph) indicating amounts of trace substances. Peaks were produced by volatile organic compounds, mainly hydrocarbons containing
313 C atoms as indicated for n-alkanes below of B and D (original)
98
n Fig. 7.8. A, B Mean directions of vanishing bearings per release; same data ordered with re-
spect to deviation from home (A) and north (B); each peripheral triangle represents the mean of
a sample of 414 pigeons, each central arrow the second-order mean vector calculated from 46
mean directions. Numbers in A and B give mean vectors and statistical significance (cf. Fig. 3.9).
C Mean homeward components obtained from the same 46 experiments; horizontal lines indicate second-order means per home loft, arrows on the right overall means (significantly different
with P<0.0001). Filled symbols and solid lines Control pigeons without filter; open symbols and
broken lines experimental pigeons kept in a container with filter (first-order total n=433 and 434
bearings). All birds departed under nasal anaesthesia. Numbers on the abscissa of C give the
mean distance from home of every two releases which were conducted pairwise with pigeons of
equivalent experience from approximately opposite sites at a similar distance. (Data from
Wallraff and Fo 1981; Wallraff et al. 1984, 1993 and several analogous experiments)
were also not disoriented; they preferred directions in the western semicircle,
in accordance with PCDs typical for all four home sites, clearer than the controls. By and large, the orientation patterns are very similar to those obtained
with nerve-sectioned birds (Fig. 7.2A). Figure 7.8C illustrates that the effect
was quite consistent in Italy as well as in Germany; in 40 out of 46 releases the
controls were better homeward-oriented than the filter birds.
These experiments are particularly conclusive because it is difficult to give
any plausible interpretation on a non-olfactory basis. Before release, neither
control nor experimental birds were injured in any way; both groups were unimpeded in smelling (even with a filter, the interior of the wooden containers
filled with pigeons was not clear of odorous substances; cf. Fig. 7.7C, E). Upon
release, the olfactory epithelia of both experimental and control birds were anaesthetized. Thus, any non-olfactory side effects caused by nasal anaesthesia
(Sect. 7.1.3) acted in essentially the same way in either group, so that such effects could not have been responsible for the different behaviour of the groups.
In some experiments, not only charcoal filters were used but also filters
made of fiberglass paper which retained almost all the aerosol particles floating in the atmosphere. These filters were ineffective (Wallraff and Fo 1981).
Thus, volatile substances in the gas phase must be useful for navigation.
Atmospheric chemosignals do not instantaneously provide the pigeons with
an achievable optimum of positional information. Even if not treated with an
anaesthetic, the birds coming from a filter container are, on average, much
poorer in initial homeward orientation than those that could smell natural local
air for several hours before release (Fig. 7.9, c versus a and b). About 515 min of
7.2 Dependence on the Air the Pigeons Breathe
99
n Fig. 7.9. Second-order homeward components of vanishing bearings in 16 releases using
charcoal filters combined with nasal anaesthesia. In each release, four groups of pigeons with different combinations of treatment were used. Left-hand half-column Condition before release;
right-hand half-column condition during release; light grey without treatment; dark grey with
treatment. (Data from Wallraff et al. 1984)
smelling before and during initial flight is obviously insufficient to reach full
performance; some time for collecting and processing of airborne signals appears to be necessary. Conversely, nasal anaesthesia applied shortly before release has little effect on initial orientation if the birds were allowed to smell the
natural air over several hours during transportation and/or at the release site
before they began flight (Fig. 7.9, b versus a). For initial orientation, smelling of
natural air over a longer period of time before release is thus more decisive
than smelling during the few minutes while the departing birds are under observation.
7.2.2
Air from Different Environments
The filter experiments have shown that ground-level air contains sufficient information to determine the approximate direction towards home. It is unknown whether air at somewhat higher altitudes contains better information.
At least, however, it seems to be not so much better that its availability during a
few minutes of flight substantially improves orientation (Fig. 7.9, a versus b) or
compensates preceding lack of information (Fig. 7.9c). On the other hand,
does any sample of ground-level air contain equally suitable positional information? Air staying, with little exchange, closely above the ground of a forest or
a field of full-grown maize provides apparently less suitable information than
100
n Fig. 7.10. Second-order homeward components of vanishing bearings in 16 (left) and 18

(right) experiments with pigeons exposed, before release, to air of different environments. All
birds were released under nasal anaesthesia. With the two series combined, the birds exposed to
air among vegetation were, with P<0.001, clearly poorer in homeward orientation than the birds
that could smell open-land air before release. (Data from Wallraff et al. 1992)
the free airspace in an open landscape to which pigeons are usually exposed
before release (Fig. 7.10). In the related experiments, two containers with pigeons were transported with filters to the release area. In the first series, the filters were dismounted there for a period of 3 h. During that time, one of the
boxes (containing the forest birds) was deposited in a fairly dense forest,
while the other box (containing the open land birds) was placed on top of the
van in open country some 25 km distant. Thereafter, with the filters mounted
again, both types of birds were alternately released in the usual manner. In the
second series, the filters were dismounted at the release site, which was at the
edge of a field of maize, mostly green and 23 m high. At the inlet of one container, the filter was replaced by a tube which was laid over a length of 2030 m
into the maize field at ground level, whereas the other box was ventilated with
free air from 24 m above ground.
Consistent with these results is the finding that pigeons kept in a weakly ventilated container with relatively stale air were not homeward oriented (Wallraff
et al. 1984). Also, the interior of a barn seemed not to contain sufficient positional information (Papi et al. 1984).
7.3 Olfactory Misguidance
101
7.3
Olfactory Misguidance
7.3.1
Spatial Separation of Sites of Air-Smelling and Release
Once it had been found that olfactory conditions before release determine initial orientation, it was a logical subsequent step to disconnect the site of smelling local air from the site of release. The pigeons treated in this way vanished in
directions that were, by and large, appropriate to the site of smelling but not to
the site at which they were actually released. This was shown in two series of
experiments independently conducted in different areas using somewhat different methods (Benvenuti and Wallraff 1985; Kiepenheuer 1985).
The inset in Fig. 7.11 explains the experimental procedure used in one of the
series. In containers equipped with filters as mentioned, two groups of pigeons
were transported to different sites (T and F) in roughly opposite directions
from home, where the filters were simultaneously removed. After the birds
were allowed to smell natural air for a period of 3 h, the filters were reinstalled.
One of the groups (false open) was then driven from site F to site T, where the
other group (true open) remained waiting. A third group (false filter) participated in the detour from F to T, but was never allowed to smell unfiltered, natural air. Pigeons of the three groups were then released alternating at site T, all
birds flying under nasal Xylocain anaesthesia as described above.
Figure 7.11 summarizes the initial orientation data of these birds. It is evident that the experimental birds oriented towards the simulated home direction as equally well as the control birds towards true home (Fig. 7.11, E versus
=
=
=
A; cH = 0.32 versus 0.28). The smaller negative value of cH in C (cH =-0.20) is
explainable by the fact that the two directions of a pair of releases (true home
and false home) were not always exactly opposite each other. The third group
of pigeons, which were at no time allowed to smell unfiltered air, ranged between the two other groups. Correspondingly to other experiments (Figs. 7.2
and 7.8), those birds completely deprived of smelling natural air showed a general compass tendency towards WSW most clearly, but, in principle, all three
groups had this PCD in common (Fig. 7.11FH).
In a similar series of experiments conducted around Tbingen, Germany,
Kiepenheuer (1985) carried pigeons in open baskets to two opposite sites, allowed them to sit there in the open air for 1 h and then kept them, for further
transport and waiting,in spacious (750-l) plastic containers,each with five birds.
These airbags had been ventilated with local air from the exposure site. All birds
were driven back to the loft, half of them exchanged and again carried to the two
sites. Here, their nares were anaesthetized with Gingicain while the birds were
still sitting in the container, from which they were taken singly for release. The
results obtained at four pairs of opposite sites (distances 1123 km from home)
are summarized in Fig. 7.12 (filled symbols).Including the data given in Fig. 7.11
102
n Fig. 7.11. Vanishing bearings observed in 10 releases, pairwise consecutively conducted at
roughly opposite sites at distances of 2455 km from home, with three differently treated groups
of pigeons. The experimental procedure is illustrated by the framed insert (H home; T true site of
release; F false site of air exposure; central arrows at T example mean vectors of initial bearings). Diagrams show the pooled individual bearings on the periphery, mean vectors per release
as arrows and second-order mean vectors derived from them as centre of a 95% confidence ellipse. The same bearings are shown as deviations from three reference directions: the actual true
homeward direction (left), the direction pointing towards home from the site where the False
open birds had been exposed to natural air (middle) and north (right). P values inside the diagrams result from using the V test referring to the upward direction (AE) or from using the
Rayleigh test (FH) (Batschelet 1981). P values between diagrams refer to two-sample comparisons of upward components of individual bearings using the Mann-Whitney U test. A versus C
is significant, on this condition, with P<0.0001; a second-order test applied to the 10 pairs of
mean vectors in A and C results in P<0.01. (Data from Benvenuti and Wallraff 1985)
103
n Fig. 7.12. Summary of site simulation results in two independent series of experiments with
pigeons from three lofts in two countries. Each peripheral symbol indicates the mean vanishing
direction of a sample of pigeons (n=427) released at the same site. Filled triangles Eight release
sites around Tbingen, Germany (data from Kiepenheuer 1985). Open symbols refer to the data
of the Fig. 7.11 series (also 8 sites); triangles home loft at Andechs (Munich), Germany; diamonds
home at Arnino (Pisa), Italy. Mean vectors resulting per loft and from all 16 bearings (heavy arrow) are also given; the homeward component c H refers to the total. Mean directions of the experimental birds (exposed to natural air at a false site roughly opposite from home) are shown
twice, in B and C, in relation to different reference directions. Deviations from home in the 16
pairs of corresponding second-order bearings in A and B are significantly different with P<0.001,
eight black and white pairs, respectively, with P<0.01 (Wilcoxon test)
as well, the figure summarizes all the related experiments conducted around
three lofts, two in Germany and one in Italy.
It is reasonable to assume that the pigeons, if sufficiently anaesthetized,
roughly continued their initial courses for a while and thereafter gradually redirected their flight according to the restored olfactory feedback. At this time, the
majority of the control birds should have shortened their distance from home,
whereas the majority of the false site birds should have lengthened it. The time
required by the two groups to reach home in the filter experiments (cf. Fig. 7.11)
differed significantly according to this expectation (Fig. 7.13A). Again, the permanent-filter birds ranged between the two other groups.
In a third series of related experiments, DallAntonia et al. (1999) used route
recorders (Sect. 2.3), tracking the birds complete paths home, of which two examples are shown in Fig. 7.13B. Without restrictions in smelling, pigeons were
brought to two opposite false sites, N and S, where they remained sitting for 3 h.
Then they were transported, in containers with filtered air, to a release site R. A
few minutes before release, the nares of each pigeon were anaesthetized with
Xylocain. Depending on where the birds had been exposed to natural air, they
tended to fly home on detours deviating either south or north of a direct course.
Not all routes were as clear as the two examples shown in Fig. 7.13B,but no counter-example is available and the difference between the groups (five and six
birds) is statistically significant. (A general westward tendency may be due to in-
104
n Fig. 7.13. Homing after air exposure at a false site. A Cumulative percentage curves of homing performances following initial orientation as shown in Fig. 7.11. Filled symbols refer to True
open, open symbols to False open, and line without symbols indicates False filter data (data
from Benvenuti and Wallraff 1985). B Two groups of pigeons from a loft near Pisa were carried
from their home (H) to sites N or S, respectively, where they could smell natural air for 3 h. Thereafter breathing only filtered air, they flew off, under nasal anaesthesia, at the release site R (outward journeys shown schematically). Map shows homing routes of one pigeon per group. Arrows
labelled 2 h indicate positions reached 2 h after release around which time anaesthesia may have
ceased. (Data from DallAntonia et al. 1999)
complete anosmia and/or to the south-westward PCD of the Arnino pigeons, as

it is known from many other releases; cf. Fig. 7.11FH and Ioal 1995.)
7.3.2
Outward-Journey Detours
These site simulation experiments can be seen as particularly distinct forms of
detour experiments during which pigeons are transported to the release site
not on a route as straight as possible but on a route widely bulged either to the
right or to the left (Papi et al. 1973, 1978b, 1984; Keeton 1974b; Hartwick et al.
1978; Tgel and Wiltschko 1992). In many cases and on average, the initial
bearings chosen by such birds deviate from the normal course in a direction
away from the detour bulge. Figure 7.14 shows a particularly clear example.
These detour experiments indicated at an early stage that departing pigeons
refer not only to positional information currently available at the release site,
but also to information previously collected during passive transportation.
Later, it was shown that such deviations occur only if the pigeons were not impeded in smelling natural air during the outward journey and thus that the
pre-release information used is olfactory in nature (Papi et al. 1984). Since we
have seen above that smelling of natural air during a longer period before release is more influential than smelling around take-off time (Fig. 7.9), the deviations are consistent with the site simulation effects. During transport, the pigeons collected olfactory information in areas outside the direct line between
home and release site, so that the remembered median position is accordingly
105
n Fig. 7.14. Pigeons transported along different detours from their loft near Florence to the un-
familiar release site vanished in directions deviating clockwise (CW) or counterclockwise (CCW)
from the direct course towards home. (Modified from Papi 1976)
displaced (see Fig. 14 in Wallraff 1990a). The degree to which a detour modifies
initial orientation depends most likely on the spatial and temporal relationships between the areas crossed and the release site itself, on the navigational
suitability of olfactory signals gained there and here, and on the strength of
non-affected home-independent directional tendencies (Sect. 3.7.1). To a great
part, these parameters were not known or not controlled; probably they were
quite variable in different experiments. It is not surprising therefore that the
observed angles of deflection vary between nil and more than 90.
7.3.3
Natural Olfactory Misguidance
Given a particular geographical relationship between release area and home
site, it can happen that normal non-manipulated pigeons fly well oriented away
from home, not only initially, as sometimes occurs, but definitely over longer
distances (Fig. 7.15C). As, under the same conditions, anosmic birds did not
show this strong directional preference, the findings suggest that olfactory inputs gained in the River Po valley made pigeons from a loft just beyond the Alps
believe themselves to be not south but north of home. The same inputs, however,
seemed to contain fairly correct (albeit not very precise) positional information
for pigeons coming from a loft 200 km farther away in the north (Fig. 7.15A,B).
These results suggest that, even within a given spatial range, atmospheric
signals do not induce correct homeward navigation from everywhere to everywhere. They may even mislead the birds towards completely wrong directions.
Although, at first glance, the results of these transalpine experiments appear
rather puzzling, it will later be shown that it is possible to include them in a
fairly simple hypothetical interpretation (Sect. 7.6.4).
106
n Fig. 7.15. Recoveries of pigeons from two German lofts released at a site near Mantua in the Po
valley. A, B Home loft near Wrzburg; C, D home loft at Andechs (ca. 30 km SW of Munich and
25 km ESE of L in Fig. 7.17). Olfactorily intact birds (OLF) were transported from Wrzburg to
Andechs in charcoal-filtered air, from Andechs across the Alps with occluded nostrils and in the
first part of the journey additionally under nasal anaesthesia; during hours in the release area
they were olfactorily unimpeded. Anosomic birds (ANO) had bisected olfactory nerves. Radius
of circles = 100 km. Symbols refer to different releases in 4 years. Note that from about 50 km
north of the release site onward the ascending slope of the Alps probably hampered northward-flying pigeons in A. (Wallraff 1993)
7.4 Spatial Range and Variability of Olfaction-Based Homing
107
7.4
Spatial Range and Variability of Olfaction-Based Homing
Many of the peculiarities of homing behaviour described in Chapter 3 should
now be seen as reflecting peculiarities of olfactory navigation. In particular, the
temporal and geographical variability of pigeon homing (Sects. 3.3 and 3.4.2) as
well as its spatial range (Sect. 3.5) must be considered under this aspect.
7.4.1
Distance Range and Efficiency in Different Regions
As a consequence of the last-mentioned findings (Fig. 7.15), we do not expect
an everywhere effectual maximum distance over which olfactory navigation
operates. Nevertheless, with respect to pigeons of a given home loft, we can try
to determine the spatial range within which they are able to deduce useful positional information from local ambient air. This range can be reliably determined only if the birds are prevented from receiving olfactory signals during
the outward journey, but have sufficient time to smell the air at the location at
which they are tested. Best for this purpose are experiments in which the birds
are transported while breathing artificial or filtered air, respectively. Results of
such experiments with first-flight pigeons were different in different regions
(Fig. 7.16). In Italy (Tuscany), air filtration during transport had little or no effect over distances of less than 80 km, but prevented initial homeward orientation at distances more than 100 km away from home (Benvenuti et al. 1994). In
Germany, transportation without access to environmental air did not reduce
orientation levels at sites 180 km distant; recoveries and return rates also did not
indicate any influence of outward-journey conditions (Wallraff 1980a,b). Other
first-flight pigeons were well oriented towards home even at 300 km distance after transportation with occluded nostrils in a sparsely ventilated car (Figs. 3.4
and 3.5; Wallraff et al. 1986a). Pigeons well experienced in homing were, in this
country, equally good in initial homeward orientation and returning performance when released at two other sites 300 km distant from home (and more
than 170 km distant from previous release sites), independently of whether they
were or were not prevented from smelling natural air during transport (Wallraff
1980a). In Italy, in contrast, experienced pigeons were clearly homeward-oriented only when allowed to breathe natural air while being displaced to sites as
far as 100200 km from home, but more poorly or not at all after transport isolated from ambient air (Wallraff et al. 1980; Baldaccini et al. 1982).
Thus, the range within which olfactory navigation operates appears to depend
on particular geographical conditions. Some findings indicate that, in fairly flat
regions, the range may exceed 300 km considerably, and may even exceed
700 km (Wallraff 1981a), while the conditions are less clear-cut in generally
mountainous regions (Italy) or with a high mountain ridge between home and
the release site. In the latter case, in transalpine displacements from Tuscany to
Bavaria, results differed depending on distance and treatment of the birds
108
n Fig. 7.16. Mean homeward components of vanishing bearings in releases of first-flight pi-
geons in Italy and Germany. Black bars and solid horizontal lines Birds breathed natural ambient
air during transport; grey bars and broken lines birds breathed filtered air (A, B) or artificial air
(C) during transport. Horizontal lines indicate second-order means of all five, six and four releases. Upon release, all birds were unimpeded in smelling natural air. Each bar is calculated
from 1115 individual bearings (A, B data from Benvenuti et al. 1994; C data from Wallraff 1980a)
7.4 Spatial Range and Variability of Olfaction-Based Homing
109
n Fig. 7.17. Recoveries of pigeons displaced from Florence (F) over 490 km to Landsberg (L) and
over 680 km to Wrzburg (W). Prior to the transalpine displacement the pigeons had made a
number of training flights mainly on the northsouth axis over distances up to 170 km. The
training site farthest north was in the Po valley near Mantua (M). A Control birds unimpeded in
smelling environmental air during transport as well as afterwards. B Birds transported with their
nostrils occluded; unimpeded olfaction afterwards. C Anosmic birds with bilaterally sectioned
olfactory nerves. Circles and solid lines Release sites near Wrzburg; triangles and broken lines
release sites near Landsberg (50 km WSW of Munich). Open and filled symbols refer to different
years. Symbols close to F indicate returned pigeons. The grey zone marks roughly the Alps; stippled area between M and F marks the approximate area familiarized by previous homing flights.
(Modified from Ioal et al. 1983)
(Fig. 7.17). Olfactorily intact pigeons released in an area west of Munich fairly
close to the Alps, about 500 km north of their loft near Florence, tended to fly
southward, but were obviously deflected by the mountains (graphs A and B, release site L); about 20% of them returned to the loft. Their apparent homeward
orientation was not recognizably affected by occlusion of their nostrils during
transport (B versus A). In contrast, from almost 700 km north (site W), southward tendencies were indicated only in those birds that were unimpeded in
smelling natural air during transport. Thus, the release area W apparently did
not contain positional information. Southward tendencies in Fig. 7.17A must
have been induced by remembered olfactory signals perceived during the outward journey, which included the familiar area between sites F and M. Figure 7.17C suggests that the southward tendencies from L, as shown in B, resulted from olfactory inputs available in that area.
Further transalpine displacements, from north to south, which provided
even more peculiar results, have already been shown above (Fig. 7.15; see also
Fig. 7.35 in Sect. 7.6.4). They demonstrate most clearly that it is impossible to
define an ordinary distance limit that is valid in all regions and all spatial relationships among sites.
110
In general, it seems that the upper range limit of olfactory navigation tends
to be at longer distances in Germany compared to Italy. In Italy the functional
range includes very short distances down to 7 km (Meschini 1983), whereas in
Germany no effect of olfactory deprivation was found at distances from home
less than 10 km, and inconsistent effects were found at 30 km (Wallraff 1981a,
1982). Over 2030 km, first-flight pigeons were considerably better in initial
orientation and homing performance in Italy than in Germany, even if they belonged to the same genetic stocks (Fo et al. 1982, 1984; Fig. 3.20). Consequently, olfactory deprivation tends to have a much more drastic effect over
shorter distances in Italy, because the breakdown occurs from a higher level
(e.g. Benvenuti 1979; Meschini 1983; Wiltschko et al. 1986, 1987d; Schlund
1992). The general impression is that olfactory navigation is most effective in
Italy over fairly short distances, whereas in Germany it becomes increasingly
operational further away from home and then remains functional over larger
regions than in Italy, albeit on a lower level of efficiency. Other countries have
been examined less thoroughly under this aspect, but it seems that England as
well as various regions of North America are more similar to Germany than to
Italy (Fig. 7.4A; e.g. Keeton et al. 1977; Wiltschko et al. 1987d; Benvenuti and
Brown 1989; Bingman and Mackie 1992; Bingman and Benvenuti 1996;
Bingman et al. 1998a; Guilford et al. 1998). Olfaction-based home orientation at
levels as high as over shorter distances in Italy has been rarely found elsewhere.
In other countries, as well as over longer distances in Italy, the atmosphere
seems to contain less reliable navigational signals. Nowhere, however, has clear
evidence been found that pigeons are able to home from unfamiliar areas by
using alternative, i.e. non-olfactory, environmental cues (Sect. 7.8.3).
7.4.2
Temporal Variability
It is reasonable to assume that the superiority of Italy over Germany in the
short distance range (Fig. 3.20) has something to do with differences in climate. On this assumption, it is plausible that, in Germany, a marked annual cycle in homing performance has been found that is correlated with ambient
temperature (Fig. 3.17). With an increase in temperature, vapour pressure of
volatile chemical compounds increases and hence above-threshold availability
of potential olfactory signals in the atmosphere might increase as well. If we
are ready to speculate further, we may even tentatively interpret the finding
that homing was correlated with short-term fluctuations of temperature in autumn but not in spring, although absolute levels of temperature were similar
(Fig. 3.17B, dashed horizontal bar). This finding might indicate that decisive
compounds originate from the vegetation, whose growth activities in spring
override temperature-dependent short-term variations, which holds fairly
high levels of evaporation in summer and enters a refractory phase in autumn
during which evaporation depends on physical conditions rather than on biological activities.
7.5 Varying Home Site Conditions in Aviary Experiments
111
If homing depends on atmospheric chemosignals, it is not only plausible but

also should be expected that homing behaviour shows day-to-day variations
which are in some way correlated with meteorological variations. Some correlations have in fact been found (Wallraff 1960; Dornfeldt 1996), but causal connections have so far not been established. Since the homing behaviour of pigeons with and without momentary olfactory access to natural air, as observed
by Kiepenheuer et al. (1993), fluctuated largely in parallel, it seems that temporal variations in true navigation are similarly intermingled with home-independent directional preferences, as has been observed in the case of spatial
variations (Sect. 3.7.1; see also Wallraff 1986). The thus apparently very complex interrelations have not yet been analysed.
7.5
Varying Home Site Conditions in Aviary Experiments
Homing, if not based on path integration (Sect. 4.2.1), requires a home site
characterized by particular location-bound properties. Pigeons, during their
long-term stay at this site, must learn these properties, but they must learn
even more. In order to find their way back from unfamiliar distant areas, they
must have acquired some knowledge about rules according to which environmental properties change depending on direction (and possibly distance)
from the home site. It has long been known that acquisition of this knowledge
does not require extended exercise flights in the vicinity of the loft, as might be
necessary in order to measure regular changes of relevant cues over some distance in various directions. Pigeons confined, since fledging, for several
months in an aviary, were well homeward-oriented when directly displaced
over 90300 km (Figs. 3.7 and 3.8; Kramer and von Saint Paul 1954; Kramer
1957, 1959a,b; Wallraff 1966a, 1970a, 1979). There was no indication that navigational capabilities were noticeably weaker than those of first-flight pigeons
which had been allowed to fly freely around their loft (Sect. 3.1.3). However,
this only holds true if the aviary, made of wire mesh, stood unobstructed in an
open landscape. By particular shieldings and other constructions, it was possible to modify the birds orientation behaviour.
7.5.1
Shielding, Deflecting and Reversing Winds
Most of the experiments described in this section do not directly prove involvement of airborne odours. They are, nevertheless, located here because
they influence the atmosphere which, as outlined above, appears to carry olfactory signals used for homing. Moreover, these experiments provide valuable
hints that might elucidate the mechanism of olfactory navigation.
No connection with the atmosphere was known when Gustav Kramer
(1959a,b), stimulated by an unintended observation of G. Pratt, started to raise
pigeons in an aviary positioned in a large cavity or surrounded by a wooden
112
n Fig. 7.18. Recoveries and vanishing bearings (inset) of pigeons raised in aviaries surrounded
by a wooden palisade at Wilhelmshaven (Wi) and released 150 km south of it near Osnabrck
(Os). These data should be compared with those of pigeons living in unshielded aviaries as shown
in Fig. 3.7, Os/Wi. (After Kramer 1959b, supplemented by Wallraff 1966a)
wall. Such pigeons, although often flying long distances, failed to orient homewards (Fig. 7.18). Later, it turned out that it was not the visual limitations that
were responsible for the breakdown of homing. If the upper part of the surrounding screen of one of the Wilhelmshaven aviaries was made of clear glass,
orientation remained poor (Wallraff 1966a). Glass screens prevented homeward
orientation also in southern Germany, whereas pigeons living behind visually
impervious screens that were pervious to airflow (Fig. 7.19) were as good in
homeward orientation as control birds from an unshielded aviary (Wallraff
1979).
These findings merely led to the conclusion that some dynamic factors of
the atmosphere (Wallraff 1970b) are in some way involved in pigeon homing.
The more specific next generation of aviary experiments were initiated by the
Papi group in Italy when the first results indicated the importance of olfaction
for home-finding. In the first of these experiments, Baldaccini et al. (1974) operated with two home sites 42 km apart. At fledging age a group of pigeons was
divided into four groups living in two small aviaries at each home site. The vertical walls of each aviary were visually impervious so that the birds could not
see the surrounding environment. One of the cages at each site, however, was
pervious to winds (similar to that shown in Fig. 7.19). For 3 days a week, all pigeons lived in and around a larger unscreened wire-mesh aviary at home site A
(Arnino near Pisa) which they could leave for exercise flights. During these
113
n Fig. 7.19. Left Schematic section of the louver palisade. The upper part of the aviary consisted
of wire mesh (M) and was surrounded by gratings of sinusoidally waved opaque plastic lamellae
mounted in wooden frames (F). The pigeons, when sitting on the perches (P), were exposed to
wind, but visually isolated from the surrounding environment. A, B Directions of recovery sites
at least 10 km away from the release site. Pigeons from the louver aviary (A) and from an aviary
with the louvers replaced by clear glass plates (B). Home site near Seewiesen southwest of Munich; release sites 97 km west (filled squares), 113 km east (filled circles), 117 km north (open
squares) and 26 km south (open circles). Heavy arrows show second-order mean vectors calculated from four first-order vectors; their homeward components are also given. Additional numbers indicate average shortening (+25 km) and lengthening (-26 km) of home distance of recovery sites against home distance of the release sites. (Drawing from Wallraff 1970b, data from
Wallraff 1979)
3 days, the birds carried a plastic mask applied to the upper beak, pressing the
nostrils so that smelling was impeded. After about 3 months the pigeons were
released halfway between home A and home B. Figure 7.20 shows initial bearings of those pigeons that were living in a cage pervious to airflow at site A or B.
It is evident that the B birds tended to fly towards home B, where they had
never flown around and never seen the environment, but where they could
smell the local atmosphere while feeling the wind. The A birds, which had coinciding visual, olfactory, wind and flight experience at site A, started preferably towards A. All of them arrived there, most of them on the day of release,
n Fig. 7.20. Vanishing bearings of pigeons simultaneously re-
leased between two home sites. At either site, A and B, one

group had been living during 4 days a week in an aviary
surrounded by visually non-transparent screens that were
pervious to airflow. The other 3 days a week all birds had
spent in and around an open wire-mesh aviary with allowance
of free flight at home site A, but with their nostrils occluded.
Filled symbols Birds from screened aviary at site A; open symbols birds from screened aviary at B. (Data from Baldaccini et
al. 1974)
114
n Fig. 7.21. A, B Aviary arrangements deflecting the wind clockwise (A) and counterclockwise
(B). Sides of cubic cages are 2.1 m long. In C, natural winds are shielded from the two lateral glass
corridors (space available to the pigeons 51.2 m). While the wind blows roughly along the axes
of the cages, an artificial air stream is produced by a fan blowing in the same direction as the outside wind in the left (control) compartment and in the opposite direction in the right (experimental) compartment. With winds deviating from the axis by more than 45 and under calm
conditions, the fans do not operate. (Papi 1986)
whereas most of the B birds were never seen again (four returned eventually to
A, five to B). The pigeons from the wind-shielded aviaries (not shown) were
oriented neither towards A nor towards B. Thus, flying around and feeling the
wind, but without the normal ability to smell, is insufficient to establish a home
site. The experiments indicate that homeward navigation from an unfamiliar
area requires both olfactory and mechanical experience of the home site atmosphere, while pinpointing the loft profits from visual experience of its immediate vicinity during exercise flights (cf. Fig. 3.8).
Other experiments aimed to manipulate wind directions, because winds
coming from different directions were thought to contain different composi-
115
n Fig. 7.22. Initial orientation of pigeons that had been living previously in aviaries with differ-
ently deflected, or non-deflected, winds as indicated (examples showing wind from south). The
diagrams (home upwards) show corresponding vanishing bearings at three different sites as distinguished by symbols (distances 9, 24 and 105 km). Arrows indicate mean vectors calculated
from three single-release vectors. (Modified from Baldaccini et al. 1975)
tions of odorous substances. Figure 7.21 shows arrangements designed to deflect or reverse the wind. The arrangements were inspired by the hypothesis
that the development of an olfactory map, correctly embedded in a compass
scale (Fig. 5.5), might be grounded on associations between varying odour
patterns and concurrently varying directions of the wind (e.g. Papi et al. 1972;
Papi 1976). Deflections or reversals of the wind would then rotate the compass
alignment of the map and thus result in corresponding deflections of the pigeons initial bearings.
The first three experiments with deflector aviaries (Baldaccini et al. 1975)
were in full agreement with this expectation (Fig. 7.22). Bird deflections were
sometimes smaller and sometimes larger than wind deflection, but the mean
was appropriate. In later repetitions, the deflections continued to show the predicted sense of rotation very consistently, but the amount of bird deflection
was mostly considerably smaller than the amount of wind deflection (Baldaccini et al. 1978; Kiepenheuer 1978c, 1979, 1982; Waldvogel et al. 1978; Waldvogel
and Phillips 1982, 1991). Unfortunately, all these experiments were conducted
at quite short distances (634 km, most releases at 920 km), where olfactory
navigation is probably not yet fully operational in all regions (Sect. 7.4.1). Since
most releases were conducted with pigeons that had previous opportunities to
fly at home or even had been released at the test site or other sites in the area,
116
n Fig. 7.23. Pigeons with the anterior commissure of the forebrain sectioned were kept, alternat-
ing every 3 days, in cages with reversely deflected winds. With winds deflected counterclockwise
(CCW), their left nostril was plugged (A), with clockwise (CW) deflection the right nostril (B). In
a series of tests the birds were released with either the one or the other nostril occluded. Vanishing bearings are pooled with home pointing upwards. Releases were successively at three sites,
13 km E (circles), 9 km NNE (squares) and 25 km SE (diamonds). Letters AK within the symbols
label the 11 individual pigeons. Depending on their homing success they participated differently
in the different releases (10 pigeons released twice at the same site with one or the other nostril
plugged). (Data and schema graphs from Fo et al. 1986)
many birds were, or could have been, more or less familiar with the landscape
they could see. Thus, it is likely that the olfactory component of their initial orientation was often fairly small, while a non-rotated visual component had similar strength or even dominated (see analogous effects in clock-shift experiments: Sect. 8.1.4). Moreover, it is not yet clear in what way wind deflections
interfere with homeward orientation, on the one hand, and with preferred
compass directions (PCDs), on the other hand. The latter are also known to be
influenced by wind directions experienced at home (Sect. 7.5.3). So far, we can
only say that the deflector effect is generally in good agreement with the hypothesis on the mechanism of olfactory navigation elucidated below
(Sects. 7.6.1 and 7.9), but a full understanding of the effect would require more
carefully and specifically designed experiments (Sect. 12.3.1 and Appendix).
Direct linkage between wind deflection and olfaction became clearly visible in
an inventive and sophisticated series of deflector experiments conducted by Fo
et al. (1986) with pigeons in which signal transfer from one brain hemisphere to
the other was prevented by sectioning the anterior interhemispheric commissure. The birds were kept alternately in one of two cages in which the wind was
deflected clockwise or counterclockwise, respectively. In one of the cages they
had their right nostril plugged, in the other cage the left one. Depending on
which nostril was plugged upon release, mean deviations from home were accordingly either clockwise or counterclockwise for the same pigeon (Fig. 7.23).
These results indicate that the pigeons had developed differently rotated olfactory maps in the two hemispheres.
Less influenceable by potentially disturbing side effects than many of the deflector experiments were the wind reversal experiments (Fig. 7.21C), most of
which were conducted at longer distances from home (Ioal et al. 1978; Ioal
117
n Fig. 7.24. Vanishing bearings of pigeons that had been living, at home, in one of three corridor
cages (cf. Fig. 7.21C) oriented in a WNWESE axis as indicated. Results of two releases. Pigeons
from cage with artificial wind roughly corresponding to natural wind (filled circles), from cage with
reversed wind (open circles) and from cage without wind (crosses). (Data from Ioal et al. 1978)
1980). The birds from the cage in which the wind was reversed against the simultaneously blowing natural wind flew quite consistently in a direction away from
home and contrary to the direction flown by the control pigeons (Fig. 7.24). Although, due to geographical constraints, most of the releases along the cage axes
were conducted to the east of home, it has been ensured by two releases from
western positions on the sea that the birds did not simply prefer opposing compass directions. It seems possible, however, that the particularly different air
coming over land or over the sea made the results particularly clear-cut.
The effects described in this section clearly concern the navigational and not
the motivational aspect of homing. While it might appear conceivable that pigeons living isolated from the outside world in a cage surrounded by solid
walls do not accept this site as an attractive home to which they would like to
return (Fig. 7.18), it is unlikely that such an interpretation might be valid for
glass walls but not for visually impervious walls as shown in Fig. 7.19. Predictable deflections and reversals of flight courses (Figs. 7.217.24) could definitely not have been caused by motivational deficits.
However, motivation apparently plays some role in the readiness to learn the
wind-related properties of the home site. Young pigeons kept in a screened aviary over a period of 34 months post-fledging were not homeward-oriented
even if they had subsequently been living for another 3 months in an unscreened aviary exposed to natural winds, whereas other birds caged unscreened from the beginning oriented towards home (Gagliardo et al. 2001a).
Yet early screened pigeons did show some homing abilities, though on a reduced level, if in the second phase they were allowed to make exercise flights in
the loft area (Odetti et al. 2003). Thus, during an early sensitive period, motivation for map learning (or imprinting) appears to be high enough to function
under suboptimal living conditions, while later on a retarded learning process
operates only under more favourable conditions (for late relearning, see also
Sect. 3.6).
118
7.5.2
Applying Artificial Odours and Winds
Inspired by the first results on the apparent role of olfaction in homing, Papi et
al. (1974) demonstrated that pigeons oriented their initial courses according to
associations they had made, at home, between artificial odours and artificial
winds. When smelling, upon release, either olive oil or synthetic turpentine,
odours to which they had been exposed in the home aviary with opposing
winds produced by blowers, the birds selected a direction corresponding to the
downwind direction they had experienced while smelling the respective odour
at home. A similar orientation downwind away from the odour source is illustrated in Fig. 7.25. In this case (Ioal et al. 1990), control as well as experimental
pigeons were held in normal air-pervious wire-mesh aviaries. Only the experimental birds experienced, from time to time, an artificial air stream from
north-northwest that carried the scent of benzaldehyde. When this scent was
added to the air ventilating all containers in which the birds were transported
n Fig. 7.25. Experimental pigeons were kept in a square aviary (2.52.5 m) through which, from
time to time, air was blown by two fans from north-northwest. This air carried the smell of
benzaldehyde evaporating from six dishes in front of the aviary. Control pigeons were kept in a
similar aviary without artificial wind and odorant. Peripheral symbols and heavy arrows indicate
vanishing bearings of pigeons smelling benzaldehyde during transport, at the release site and
during initial flight. Filled circles Controls; open circles experimental birds. Thin dashed arrows
show mean vectors from pigeons tested without application of benzaldehyde. (Data from Ioal et
al. 1990)
119
and held at a release site, and when thereafter a solution of benzaldehyde was
painted on all pigeons ceres and beaks, the control birds oriented unimpressed towards home, whereas the experimental birds flew southward irrespective of where they were released. Without application of the scent before
and upon release, however, the experimental birds were as clearly homeward-oriented as the controls.
These experiments demonstrate that pigeons at home consider the direction
of the wind that is correlated with a particular olfactory sensation. Re-experiencing this sensation at a release site, the birds behave as if they were at a position in an upwind direction with respect to this associated wind. A discussion
of this finding follows in Section 7.6.5.
7.5.3
A Digression to the Preferred Compass Direction (PCD)
Although interrelations with olfaction are as yet unclear, it should be noted
here that modified wind conditions in an aviary not only affect homeward orientation, but also modify the apparent PCD (Sects. 3.1.1 and 3.7.1). Screens
leaving an air-pervious corridor aligned roughly rectangularly to a PCD (as
observed in pigeons living in a normal unscreened aviary) produced a reversed PCD, whereas a corridor along the normal PCD axis had no effect
(Wallraff 1978a, 1979). Screening winds from only one direction led to a modified PCD deflected towards the unscreened direction (Ioal 1996; see also Ioal
and Benvenuti 1983). It is unresolved, however, whether an experimentally induced or modified PCD is equivalent to the spontaneous PCD or whether it results from a superimposed distorted olfactory map (Wallraff 2001, p. 5 and
Fig. 2). Smelling air coming from a certain direction but not being able to determine that direction would strongly interfere with an assumed map-building process (Sect. 7.6.1).
The spontaneous PCD is not generally directed towards or away from the
most frequent wind direction at the home site. Pigeons housed in lofts only
some 825 km away from each other preferred very different compass directions (Schmidt-Koenig 1963a; Wallraff 1970a). Exposure to selected natural
winds from opposite directions did not influence the PCD (Wallraff 1978a). Pigeons coming from a circumferentially shielded aviary not only failed to orient
homeward (Sect. 7.5.1), but also showed, if at all, an underdeveloped PCD
(Ioal et al. 1978; Wallraff 1978a, 1979). Good exposure to winds possibly tends
to favour the expression of a PCD (cf. Wallraff 1990a, p. 104).
Most of the deflector-loft experiments were conducted at a time when release
from sites in central symmetry had not yet become a general rule. Thus, homeward orientation, on the one hand, and PCD, on the other hand, as could be compared in other contexts (e.g. Figs. 3.4, 7.2 and 7.8), cannot be compared as regards effects of wind deflection. Winds in combination with olfaction obviously
determine the development of the home-finding mechanism (Sects. 7.5.1 and
7.5.2), but it is as yet unclear to what degree one or both factors determine the
120
development of the PCD. This latter development occurs also in anosmic pigeons (Wallraff 1978a; Papi et al. 1989), but complete independence of olfaction
is not yet definitely proven. Also, it has not yet been tested whether screened or
deflected winds modify the PCD of permanently anosmic birds, i.e. whether the
wind-induced deflections so far produced in pigeon orientation reflect only influences on olfactory map building or on PCD development or whether both developments are interrelated (Sect. 12.3.1 and Appendix). Olfactory deprivation
only upon release, but not previously at home, did not abolish the effect exerted
by preceding wind deflection in the home aviary, possibly because the PCD had
been deflected (Kiepenheuer 1979; for discussion see also Wallraff 1981a,
1990a).
7.6
Spatial Structures in the Chemical Atmosphere
The manifold experiments described above (Sects. 7.17.5) leave hardly any
doubt that the atmosphere contains olfactory (= chemical) key signals used by
pigeons for home-finding. This outcome was unexpected and is not immediately plausible. In order to achieve some plausibility, if at all possible, it was
necessary to search for atmospheric-chemical structures that might provide a
potential physical basis for the observed behavioural performances. A precondition of an empirical approach was a working hypothesis specifying some
guidelines according to which acquisition of potentially elucidating data could
be obtained.
7.6.1
A Working Hypothesis
In Section 4.2.2, we proposed that birds, being able to home from unfamiliar
areas, should have established a gradient map telling them in which direction
from home specific physical quantities increase or decrease with some regularity. In the previous sections of this chapter, we concluded that trace gases
dispersed in the atmosphere appear to carry the positional information that is
necessary for birds to home. If both the proposition and the conclusion are
valid, spatial gradients in airborne trace gases should exist which are exploitable for navigational purposes. Owing to the instability of the atmosphere and
to the low concentrations of such trace gases, this supposition appears unrealistic. Owing to the bulk of experimental evidence, however, there is hardly an
alternative approach towards solving the pigeon homing problem except to
develop an hypothesis supporting this supposition despite intuitive resistance.
A hybrid map consisting of olfactory and other components is also conceivable, but there are no hints suggesting that, in unfamiliar areas, other cues
might be necessary for home-finding as well (Sects. 4.2, 5.4 and 6.3.3). Therefore, only atmospheric factors shall be considered.
7.6 Spatial Structures in the Chemical Atmosphere
121
The classical idea of bi-coordinate navigation proposes two gradients intersecting at a sufficiently large angle, ideally at an angle of 90 (cf. Fig. 5.5).
Straightforwardly applied to the atmosphere, the concentrations of two trace
compounds should have fairly stable gradients in two different directions extending monotonically over several hundred kilometres. Moreover, birds should
be able to measure absolute concentrations of two chemosignals independently
of each other. Both these propositions can hardly be expected to be met in the
real world. Physically, absolute concentrations are very variable depending on
weather and wind. Physiologically, determination of absolute intensities is problematic, and separate determination of two absolute intensities within the same
modality even more. Somewhat closer to reality might be the expectation that
birds respond to ratios among quantities of two or more compounds independently of their absolute concentrations. Based on such ratios, we distinguish between different fragrances of flowers, fruits or perfumes whose specific smell
results from the proportional mixture of a number of chemical compounds.
Within a wide range of absolute concentrations, we perceive a stable odour quality of a particular mixture. We do not recognize its components separately, but
we feel a change in quality when ratios between compounds change. In the atmosphere, ratios are less variable than absolute concentrations (cf. Fig. 7.27).
If birds were using gradients of ratios between compounds to determine position, two compounds would not be sufficient, because a ratio between two
quantities could provide only one coordinate. With a minimum of three compounds, however, any position would have a unique ratio pattern, provided
that the ratio gradients are perfectly monotonic and sufficiently divergent, as
illustrated schematically in Fig. 7.26. Position determination would not be affected by varying absolute concentrations (density of dots in Fig. 7.26A) which
might, for instance, fluctuate with temperature. Decisive were the proportions
among the contributing compounds (pie charts in Fig. 7.26B). If birds knew in
which direction the ratios change in what way, they could deduce the direction
they should fly in order to reduce the proportional difference from the home
spectrum (peripheral vector diagrams in Fig. 7.26B).
Referring to ratios rather than to absolute quantities does not solve the problems connected with varying wind directions. If winds were to rearrange all
proportions completely, there would hardly be any chance that odours might
be applicable for position determination. If winds were to displace a certain
gradient pattern as a whole, however, its internal structure might remain fairly
stable, and then winds might even be helpful. With a northeasterly wind, for instance, the pattern shown in Fig. 7.26A would be displaced so that the relatively
many black molecules in the northeast would be shifted towards the location
of a pigeon loft in the centre. As a navigationally positive consequence, the pigeons might learn, at home, that winds from northeast are correlated with a
relative increase in compound black, winds from the southwest with a relative
decrease. When later released at a site with an unusually large portion of black,
the birds, remembering this correlation, might be induced to fly away from
their current position towards southwest. A mechanism of this kind would be
122
n Fig. 7.26. Schema of an olfactory navigation system based on (unnaturally steep) ratio gradients
of three fictitious compounds in the atmosphere. A A two-dimensional field within which the portions of the compounds, distinguished by symbols, vary along differently oriented gradients (arrows indicating upslope directions). A pigeon loft is located in the centre. B The pie charts give percentages of compounds in the centre as well as at eight sites around it (radius = three-quarters of
half side length in A, e.g. corresponding to 150 km). In the peripheral graphs, differences to the
home ratio are drawn as percentages of this ratio and attributed to the gradient directions (numbers give these percentages). If the difference is negative, the resulting vector points upslope, if it is
positive, it points downslope (thus tending to reduce the difference by moving in that direction).
Thick arrows indicate flight directions calculated from the three site-specific vectors. Note that
along the westeast axis and along the diagonal axes the portion of one of the compounds remains
almost constant (see near-zero difference values and associated absence of vector bars). (Wallraff
2004)
in good agreement with the aviary experiments described in the previous section (Sect. 7.5). However, it would be connected with the serious negative consequence that the spatial position of the whole two-dimensional pattern of ratios would be unstable in time. No invariable set of ratios would characterize
the home site or any other location. Thus, additional preconditions must be introduced to make the mechanism operational.
The simplest solution would be that wind-induced shifts of proportions, although recognizable for the birds, are small against differences depending on
position. The problem of ambiguity (does the relative increase in black indicate wind from or displacement to northeast?) would then arise only at short
distances where olfactory navigation may have its lower limit. Alternatively, the
birds could take the current wind conditions into account and consider a flexible, wind-dependent ratio spectrum as a home reference. If they do so, however, they apparently do it not by means of direct measurements of wind direction using their sun compass upon release. In this case, the sun compass would
be involved in the process of position determination. Clock shifts would cause
false conclusions about wind direction, hence about the home spectrum to be
taken as a reference and hence about the birds position in relation to home.
123
Recalculation of initial bearings only on the basis of differences in sun azimuth

would not lead to a level of home orientation as high as that of control birds
(Fig. 5.2, C versus A). Thus, the birds might use indirect signals involving more
reliable information on prevailing wind conditions in a larger spatial and temporal scale than short-term measurement of the momentary local wind (which
sometimes does not exist at all). It has been shown that, theoretically, such information could be deduced from such atmospheric trace substances whose
proportional contribution to a set of utilized compounds varies with wind direction but varies very little depending on position (Wallraff 1989c).
All these considerations concern conceivable basic structures (for application in a navigation model, see Wallraff 1989c). Even if they would fit reality in
principle, it cannot be expected that real structures are so schematically perfect. Also, it need not be proposed that they are. Home orientation of pigeons is
quite inaccurate, temporally variable and intermingled with substantial stochastic noise (Sect. 3.7.3). There are reasons to assume that noise is more inherent in the environmental signals used rather than in the animals activities
of processing these signals (Wallraff 1994a). To reach the levels of performance
obtained by pigeons, spatial signals need not be very reliable and unambiguous. A larger number of considerably noisy gradients can lead to similar performances as a smaller number of more regular gradients (cf. Fig. 4.2). Thus, it
would be advantageous if birds would use more than the theoretical minimum
of three airborne compounds.
7.6.2
Gradients in Ratios of Trace Gases
With the above working hypothesis in mind, it was possible to ascertain
whether the atmosphere does in fact contain positional information potentially
useful for navigational purposes. Over four summer months in three years,
airborne trace gases were collected within a radius of 200 km around Wrzburg, Germany, which was previously the home-site centre for many homing
experiments with pigeons (e.g. Figs. 3.4 and 7.3). Using small tubes filled with
adsorbent carbon material, volatile organic compounds (VOCs) were sampled
at the centre as well as at 96 sites regularly distributed around it. The collected
samples were analysed by gas chromatography and statistically evaluated
(Wallraff and Andreae 2000). Usually 200350 compounds were recorded per
chromatogram, but statistical evaluations were focused on 16 compounds that
were clearly identified in each of the 224 analysed samples obtained in the
course of 16 crosswise symmetrically arranged trips under variable weather
conditions with winds from all directions. Amounts of VOCs contained in the
different samples varied considerably, but the shapes of the chromatographic
peak patterns were similarly reproduced in all samples (Fig. 7.27). Data of interest were the proportional variations of peak areas within these patterns and
their spatial relationship to the geographical positions at which the samples
had been collected.
124
n Fig. 7.27. Parts of two typical gas-chromatograms showing the peaks of 16 more thoroughly
analysed chemical compounds in the atmosphere. Numbers give C (carbon) indices used for
identification. Ordinates are in relative, but corresponding units; note the different ranges. Boxes
contain example calculations using areas (relative units) and resulting ratio of one pair of compounds. (Wallraff and Andreae 2000)
The basic unit for calculations was the amount of one compound as a percentage of the sum of two or more compounds within the same chromatogram
(see boxes in Fig. 7.27). If the 16 omnipresent VOCs were combined pairwise,
120 combinations were available. For each compound within a pair, the mean
percentage resulting from 192 chromatograms (two per peripheral site) was
determined. From the standardized differences from this mean, together with
the spatial distribution of the sampling sites, a parameter of eccentricity was
computed which indicates the degree of spatially oriented separation of
above-mean and below-mean ratios (for details of computations see Wallraff
and Andreae 2000). In the case of a spatially uniform distribution of the differences, eccentricity would be zero. Most of the actually observed values are considerably larger (filled columns in Fig. 7.28). However, with a limited number
of measurements and sites, even random distributions would usually deviate
from zero to some degree. Possible random deviations can be determined by
linking the chromatograms not to their actual sites of origin but to randomly
selected other sampling sites. Average deviations from zero resulting from 105
independent replications of such mixing procedures are shown by the crosshatched histogram in Fig. 7.28. The highest actual values were not reached in
any of the 105 random permutations; the 99.9% range of their medians does
not include the actual median. The circular inset in Fig. 7.28 shows the individ-
125
n Fig. 7.28. Statistical expression of gradient character in spatial distributions of ratios among
pairs of atmospheric trace compounds, measured in terms of eccentricity. Filled columns give
the frequency distribution of eccentricities of 120 ratios among 16 pairwise combined VOCs resulting from the actual geographical distribution of air samples. Cross-hatched columns show
mean distribution of the same ratios resulting from 105 permutations in which air samples were
distributed to the sampling sites at random. Filled circle at 16.5 km indicates the median of the
actually observed distribution, dot at 7.7 km the second-order random median with the 99.9%
range of medians per replication. Circular inset (radius 200 km, north above) shows the ratios
among two compounds, C4.0 and C5.3, at the 96 sampling sites. Sites where the portion of compound C4.0 was above mean are marked with filled circles, those where it was below mean with
open circles (their sizes indicating differences from the mean). Ellipses give 95% confidence areas
of mean locations of black and white sites. (Modified from Wallraff and Andreae 2000)
ual sampling sites, with the corresponding standardized ratios between the
two compounds providing the highest eccentricity value.
Comparable results were achieved when ratios were determined not among
pairs of substances, but among triplets, sextets etc., as would be necessary to
derive positional information (Sect. 7.6.1). Also, not only the 16 omnipresent
compounds were tested. Calculations including VOCs that were found in at
least 90 or 80% of the chromatograms (up to 72 compounds) led to basically
similar results. These results, however, were less reliable, because they included
very small peaks whose identification and quantification involved larger
ranges of possible error.
Each of the 16 substances, being always one partner among 15 pairs, is differently localized in the black histogram of Fig. 7.28. If the six highest ranking
compounds, which are farthest outside of the range reachable by chance, are
separated and their ratios calculated not pairwise but as a percentage of the
sum of all six, their relative amounts form mountainous landscapes with differently oriented overall slopes (Fig. 7.29). The distributions are quite noisy,
126
n Fig. 7.29. Relief maps showing standardized ratios of six compounds among each other as
measured in the region around Wrzburg (i-hexane = 2,2-dimethylbutane in Fig. 7.27). Areas
where the relative abundance of the respective compound was above its overall mean are light
grey, those with values below average dark grey. Small diagrams show means SEM in classes of
distance along the computed gradient slope whose upwards direction is indicated (ordinate =
difference from overall mean in units of standard deviation). (Wallraff and Andreae 2000)
but in principle they fit the expectations illustrated in Fig. 7.26. Each site in the
inspected region of Germany is characterized by a particular ratio spectrum of
the six substances.
Most likely, none of these six compounds is actually used by pigeons to find
the way home. Except for isoprene, they are all anthropogenic hydrocarbons.
However, inhomogeneous distribution of sources and sinks, chemical conversions, variable lifetimes and atmospheric transport patterns are equivalent in
anthropogenic and in naturally emitted biogenic VOCs. If ratio gradients exist
among some airborne compounds, they can be expected to exist among other,
also natural, VOCs in a similar fashion as well. The chemical tracers analysed
in detail were predominantly those that were present in the air at fairly high
concentrations and hence were most accessible to the applied method. It was
evident, however, that several other, not always detected and not chemically
identified VOCs have similar gradient patterns in the ratios among each other.
The gradients so far recognized using fairly crude techniques are probably not
the clearest ones actually existing in the atmosphere.
Less remarkable than the revealed spatial order is the observation that ratios
among VOCs vary temporally depending on wind direction. Such influences
could have been expected. It is remarkable, however, that at least some spatial
gradients hold their alignments fairly stable even with opposing winds
(Fig. 7.30) or air masses arriving from different directions. It is further remarkable that a correlation exists between upwards direction of the spatial ratio gradient of a given compound and wind direction under which the ratio increases (Fig. 7.31). In the analysed data, the two directions diverge by a mean
angle of approximately 45. It is not yet clear whether this angle is generally
127
n Fig. 7.30. Mean percentages of the six substances (see Fig. 7.29) in various directions from the
centre with varying directions of the wind during air sampling (wind speed 5 km/h). Directions
of sites from the centre (irrespective of distance) were combined in angular classes and running
means of the percentage per compound determined in steps of 5 around the circle. Curves were
calculated for two pairs of mutually exclusive subsets of chromatograms: with winds from semicircles north and south (thick curves without dots) and semicircles west and east (thin curves with
dots). Site-independent means per wind class are shown by thin horizontal lines. Percentages at
the ordinates indicate ranges of variability for each substance. (Wallraff and Andreae 2000)
representative or is more or less biased by chance-based predominance of anticyclonic weather conditions during data sampling. Also, in some individual
(and perhaps decisive?) substances, the angle is close to zero, as would be expected according to the above hypothesis (Sect. 7.6.1) which considers the aviary experiments (Sect. 7.5). At any rate, a systematic relationship seems to exist
between VOC ratio variations as correlated with wind, on the one hand, and location, on the other hand. Thus, also this hypothetical prediction has been met,
128
n Fig. 7.31. Angular relationships between wind-dependent and site-dependent directions of

eccentricity of ratios among 525 pairs out of 72 compounds (selected according to frequency of
presence in the chromatograms and to amounts of eccentricity). Example for reading: in 25% of
the compound pairs, the maximum relative increase of one of the partners occurred with winds
deviating 4515 clockwise from the direction of its maximum spatial increase (e.g. ratio of compound A relative to B, which was spatially largest in regions northwest of the centre, was temporally largest with winds from north). (Modified from Wallraff and Andreae 2000)
albeit not in a most simple and ideal way. The presently available data set is too
small to enable definitive conclusions on long-term geometrical relationships
between wind and the spatial distribution of trace gases.
7.6.3
Applicability of Atmospheric Gradients to Navigation
In a next step we ask whether the six distributions shown in Fig. 7.29 can be
used as a map basis for home-finding. Assuming that computer-borne model
pigeons are thought to have gained, before displacement, some approximate
knowledge of the compass alignment of the six gradients, it is possible to deduce, for any site within the investigated area, from the six local ratios a direction that aims to approach the centre (Fig. 7.32; details of computations in
Wallraff 2000a). Vectors in Fig. 7.32A are comparable with those shown above
in the theoretical schema (Fig. 7.26B). When the whole field of 200-km radius
is inspected (Fig. 7.32B), some of the computed directions are seen to be completely wrong, but at 94% of the sites the deviation from home is less than 90
and the overall homeward component is, with a value of 0.77, better than usually found in experiments with pigeons. In Fig. 7.33, computer results based on
observed atmospheric data are compared with observations obtained with
real pigeons. Figure 7.33A not only summarizes (with its heavy curve) the directions exhibited in Fig. 7.32B, but also shows that even VOC data obtained
129
n Fig. 7.32. A Computed initial bearings at 12 sites 150 km from Wrzburg, deduced from the
atmospheric data shown in Fig. 7.29. From knowledge of gradient directions (which birds hypothetically determine at the home site by correlating winds with odours) and local measurement
of six ratios as differences from their overall means, six individual vectors were calculated and
combined to a resulting vector (arrow), whose length may correspond to the degree of angular
scatter in a sample of real pigeons. B With the same atmospheric data, estimates of homeward directions were computed for an array of 196 sites within a radius of 200 km around Wrzburg. The
sector between direction towards the centre and computed direction is indicated by solid black.
Areas of circles are proportional to lengths of computed homing vectors (relative units). The
overall mean vector resulting from the 196 directions deviates from home by 2 and has a length
of 0.78. (Wallraff 2000a)
while winds came from opposing directions (thin curves) provide results in
the range of performances of living birds.
Complete homing routes produced by the model are shown in Fig. 7.34, A
and B. The direct parametric outcome as computed from the measured VOC
ratios (A) represents individual flight paths derived from exactly these data by
means of a given algorithm. In nature, two sources of variability can be expected. First, the atmospheric environment is unstable. Due to temporal fluctuations, each pigeon meets somewhat different ratios (cf. Fig. 3.19). Second,
the birds do not perform precise measurements, but deduce their directional
decisions from probabilistic estimates. Owing to these external and internal
sources of noise, each individual bird flies its individual path as simulated in
Fig. 7.34B. If not the whole paths are documented, but only end points of interrupted flights, a picture simulating distributions of recoveries can be created
(Fig. 7.34C; compare Figs. 3.8 and 7.3).
Results still in the range of performances of living pigeons cannot only be
reached by using the six paradigmatic VOCs used to produce Figs. 7.327.34,
but also by using several other combinations of three or more compounds. Of
course, however, not every arbitrary combination is suitable.
In the above simulations, the directions of the VOC gradients were taken
from Fig. 7.29 and considered as known. Experiments with pigeons confined
130
n Fig. 7.33. Homeward orientation of computer model pigeons using VOC ratios of Fig. 7.29 (A)
and of real pigeons (B). A Heavy solid curve shows the frequency distribution of absolute angular
deviations from the centre as calculated in Fig. 7.32. Thin curves show corresponding results obtained with winds ( 5 km/h) coming from only one hemicircle as indicated (e.g. from north
89). Dotted curve refers to wind speeds >5 km/h irrespective of wind direction. Column n gives
the number of air samples on which the computations are based, column %<90 the percentage
of sites at which the computed direction deviates by less than 90 from the direction towards the
centre. B Corresponding frequency distributions of mean vanishing bearings of real pigeons in
nine series of experiments conducted at fairly symmetrical release sites around home. Open symbols refer to inexperienced pigeons displaced for the first time. For details of sources see original.
(Wallraff 2000a)
131
n Fig. 7.34. Homing routes of computer model pigeons using the VOC ratios of Fig. 7.29; eight
release sites 150 km distant from home near Wrzburg. A Parametric output of the model without noise. B At each site, ten pigeons starting, determination of positions distorted by some stochastic noise. C Simulation of recovery sites: end points of shorter routes; ranges of random errors greater than in B. (Wallraff 2000a)
in aviaries in which winds were manipulated in various ways (Sect. 7.5) suggest that the birds achieve this knowledge by associating varying VOC ratios
with contemporaneous wind directions. Figure 7.31 shows that the two parameters are in fact correlated. It is not yet clear whether the birds can ignore the
current weather conditions (especially wind directions) during the time of
their homing flight (as may be suggested by Fig. 7.33) or whether they have to
take these conditions into account, possibly by using additional information
derived from VOCs as suggested in an earlier model (Wallraff 1989c).
7.6.4
Assumed Dependence on Geographical Peculiarities
Thus far, existing data suggest that pigeon homing from unfamiliar areas is
based on olfactory atmospheric signals everywhere on the earth (Sect. 7.4).
They also suggest that navigational performances achievable by using such
signals are not everywhere equivalent (Figs. 7.4 and 7.16). Considerable regional variability is, in fact, to be expected when soft atmospheric cues are
used which depend on geographical conditions such as climate, vegetation, flat
or mountainous country, sea and land constellations etc. (and not hard astronomical or geophysical cues implying global rules).
There are particular indications that the ranges of sufficiently monotonic gradients are limited by orographical structures, i.e. that higher mountain ridges
may interrupt the continuity of suitable indicators of position. With the olfactory ratio gradient model in mind, we can now interpret the results of the
above-mentioned transalpine displacements (Sects. 7.3.3 and 7.4), which are
summarized in Fig. 7.35B. Assume that the portion of a compound A, as a percentage of A+B, regularly increases (or decreases) over the relatively flat area of
Germany, may perhaps hold a plateau over the Alps and has a reversed slope
down to the River Po valley.Such relationships among some aerial trace gases do
in fact occur (Fig. 7.35A,C). Pigeons from a loft at Andechs close to the northern
132
n Fig. 7.35. Diagrams in B summarize directions of recoveries after transalpine displacements
as shown in detail in Figs. 7.15 and 7.17 (north above). Double-headed arrows point towards
home, heavy arrows give mean vectors calculated from recovery directions (radius of circle =
vector length 0.25). Positions of three home lofts are marked by squares. In A, 44 sites are indicated at which samples of VOCs were collected approximately along the AndechsMantua meridian at different altitudes during two southwardnorthward trips in July 1995 (diamonds) and
2001 (squares) and during the investigations described by Wallraff and Andreae (2000) (plus
signs). In C, running means of ratios among three pairs of substances as derived from these data
are shown. Each curve gives the amount of C4.0 (= n-butane, see Fig. 7.27; upper abscissa) as a
percentage of the sum of C4.0 and one of three other compounds (whose percentages are given at
abscissa below). Straight lines are drawn through the values measured at the release site Mantua
whose northsouth position is labelled zero (see Appendix for more details)
133
edge of the Alps, when released in the Po valley near Mantua, would perceive a
ratio among A and B which north of the Alps is correlated with a change of position from Andechs towards north. Thus, the birds would conclude to be north of
their loft and therefore fly southwards away from home. Pigeons from a loft at
Wrzburg, released at the same site in Italy, would observe a ratio that they similarly would meet some 100 km or so south of home.Consequently,they would fly
northwards, i.e. towards home. Italian pigeons housed at Florence, when released in Germany in the neighbourhood of Andechs, may find there a ratio indicating, also south of the Alps, a position north of home. Consequently, they fly
correctly southwards.When released 200 km farther north near Wrzburg,however, Florence pigeons appear to be unable to draw any meaningful conclusions
from the local olfactory spectrum and therefore behave disoriented.
Of course, this interpretation of the transalpine experiments is highly hypothetical. Not all ratios among compounds follow the patterns shown in
Fig. 7.35C, but basically similar up-and-down gradients are more frequent
than expected on a pure random basis. On the whole, geographical variability
of olfactory navigation, including varying levels and distance ranges of functionality and including false orientation under particular release-site homesite constellations (Sect. 7.4), is well compatible with the ratio gradient hypothesis (Sect. 7.6.1).
7.6.5
The Short Distance/Long Distance Problem
Initially, even the discoverers of olfactory navigation did not believe that homing over hundreds of kilometres could be based on atmospheric signals perceived in such far-distant areas. Homing over such long distances was assumed
to make use of other (yet unknown) navigational cues and/or of olfactory signals sampled during the first part of the outward journey (e.g. Papi et al. 1973;
Papi 1976). Now, the first possibility appears outdated; even over long distances, no other environmental signals can replace lacking olfactory signals.
Utilization of olfactory outward journey information, on the other hand, is
necessary over somewhat longer distances in some regions only, while in others the navigationally exploitable atmosphere covers a radius of several hundred kilometres (Sect. 7.4.1). Not for these long distances, but for short distances around the loft, especially for very short distances, an early presented
explanatory hypothesis (Papi et al. 1972; Papi 1976), later labeled the mosaic
hypothesis, appeared appropriate, for which the experiment shown in
Fig. 7.25 can serve as a model. While winds blow from a particular direction, a
pigeon loft may lie within the plume of an odour source such as, for instance, a
factory, a field of flowering rape, a forest or a lake. The scent from that source is
very prominent with winds from that particular direction and almost or completely lacking with other winds. When smelling that odour at an unfamiliar
site, a pigeon may conclude to be displaced toward the respective odour source
and thus fly the wind direction it has learned to be correlated with that odour
134
at home. A pigeon from another loft located only a few kilometres away on the
other side of the factory or forest, however, would fly in the opposite direction
when smelling that odour. So what would their home-site experience help the
two pigeons when they were displaced over 100 km in the same direction from
both lofts?
For homing over longer distances such conspicuous local odours, in an
all-or-nothing way correlated with particular winds, should be confusing
rather than helpful. A navigation system applicable to homing from remote
unfamiliar areas requires long-range gradients originating from omnipresent
compounds and not an unpredictable patchy pattern of a variety of odour
sources [which might, at best, constitute a basis of an olfactory mosaic map
within a familiar area (Fig. 4.1A,B)]. It is, however, difficult to imagine that a
simple system as proposed in Fig. 7.26 provides useful positional information
at distances of 200 and 20 km alike. Gradients useful over 200 km can hardly be
expected to be so uniformly monotonic and noiseless that they can be
navigationally exploited less than 20 km away from home with similar reliabil-
n Fig. 7.36. Statistical expression of gradient character in spatial distributions of ratios among
120 pairs of 16 atmospheric trace compounds in three concentric ranges of distance around a
centre near Wrzburg, Germany. The material is identical with that used to generate Fig. 7.28.
The 96 sampling sites were divided into three equal parts at distances 2585 km (mean 55 km),
85145 km (mean 115 km) and 145205 km (mean 175 km). Vector lengths were calculated from
spatial distributions of 120 ratios among two compounds, each distribution obtained at a set of
32 symmetrically arranged sites. Curves are analogous to those in Fig. 7.28 with the difference
that only direction and not distance from the centre was considered. As a result, the three curves
of average random expectation (indicated by thin lines) are almost identical for the three classes
of distance. Most divergent from these curves (P<0.0001) is the frequency distribution obtained
from the air samples collected farthest away from the centre (filled circles) and least divergent the
distribution resulting from the sites less than 90 km away (stars); median distances (open circles)
range in between. (Extracted from material as reported by Wallraff and Andreae 2000)
135
ity. In fact, the 16 analysed VOCs found in the atmosphere everywhere within a
200-km radius around Wrzburg (cf. Fig. 7.28) revealed increasingly regular
gradient characteristics with increasing distance from the centre (Fig. 7.36).
Correspondingly, first-flight pigeons housed at that centre tended to be better
homeward oriented with increasing distance from home (Fig. 3.5).
In Tuscany (Italy), olfactory conditions seem to be different. Homeward orientation over short distances (down to 7 km: Meschini 1983) is much better than in
Germany (Sect. 3.4.1; Figs. 3.3 and 3.20; see also Wiltschko et al. 1987d). The upper distance limit, however, up to which reliable positional information can be
extracted from the atmosphere, was found to be reached already around 100 km
(Benvenuti et al. 1994; see also Baldaccini et al. 1982; Sect. 7.4.1; Fig. 7.16). We
may speculate that, owing to the more pronounced and smaller scaled geomorphological structures in this region (coasts and mountains), useful atmospheric gradients are steeper, but their ranges of sufficient monotony more limited. To achieve an olfactory navigational system that is well operational over
short and long distances, it would probably need to be more complicated than
assumed in the above simplistic model (Fig. 7.26). Possibly, measurement of not
only direction but also velocity of winds might appropriately integrate different
substances that are most suitable for different ranges of distance (Sect. 7.7.2).
Even within a radius of only some 50 km, it is difficult to see how a system
based on non-quantified presence or absence of particular odours (according
to the early mosaic hypothesis by Papi et al. 1972; Papi 1976) might operate at
any site within this range and with reference to different home sites (cf.
Wallraff 1974a, 1980b, 1991a, 2001). It is, therefore, difficult to interpret the
benzaldehyde experiment described above (Fig. 7.25). Either a particular
odour perceptible at high concentrations with wind from one direction and
(almost?) absent with other winds would have a local source (and thus would
not contain positional information for areas far away from home) or its presence or absence would depend on current wind direction everywhere within a
large region (landward and seaward wind in coastal areas; Scirocco coming
from south, Mistral from north etc.). Presence of that odour alone would then
not be site-specific and could thus not indicate a direction in which to fly.
Against this background, I see three little-attractive possibilities to interpret
the benzaldehyde experiment: (1) The home-site condition induced an escape
response in a direction that at home would have led away from a strong unpleasant odour source; it has nothing to do with homing at all (and its causation would not be easily understandable). (2) The pigeons behaved as predicted by the mosaic hypothesis. Then their behaviour simulated a natural
response to a short-distance displacement towards a local odour source, but is
not relevant for olfactory long-distance homing. As the latter exists, we would
have to presume the co-existence of two different mechanisms of olfactionbased homing. (3) The pigeons incorporated the unnaturally extreme odour
and wind condition into their sensitive odour and wind mechanism as well as
possible according to the rule: fly in the direction in which, at home, the wind
blew while the olfactory input was most similar to the current olfactory input.
136
They would then have adjusted the highly artificial all-or-nothing relation to
a quantifying scale which usually operates with much weaker more-or-less
proportional relationships.
Compared with the other alternatives, the last interpretation appears to be
the most plausible. In natural life, wind-correlated local odour plumes would
not severely disturb homeward orientation, because the birds would rarely encounter a similarly composed odour elsewhere again (as they did in the
Fig. 7.25 case), and if so, they would leave it quickly and then reorient their
course according to the more regular VOC ratios (as they obviously did in the
Fig. 7.25 experiment: no difference between experimental and control pigeons
in homing performance).
7.7
Avian Olfactory Perception: A Suitable Tool for Navigation?
At first glance, it is not only the atmosphere that appears unsuited as a medium
allowing spatial orientation. It is also the birds sense of smell that appears unsuited to make appropriate use of volatile substances diluted in the atmosphere. At least four major problems would have to be solved in this respect.
They focus on the possibility that olfactory signal processing as applied in
avian navigation might differ from what we know from introspection and
about classical olfaction.
7.7.1
The Problem of Sensitivity
Although it is now well known that the sense of smell is not generally underdeveloped in birds and that many species apply it in a variety of ecological contexts
(e.g. Wenzel 1973; Roper 1999), it has not been shown so far that it is sensitive
enough to perceive atmospheric trace gases at the very low concentrations at
which most of them mostly occur (see Waldvogel 1989). While in rural areas
concentrations of biogenic VOCs (e.g. isoprene and terpenes) and several sorts
of other low and medium molecular weight hydrocarbons (except methane and
ethane) rarely exceed 1 ppb (part per billion) (e.g. Rudolph and Khedim 1985;
Kesselmeier and Staudt 1999), the lowest olfactory threshold for any substance
measured by conventional methods before 1999 was at a level of 100 ppb in pigeons and 10 ppb in any one avian species at all (Roper 1999). However, conditioned or unconditioned laboratory responses or non-responses to some arbitrarily or in other contexts chosen chemicals at varying doses tell us little about
the possible sensitivity to thus far unidentified key compounds that might be
used under natural conditions in the context of navigation. Moreover, decisive
substances may act over longer time periods in a subconsciousway not eliciting
responses in such experiments (Sect. 7.7.4).
7.7 Avian Olfactory Perception: A Suitable Tool for Navigation?
137
7.7.2
The Problem of Odour Discrimination
According to the hypothesis outlined above (Sect. 7.6.1), it would not be necessary to distinguish between hundreds of compounds or bouquets of compounds. A limited number of specific compounds might be sufficient. Their
relative proportions should be distinguished, if not by individual distinction
(as in the model calculations above: Figs. 7.26 and 7.32), then by discerning the
resulting composed-odour quality which varies systematically with winds
from different directions (Sect. 7.6.2). In Fig. 7.26B, a bird would then not discern the sectors of the pie graphs separately, but would sense each pie having a
different flavour resulting from the respective proportional mixture. However,
separate relative quantification would probably make the system more effective and would probably be necessary if it is not as simple as assumed in the
Fig. 7.26 schema. It is likely that the number of included compounds is greater
than assumed there and that their ratio gradients are considerably noisy,
showing small-scale spatial and temporal fluctuations. Also, the distance
ranges over which the gradients are sufficiently steep and monotonic may differ. To allow short-distance and long-distance navigation (Sect. 7.6.5), the
mechanism might refer to different sets of compounds and thus would have to
discriminate between them.
7.7.3
The Problem of Compound Selection
Certainly most of the trace substances diluted in the atmosphere are unsuited
as indicators of position (cf. Wallraff and Andreae 2000) and therefore should
be ignored by navigating birds. How do they select those that are suitable? As it
was possible to elicit predictable orientation responses to three arbitrarily
chosen odorants (Sect. 7.5.2), it seems unlikely that a certain set of chemical
compounds used for navigation is genetically fixed in a qualitatively exclusive
manner. Nevertheless, preference for specific compounds could have been obtained via sensitivity. The expected outstanding sensitivity (Sect. 7.7.1) could
be limited to the navigationally most suitable substances. Then it may happen
that an arbitrary scent presented at a high concentration (and being conspicuous because of its strong correlation with wind direction) induces a stimulus at a level similar to that of pre-adapted specific chemosignals at very low
concentrations. Search for the species of trace gases actually used by pigeons
for homing may thus go via determination of thresholds of neural responses
(Sect. 12.3.3).
138
7.7.4
The Problem of Adaptation/Habituation
Usually, the olfactory sense reduces its sensitivity to a given odour if it is exposed to this odour over a longer period of time (e.g. Dalton and Wysocki
1996). Such adaptation must not occur with respect to chemosignals used for
navigation, which are permanently present in the air and need to be permanently evaluated in an appropriate manner.
This latter demand, in particular, makes it likely that olfactory signal processing as applied for navigational purposes differs from classical olfaction. It
would not be unique in this respect. Particular airborne chemicals at very low
concentrations can activate specific brain regions and regulate physiological
and psychological functions in humans without being consciously discernible
as odours (e.g. Lorig et al. 1991; Stern and McClintock 1998; Sobel et al. 1999;
Jacob et al. 2001a,b, 2002). Wang et al. (2002) conclude that, while the central
process of conscious perception habituates quite rapidly and completely, the
receptors do not as completely adapt and continue to relay information to the
brain. Salmon migrating upstream are also permanently exposed to the olfactory tracers around them. It appears conceivable, at least, that birds deduce, at
home, some north-wind feeling, east-wind feeling etc. from the wind-associated proportional composition of a set of related airborne compounds without
being consciously aware of them.
Speaking about olfactory navigation means only that the input channel of
the olfactory system is involved, but not necessarily that the birds rely on
smelling of odours in our everyday meaning. Possibly, we should better avoid
speaking of odours in this context at all, because odour is by definition associated with a conscious sensation. Instead, we may more neutrally state that navigating birds rely on atmospheric chemosignals.
7.8
Arguments Against Olfactory Navigation and Replies
Because olfactory navigation has been a matter of considerable controversy, it
seems advisable to devote an extra section to the contra-arguments. More detailed discussions can be found in the Appendix and elsewhere in the literature.
7.8.1
Intuitive Incredibility
When confronted for the first time with the issue that birds might navigate
home over hundreds of kilometres by means of airborne trace gases, almost
everybody responds with disbelief. This immediate response is understandable because intuitively it seems inconceivable that the unstable atmosphere
contains information on positional relationships over such long distances.
There are some publications that seem to provide evidence that olfactory navi-
7.8 Arguments Against Olfactory Navigation and Replies
139
gation is in fact unfeasible (Becker and van Raden 1986; Waldvogel 1987;
Ganzhorn and Paffrath 1995). However, these claims are based on the premise
that birds should be able to deduce positional information from separately
measured concentrations of at least two individual compounds. As it is a priori
unlikely that this premise is appropriate (Sect. 7.6.1), it is also inappropriate to
advance doubts about the feasibility of olfactory navigation with reference to
these publications, as has been done repeatedly (e.g. Schmidt-Koenig 1987,
1991, 2001; Waldvogel 1989; Schmidt-Koenig and Ganzhorn 1991; Wiltschko
1996).We cannot expect that every kind of data extractable from the atmosphere
can be exploited for navigational purposes. It is a particular challenge to find
those relationships that are really relevant. The method of analysis described in
Section 7.6 may not be optimal, but at least it shows that navigationally exploitable structures can basically be found in the atmosphere.
Not only does credibility of a conclusion depend on the power of scientific evidence but also it contains a human factor. It is much more common and popular,
for instance, to speculate on magnetic maps rather than on olfactory maps of
birds, although empirical evidence is clearly against the former (Sect. 6.3.3) and
in favour of the latter (Sects. 7.17.6).The probable reason for this discrepancy is
the fact that everybody knows the geomagnetic field is a useful tool to determine
directions everywhere on earth, whereas the labile atmosphere appears utterly
unsuited as a basis for spatial guidance. Moreover, since magnetism is outside
our conscious sensory feeling and its reception is not yet well understood even
in animals, we view it as a kind of witchcraft which is most suitable to explain
mysterious phenomena. In contrast, due to our subjective familiarity with the
comparatively trivial world of smell, we know by experience that spatial orientation over hundreds of kilometres is impossible within this world. However,
neither is the earths magnetic field so clearly a useful tool for all navigational
purposes, including determination of positions (Sect. 6.3.3), nor can we be certain to be introspectively familiar with all facets of the olfactory world.
Nevertheless, olfactory navigation is often considered impossible and consequently as non-existent. Coming from this conviction, it is necessary to argue
why the experimental data apparently proving its existence are either untrustworthy or explicable in some different way. The next two sections deal with objections assembled in the literature, but I do not repeat all the aspects and arguments that I have already discussed previously in detail (Wallraff 1980b,
1983, 1988a, 1990a, 2001, 2003; see also Papi 1986, 1989, 1991; Able 1996) nor do
I cite all publications again to which the earlier discussions refer.
7.8.2
Potential Non-Specific Side Effects
Most frequent, up till now (e.g. Wiltschko 1996; Walker 1999), has been the objection that olfactory deprivation, in particular by invasive methods, may
cause general neural defects, or defects in another sensory system, thereby disturbing the pigeons orientation behaviour in a way that has nothing to do with
140
olfaction per se and/or with the mechanism of home-finding. In principle, the

objection is reasonable. Moreover, non-olfactory deficits or disturbances
caused by interferences with the olfactory system have in fact been found
(Wenzel and Salzman 1968; Wenzel et al. 1969; Wenzel and Rausch 1977;
Dornfeldt and Bilo 1990). However, the risk of producing uncontrolled effects
has been taken into account in many experimental studies which took care that
differences in treatment between experimental and control pigeons did not
simply concern smelling or non-smelling but were restricted to specific functions excluding possible non-olfactory or non-navigational side effects or, at
least, making alternative explanations extremely far-fetched and implausible
(Sect. 7.17.3). Unfortunately, this fact has usually not been mentioned when
the case of olfactory navigation has been debated with an opponent attitude.
The readers thereby gain the impression that careless conclusions were drawn
from ambiguous experiments. It is emphasized, for instance, that intranasal
anaesthesia has general systemic effects, may affect motivation to home, may
cause defects in brain activities and vestibular functions or might affect
magnetoreception. It remains unmentioned, however, that in decisive homing
experiments both the experimental and the control pigeons were treated with
an anaesthetic in essentially the same way, so that differences in orientation
behaviour could not have been caused by such side effects even if they did actually occur (Sects. 7.2.1 and 7.3.1).
Not each kind of experiment is immune to possible interpretation according
to unintended side effects if it is considered isolated from other experimental
approaches. However, if it fits in the context of a broad assembly of other investigations using different methods, it is unlikely that the results in this one particular case were disparately caused. All the findings described in this chapter
concur quite well with a coherent concept of navigation based on atmospheric
chemosignals. As long as opponents, who doubt this concept as a whole, pick
only at selected experiments without offering alternative explanations of the
entire set of results as a whole (e.g. Schmidt-Koenig 1987, 1991; Waldvogel
1989; Wiltschko 1996; Gould 1998), criticism remains counterproductive. Nevertheless, as emphasized by Wiltschko (1996), lack of an alternative does not
imply evidence that a given conclusion or hypothesis is valid. However, as long
as the conclusion or hypothesis is based on strong experimental support, cannot be falsified and has no competitor, there is no reason to abandon it and to
replace it by nothing.
7.8.3
Inconsistent Results
It has been repeatedly said that some findings indicating the importance of
olfaction in pigeon homing could not be replicated by other researchers (e.g.
Wiltschko 1996; Schmidt-Koenig 2001; Gould 2004). There are, however, hardly
more real inconsistencies than those shown in Fig. 7.37. Even in these 104 releases, olfactorily impeded pigeons were poorer than control birds in initial
7.8 Arguments Against Olfactory Navigation and Replies
141
n Fig. 7.37. Experiments using occlusion of nostrils and subsequent nasal anaesthesia for olfac-
tory deprivation. Second-order means (above) and medians (below) from 26 quartets of releases
at symmetrically distributed sites. Both labellings of the abscissa are valid for both diagrams. G or
X indicates use of the anaesthetic Gingicain or Xylocain, respectively. Filled symbols and solid
lines Control birds; open symbols and dashed lines experimental birds. Above Mean homeward
components (upper bar in quartet 3 refers to G, lower bar to X). Third-order means per group of
quartets are indicated by horizontal lines, overall means by arrows on the right. Below Median
homing speeds, as far as available. Left ordinate Mean of two central values of four medians; right
ordinate (referring to samples marked by >) relative units derived from percentage homed on
the day of release later never. Home lofts in Italy (Tuscany): F Florence, S Sinalunga, M Montefoscoli, A Arnino (Pisa), P Peccioli; in Germany: W Wrzburg, otherwise in Frankfurt am
Main (two lofts: one at ground level in a garden among buildings and trees, one on the roof of a
building); in the USA: Ithaca, New York. Note that (in contrast to Fig. 7.4) each symbol summarizes results from four symmetrical release sites; ranges of variability are therefore reduced
against single releases (sources: quartet #1: Ioal 1983; #2: Meschini 1983; #34: Wiltschko et al.
1986; #56: Benvenuti et al. 1990b; #78: Wallraff 1982 and unpubl.; #913: Wiltschko et al. 1987b;
#1418: Wiltschko and Wiltschko 1989; #1922: Wiltschko et al. 1987d; #2326: Benvenuti and
Brown 1989)
homeward orientation and homing speed, with similar consistency to those in

the experiments described above (Sects. 7.1 and 7.2). Unlike those results, however, reduction of initial homeward orientation did not always lead to a level of
zero. Olfactory impairment in the experiments presented in Fig. 7.37 has been
produced by a sequential combination of occlusion of the nostrils (during transport and waiting at the release site) and nasal anaesthesia (initiated shortly before release). The latter treatment has not been, and partly cannot be, fully standardized; at the time of release the degree of olfactory deprivation was probably
variable (Sect. 7.1.3 and see Appendix; see also the effects of Xylocain and
Gingicain in Fig. 7.37A, quartet #3). More important than the few minutes when
the departing birds are under observation is long-term smelling before release
(Figs. 7.8 and 7.9). During that time, olfactory sensitivity was only reduced by
142
the occluded nares but not eliminated (Fig. 7.6). Thus, the degree of olfactory deprivation of the experimental birds in Fig. 7.37 ranged somewhere between the
control and experimental birds in Fig. 7.8 which differed in treatment before release but which all took off under nasal anaesthesia.
Impaired but not excluded olfaction, even if generated by experimental treatment as standardized as possible, can be expected to cause inconsistent impairment of homeward orientation depending on varying uncontrolled concomitant circumstances. Some of these circumstances are always uncontrolled. We
know neither the environmental olfactory signals actually used by the birds to
determine position nor the way by which reduction of one directional tendency
(homeward) affects the output resulting from variably weighted directional tendencies (cf. Sect. 3.7.1). Also in these experiments with varying initial orientation data, if conducted in clearly unfamiliar areas, homing speeds were consistently lowered in the birds flying under nasal anaesthesia (cf. Fig. 7.5 which
refers to quartets #1922 of Fig. 7.37). Pre-release conditions could only influence initial orientation, while the subsequent homing flight required enduring
olfactory feedback (which returned with ceasing anaesthesia, so that a level of
zero could never be expected).
Several other inconsistencies between findings as well as between findings
and theoretical expectations, as emphasized by Wiltschko (1996), are unsuited
to raising doubts about the importance of airborne odours as navigational
cues. Rather, they result from imperfect planning and execution of experiments which often did not take all the factors into account that contribute to
the homing behaviour of pigeons. Over the last decades, we have been forced to
learn by experience that we must proceed much more carefully than naively
expected in the earlier years of research on pigeon homing. We have learned
(1) that it is in most cases important to use release sites in central symmetry
around home and to evaluate the obtained results as a whole (Sects. 3.1.1 and
3.7.1), (2) that the directions taken by control pigeons alone, under neglect of
the homeward direction, are an insufficient reference to assess the behaviour
of manipulated birds (Sect. 7.9.1; see Fig. 4 in Wallraff 2001), (3) that we should
not underestimate the possible role of visual landscape features as home-guiding and as distracting factors (Chap. 8), (4) that non-familiarity with the surrounding area is not guaranteed alone by the fact that the pigeons had not been
released before at the exact current release site (Sect. 8.1.1), (5) that complete
anosmia cannot be achieved by occlusion of the nostrils and is not reliably
achieved under all circumstances by intranasal anaesthesia (Sect. 7.1, Fig. 7.6),
and (6) that smelling during a longer period before release is more important
than smelling during the first minutes of flight (Figs. 7.8 and 7.9).
The experimental data obtained over the years are quite heterogeneous as
regards the degree of reliance under all these aspects. Thus, discrepancies between findings or between findings and predictions should not be misused in
declaring olfactory navigation a doubtful matter if imperfect experimental
methods were, or could have been, involved (cf. Able 1996). For some controversies over results obtained with deflector lofts, see Section 7.5.1.
143
7.9
Olfactory navigation,as described above,does not fit into preformed concepts of
animal homing or other forms of goal-finding, although the sense of smell is
widely used for purposes of spatial orientation. Usually, however, it is orientation towards the source of a particular scent by moving against the wind (e.g. towards a female moth, towards a foraging site, towards a nesting burrow etc.) or
against a water current (e.g.salmon swimming upstream to their home river),by
determining the gradient-upwards direction during search movements or by
following an odour trail fixed to the ground substrate. Pigeon navigation, showing homeward orientation from the beginning with varying headwinds, tailwinds or sidewinds over unfamiliar terrain, is different. Goal-oriented navigation in its narrower meaning, on the other hand, is usually associated with ideas
of coordinates forming a wide-ranging grid. Volatile substances dispersed in the
unstable atmosphere appear to be unsuitable as spatial coordinates. Thus, the
idea of olfactory navigation would never have been created as a theoretical concept of avian homing; it was an unexpected outcome of experimental research.
7.9.1
Experimental Evidence of Olfactory Navigation
A variety of different experimental approaches have revealed two preconditions
that must be met in order to observe homeward-oriented flights of pigeons displaced into unfamiliar distant areas: (1) during their preceding long-term stay at
the home site, the birds must have been exposed to natural air and wind conditions (Sect. 7.5); and (2) before, during and/or after release, the birds must have
olfactory access to natural environmental air (Sects. 7.17.3). There are no indications that other environmental cues (magnetic, astronomic, infrasonic etc.)
might be additionally involved in position determination (Sects. 4.2 and 6.3.3).
Nevertheless, the sun and the geomagnetic field are also involved in the homing
process, though only as directional compass references (Sects. 5.4 and 6.3.2).
Thus, the experiments strongly suggest that atmospheric chemosignals are
necessary as well as sufficient as indicators of position to enable homing over
considerably long distances in unfamiliar territories. This inference holds independently of any knowledge about the mechanism enabling olfactory navigation.
It should be emphasized here that olfactory deprivation or misguidance of
any kind acts only on homeward orientation and not on the other components
that determine initial orientation patterns (Sect. 3.7.1). Thus, when comparing
control and experimental pigeons only with each other, considerable similarities are often more obvious than the differences, because both groups have
PCDs and deflections by landscape features in common (Fig. 8.6 in Sect. 8.2;
Appendix). Evidence of an effect on homeward orientation is attainable only if
a comparison includes the homeward direction as a reference.
144
7.9.2
How Does Olfactory Navigation Operate?
This question cannot be answered as yet. Only a hypothetical approach has
been made so far (Sect. 7.6.1) and atmospheric conditions have been found
that are consistent with this approach (Sects. 7.6.2 and 7.6.3). The hypothesis
includes three basic components: (1) a position-independent directional reference, (2) winds coming from various directions as observed at home, and (3)
fairly wide-ranging gradients of ratios among atmospheric trace compounds.
There is no problem to combine the first with the second component: birds
may easily determine wind direction as an angle to the sun (Sect. 5.4) and/or to
the geomagnetic field (Sect. 6.3.2). That pigeons actually refer to wind directions has been shown by experiments deflecting or reversing winds in an aviary in which the birds lived at home (Sect. 7.5.1). The proposition that pigeons
refer to ratios among quantities of airborne compounds is hypothetical. It is
not hypothetical, however, that pigeons refer to chemosignals (Sects. 7.17.3)
and that distributions of ratios among some atmospheric trace gases include
differently oriented spatial gradients (Sect. 7.6.2). Combining these findings
results in a model as illustrated in Fig. 7.26: birds associate, over several weeks
at home, a specific ratio pattern (= composed odour quality) with a specific
wind direction. When a bird is subsequently released in an unfamiliar environment, it flies in that compass direction towards which the wind blew while the
relevant olfactory signals at home were most closely related to the signals it
had received away from home over some time before release. In the course of
its further flight, it updates the incoming signals and corrects its route accordingly.
Owing to the ambiguity of expected ratio changes caused by a change in
wind direction, on the one hand, and a change of position, on the other hand
(Sect. 7.6.1), this brief description of the hypothetical mechanism is probably
an oversimplification. Nevertheless, it may include the core of the system
whose full understanding requires more detailed investigations of the atmosphere particularly with respect to the relationship of positional direction
from home versus wind direction at home (see Wallraff 1989c for a closer inspection of this problem).
7.9.3
Do Pigeons Have an Olfactory Map?
Do not ask whether they have a cognitive map, but how they find their way
about. In this title of an article by Mackintosh (2002) we could replace cognitive map by gradient map, olfactory map etc. Actually, however, it is hardly
possible to avoid the term map while writing about somewhat more complex
mechanisms of goal finding, because the term is so commonly used in the literature and because it is so convenient. It depicts ostensively and briefly that we
are dealing with configurations extending into the two-dimensional space.
145
However, even if we can show that an animal navigates by referring to such spatial configurations, it does not necessarily mean that their spatial structure is
represented in the animals brain in some analogy to our maps on paper. Nevertheless, to some degree, such an analogy may exist in the case of a topographical
map or a cognitive map which refers to landmarks individually learned during
excursions in the two-dimensional space (Sects. 8.1, 8.3, 9.1 and 9.5). Theoretically,also a gradient mapor grid mapmay be conceivable as a representation of
spatial structures, provided that the animal could measure and register directions of increasing and decreasing scalar quantities of some parameters while
moving around its home site and thus could associate the recorded inputs with
the explored spatial dimensions. It could then extrapolate the scanned gradients
into unfamiliar farther distant areas (cf. Fig. 4.1C,D).
It is unlikely that pigeons have, in this sense, an olfactory map, an olfactory
gradient map, a navigational map or something similar. In order to develop
their homing mechanism, pigeons need not fly around (Sect. 3.1.3) and the
mechanism can be manipulated by manipulating wind directions while the
birds are confined in an aviary (Sect. 7.5.1). How should such position-fixed
pigeons create a representation of far-reaching spatial structures? In fact, they
need not have any knowledge of such structures in order to exploit them for
navigational purposes. According to the hypothesis outlined in the last section, a bird merely needs to correlate, at home, varying olfactory sensations
with varying wind directions and to fly, at a distant site, in that compass direction toward which the wind usually blew while the sensation was most similar
to the current one. In doing so, the pigeon need not have a map-like representation in its brain and need not know anything about gradients (this holds true
even if the description given here is too simplistic).
If pigeons do not have a navigational map, Kramers map-and-compass concept (Sect. 5.2) must also be abandoned. Deflections observed in clock-shift
experiments (Sect. 5.1) do not prove operation of two independent devices in
subsequent steps, as assumed over 50 years, but they reveal some correlative
linkage between positional and directional indicators. The birds need not
know that the positional indicator is what it is, i.e. that it has anything to do
with wide-ranging spatial structures (cf. Wallraff 2004).
All researchers in the field, including myself, are accustomed to using the
term map not only when referring to familiar topographical features, but in all
cases concerning the determination of the homeward course from a distant
site, and hence some sort of relative position determination, while direct sensory contact with the goal is lacking. Therefore, and because the term is brief
and convenient, I am reluctant to avoid it henceforth completely. I still tend to
use it in such cases in which a navigation system operates with environmental
structures that we humans can in some way graphically reproduce on a map
(showing landmarks, a grid of isolines etc.). We should be aware, however, that
then the term only circumscribes the external spatial structures used by animals and does not indicate their internal mode of usage. The animals behave
as if they had a map. I would not argue against erasing the term in the context
146
of olfactory navigation if a handy alternative terminology would be presented.

Presently in this book, I continue to use the term map in various contexts, although I doubt that it is always appropriate in a strict sense.
A perfect gradient map, i.e. a coordinate system based on monotonic gradients, theoretically provides information not only on the directional relationship
between two locations but also on the distance between them. However, to know
the distance towards home is not indispensable for home-finding. Flying in the
right direction until reaching a familiar target area would suffice. So we may ask
whether the pigeons map-like system involves distance as a parameter.
If the system works as hypothesized and with atmospheric data similar to
those measured so far, the pigeons should feel to be away from home with a
distinctness that depends on the degree of divergence of the currently observed olfactory spectrum from the spectrum the birds would expect at home.
Within some range around home, differences from home ratios should usually
increase with increasing distance. If Fig. 7.29 is representative, however, the
correlation is not very strong and may fade away or may be even reversed with
further increasing distances (see small diagrams in Fig. 7.29; see also Figs. 7.35
and 7.36), so that finally homeward orientation breaks down (Sect. 7.6.4).
Thus, the birds would not be distinctly informed on their distance from home,
but their as-if-map might imply, within a limited range, some rough correlation with distance. If we concede them a map at all, it would be a sort of radial
map (Gould 1998) approximately indicating the radius along which an animal
is displaced, but not including a reliable measure of distance.
In addition to these considerations, I would not exclude the possibility that
experienced pigeons, in the course of many homing flights from various directions and distances, might develop, beyond their as-if-map, an olfactory map
that actually includes a representation of wide-ranging olfactory signals as explored during those flights. This representation might have either the form of a
topographical map of familiar olfactory sensations or the form of a real gradient map based on recognition of signals gradually changing differently in different directions or the form of a hybrid map including both kinds of information (Sect. 10.2.1). As inexperienced pigeons can orientate homeward without
any exploration of surrounding areas, however, a potential map of that kind
could only be a secondary perfectioning and not the primary basis of
home-finding over unfamiliar territories.
7.9.4
Unsolved Problems Remaining Challenges
Olfactory navigation is far from being understood. We do not know any one of
the proposed volatile atmospheric compounds actually used by pigeons for
home-finding and, consequently, do not know their spatiotemporal distribution in the airspace. Also, we do not know the mechanism of olfactory signal
processing that is involved in home-finding. We do not even know whether it is
true that the mechanism makes use of atmospheric ratio gradients, as specula-
147
tively hypothesized above. Deciphering the pigeons navigation system remains a challenge. However, now there is, at least, little doubt about the category of environmental signals that we must observe and analyse in order to
find a solution to the birds mysterious homing abilities (Sect. 12.3.3).
The Role of the Visual Landscape
It would be surprising if homing pigeons during their exercise flights around

the loft and during their homing flights did not take any note of the manifold
visual features of the landscape such as mountains, hills, lakes, forests, rivers,
villages and towns. Also, it seems very likely that pigeons, like many other animals including humans, make active use of such features for orientational purposes once they are familiarized with the spatial relations between them. Thus,
we must a priori consider the possibility that in a familiar area, olfactory signals might not be the only sources of positional information.
8.1
The Landscape as a Home-Guiding Factor
8.1.1
Non-Olfactory Homing
When the same pigeons, in the course of a number of releases at varying sites,
are released at a given site for a second or third time, they usually fly off in directions similar to those flown when they started for the first time at this particular site. Site-specific deviations from the homeward direction (release-site
biases; Sects. 3.1.1, 3.2.3 and 3.7.1) remain fairly stable and homing performances are only slightly improved (Wallraff 1959a; Keeton 1973). Thus, possible familiarity with the landscape does not have an immediately obvious effect.
Other factors determining the birds behaviour are still at work (Sect. 3.7.1)
and even the revisited landscape itself might stimulate the birds to repeat the
remembered route.
Differences between first-time visits and repeated visits of a given site become
apparent, however, when the birds are deprived of olfactory access to ambient
air. Such pigeons fail to orient homeward when they had not before been released at that site or in its vicinity (Sects. 7.1 and 7.9.1), whereas they are fairly
well homeward-oriented at sites where the visual surroundings are potentially
familiar to them from earlier homing flights (e.g. Papi et al. 1980b; Papi 1986;
Benvenuti et al. 1992b; Schmid and Schlund 1993). The experiments shown in
Fig. 8.1 aimed to investigate this aspect more systematically by using controlled
patterns of familiarization. In a training phase, two groups of pigeons were released at a number of corresponding sites within different areas. For testing,
both groups were released simultaneously at novel sites within these two areas,
150
8 The Role of the Visual Landscape
n Fig. 8.1. A In a preparatory training phase lasting more than 2 months, two groups of hitherto
naive pigeons, distinguished by triangles and squares, were released at various sites within different, but centrally overlapping parabolic areas; at sites indicated by circles both groups were released together. The home loft (centre) was at Andechs, about 30 km southwest of Munich. After
training, both groups were released together at the two test sites TSW and TNE, 54 and 53 km from
home. Circles around these sites indicate the maximum distance of bearing observation (2 km)
and minimum distance to training sites (10 km). The topographic appearance of the test areas
was very different. In the test releases, four groups of pigeons were used with different combinations of F+ and F (= familiar and unfamiliar with the area) and of O+ and O (= with and without olfactory access to environmental air; O produced by air filtration before release and nasal
anaesthesia upon release; Sect. 7.2.1). In B and C, results obtained at the two release sites are
combined. B Mean homeward components of the vanishing bearings (*** P<0.001, n.s. not significant). C Cumulative percentage curves of homing performance. Homeward orientation requires either F+ or O+ or both. Note that the last pre-test release, at the neutral site 25, did not
reveal noteworthy differences between the two groups in initial orientation and homing. (Modified from Wallraff and Neumann 1989)
8.1 The Landscape as a Home-Guiding Factor
151
one group of birds with and another group without olfactory access to the local
air. The results make it clear that non-olfactory homing over 50 km can occur,
but only within areas that the birds could explore while flying there previously
(F+O versus FO in Fig. 8.1B, C). As long as they were able to smell natural environmental air, no effect of familiarization was observed (F+O+ versus FO+).
At least in these particular experiments, olfaction-based homing appeared
somewhat superior to non-olfactory homing (FO+ versus F+O), so that the
latter was completely masked when both sources of information were available
(F+O+). These experiments suggest, but do not prove, that homeward orientation without olfactory signals is achieved by referring to the familiar visual landscape. As the pigeons had not yet been released at the test sites themselves, but
only in a wider area around them, they would then have used a fairly wide-ranging image of the landscape.
Figure 8.2 gives additional hints supporting this assumption. The birds in
these experiments, conducted at the same two sites, were less extensively and
less differently trained (in the neighbourhood, only two releases 10 km away
from the test site; see sites 13 and 14 in Fig. 8.1A), and they were all released under intranasal anaesthesia. Thus, the O+ pigeons were equally impaired in
olfaction during flight and equally stressed or non-stressed by this treatment as
the O pigeons. Nevertheless, the homeward components obtained after ca. 2 or
3 min or more of flight, when the birds vanished from sight (black columns in
Fig. 8.2),were similar to those in Fig. 8.1.However,Fig. 8.2 gives an additional aspect. Within the first 40 s of flight, the F+O birds, which were disabled to collect
olfactory information, did not yet show any homeward orientation, whereas the
FO+ birds, depending solely on previously received chemosignals, were fairly
n Fig. 8.2. Mean homeward components of initial bearings at the same two test sites TSW and
TNE as shown in Fig. 8.1. Pigeon groups equivalent to those in that figure, but preceding training
pattern somewhat different and not only O but also O+ pigeons released under nasal anaesthesia (before release O in filtered, O+ in unfiltered air; see Wallraff et al. 1993). Bearings were observed 20 and 40 s after release and at vanishing from sight. (Wallraff 2001)
152
well homeward-oriented from the beginning. This difference is plausible on the

assumption that the F+O pigeons required some time to gain height and survey over the landscape before they could draw conclusions about their position
(cf. Guilford et al. 2004), whereas the FO+ pigeons had already drawn this conclusion prior to release and could not profit from looking around in an unfamiliar area. (The intermediate behaviour of the F+O+ birds suggests that even with
olfactory information the pigeons take the familiar visual landscape into account and try to bring it into relation with this information before they decide on
a course to fly. The topographical characters of the two areas of release were very
different, so that the pigeons might have immediately recognized whether or not
they could have been in the now visible area previously.)
8.1.2
Downgrading of Visual Signals
Other approaches aimed to exclude or reduce visual contact with landmarks by
means of frosted contact lenses attached to the pigeons eyes. When initial bearings of birds wearing such lenses were pooled with respect to their deviation
from the mean bearing of control pigeons with unimpaired vision,no worsening
was apparent even in very experienced birds released only 15 km from the loft,
where the landscape should have been familiar (Schlichte and Schmidt-Koenig
1971; Schmidt-Koenig and Schlichte 1972). As the reported data do not include
the direction towards home, however, they do not reveal to what degree homeward orientation or other directional tendencies (Sect. 3.7.1) made the results
similar (cf. Streng and Wallraff 1992). The only published findings achieved with
this method that include home as a reference and allow us to compare visually
impaired and unimpaired birds under otherwise equal conditions are shown in
Fig. 8.3. The pigeons with diminished image vision generally shortened their
distance to the loft, but compared with the controls, their routes were much less
clearly homeward-oriented. It appears possible that the tracks of the experimental birds reflect the level of performance achievable by pure olfactory navigation
over the short distance used (Sect. 7.4), but it is also possible that they simply reflect general behavioural disturbance.
In similar experiments using pigeons of another loft, two birds with frosted
lenses reached the close vicinity of this loft and one of them circled at a distance of 0.52 km around it (Schmidt-Koenig and Walcott 1978). It remains
undecided whether apparent recognition of the familiar home locality in this
one case indicates a very accurate non-visual navigation system or recognition
of local odours [independently of the assumed gradient system (Sect. 7.6),
which cannot be expected to operate down to such short distances] or some remaining capability of identifying visual contours of the landscape (perhaps
the most likely explanation; see p. 216 in Streng and Wallraff 1992).
Braithwaite and Guilford (1991), Braithwaite (1993), Braithwaite and Newman
(1994), Burt et al. (1997) and Biro et al. (2003) restricted variation of visual conditions to the time before flying. For a period of 5 min prior to release, each pi-
153
n Fig.
8.3. Flight routes of pigeons released under sunny conditions 15 km

north of their loft in Gttingen, Germany,
and tracked by radiotelemetry. Solid lines
refer to birds wearing frosted lenses,
dashed lines to birds wearing clear lenses.
Circles indicate determined positions of
birds in flight; asterisks indicate locations
where birds remained sitting. (Modified
from Schmidt-Koenig and Walcott 1978)
geon was placed in a box with either clear or opaque sides, allowing or preventing a view of the surrounding landscape. Birds that were allowed to preview the
landscape returned significantly faster from sites in a range of 15 km from
home, provided that they had previously been released at that particular site.
Correspondingly, the first parts of their tracks were more directly homewardoriented (Biro et al. 2002). Thus, these pigeons obviously did consider the visual
landscape and profited from its familiarity. However, as the experiments were
conducted in the very close vicinity of the loft, they have little relevance to the
more intriguing homing abilities over longer distances in less familiar surroundings.
8.1.3
Homing with Olfaction and Vision Impaired
In order to test whether non-olfactory homing (Sect. 8.1.1) is based on visual inputs, it would be necessary to combine visual and olfactory deprivation. If it
could be shown that anosmic pigeons in a familiar area are homeward-oriented
even if they are prevented from receiving potentially relevant visual signals, it
would be necessary to propose that a third sensory modality is involved. Clearly
no indication of such a necessity has been found in experiments combining
transitory exclusion of image vision (by temporarily fixed translucent-paper
spectacles) with transitory anosmia (by intranasal anaesthesia after transporta-
154
tion in filtered air), which were conducted, without intense successive-release

training, at six familiar sites 2530 km from the loft (Streng and Wallraff 1992).
Initial orientation as well as homing performance were found at decreasing levels in the following order: nothing impaired, only olfaction impaired, only vision
impaired, both senses impaired. The pigeons with two-fold impairment did not
show any trace of initial homeward orientation and their homing speeds were by
far the lowest. Thus, the findings support the hypothesis that non-olfactory
homing from familiar sites is visual homing. Nevertheless, I consider the results
indicative, but not definitely conclusive. Even the birds deprived only of image
vision were not significantly homeward-oriented at vanishing (but homed significantly faster than the double-treated birds).Unlike anosmia,image blindness
disturbs almost all kinds of elementary behavioural activities, including walking and flying, so that specific deficits in orientation can hardly be separated
from non-specific behavioural deficits. Even though additional anosmia decreased performances further, thus indicating that with olfaction they were not
yet at the deepest possible point, it cannot be excluded that the nasal anaesthesia
merely strengthened the anyhow stressful conditions acting on the pigeons in a
non-specific way.
A more promising approach by Gagliardo et al. (2001b) avoided direct interference with vision itself. It is one of the thus far rare experimental series in
which the birds were successfully tested in a round cage (diameter 1.8 m) before they began flight (Sect. 2.5). This method allowed visual shielding of the
landscape. Normal and anosmic pigeons were tested at three previously familiarized sites, one time with the surrounding landscape screened by means of
large plastic sheets approx. 2.5 m away from the cage and one time without
such a screen. Homeward relatedness of the directions in which the birds exited from the circular arena (Fig. 8.4) was basically similar to that shown in
Fig. 8.1B. The fact that even the O+ birds were only weakly home-oriented behind the screen should possibly be seen against the background that distances
n Fig. 8.4. Mean homeward components of directions
towards which pigeons exited from a circular arena in

a familiar surrounding which was either visible to the
birds or shielded (*** P<0.001, * P<0.05). The O
birds were made anosmic by means of zinc sulphate
(Sect. 7.1.2). (Data from Gagliardo et al. 2001b)
155
from home were very short (718 km), so that, even in Italy, olfactory navigation might have operated suboptimally (Sect. 7.4). These findings support the
hypothesis that non-olfactory homeward orientation at familiar sites is based
on observation of the visual landscape.
8.1.4
Interrelations Between Landscape and Sun Compass
Since it proved problematic to obstruct the pigeons view of the landscape during flight, it appeared advisable to look for more indirect methods to reveal the
possible role of visual cues to home-finding over familiar terrain. Experiments
with clock-shifted pigeons (Sect. 5.1) may be considered suitable for this purpose, because they tend to deflect initial bearings away from the route towards
home as guided by the landscape. As such deflections were regularly observed
not only at unfamiliar, but also at familiar sites, they were considered to indicate that visual landmarks are not used for orientation, because the birds
readily fly away from the visible structures that would have guided them home
(e.g. Keeton 1974a; Schmidt-Koenig 1979; Schmidt-Koenig and Ganzhorn
1991; Wiltschko 1991, 1996; Wiltschko and Wiltschko 1998). When inspected
more closely, however, just the clock-shift experiments strongly suggest that
pigeons do make use of familiar topographical features for home-finding.
It has long been known that angular deflections of initial bearings caused by
clock shift, although consistently occurring with the predicted sign, are often
smaller than predicted on a pure sun-compass basis. In the summaries of
Schmidt-Koenig (1979; Schmidt-Koenig et al. 1991), the mean deflection of 383
pigeons with their clocks shifted 6 h forward is only 70, whereas the mean difference between expected and actual sun azimuth was in the range of approximately 110120 (see Wallraff 1974a; Wiltschko et al. 1994). Moreover, mean angular dispersion per release was greater in birds with shifted than in those with
non-shifted clocks, indicating that the clock-shifted birds felt exposed to less
clear-cut conditions than the controls. Such effects can be expected if the birds
were influenced by both the pattern of familiar landmarks, indicating one direction as appropriate, and other factors, including the sun, guiding them in another direction. The conflicting condition to which a sample of individually flying birds is exposed may then result in a mean bearing somewhere between full
and no deflection and in an increased angular dispersion between the birds (see
Wallraff 1991a). Such a conflict does not occur in unfamiliar areas, where the
landscape contains neither positional nor directional information. It is very
likely that in many of the experiments conducted by Schmidt-Koenig (1958,
1961), Wiltschko et al. (1994) and Wiltschko and Wiltschko (2001), the old experienced pigeons they used were more or less familiar with the area in which they
were released. Moreover, from previous releases under clock shift, many birds
may have learned to distrust the shifted sun compass and tended instead to follow, wherever possible, the familiar landscape directly and to pay more attention
than usual to the non-shifted magnetic compass, which remains in directional
156
agreement with the landscape. Against this background, the effect of attached
magnets on the degree of deflections caused by clock shift (Fig. 6.2) may become
understandable (see Appendix).
These are ex post guesses being in general agreement with the experimental
conditions described by the authors. In other experiments, the relevant conditions were a priori controlled according to the following rationales. In an unfamiliar environment, pigeons are thought to deduce from airborne chemosignals in what angle to the sun they should fly (Sects. 7.9.2 and 7.9.3). If the
sun compass is shifted by clock shift, they follow the angular shift without hesitation, not realizing that something is wrong (Fig. 8.5A; see also Figs. 5.1 and
5.2). A familiar landscape, however, not only providing information on the
birds own position but also presenting a spatial structure over a large surrounding area, would by itself indicate the right way. During familiarization,
the sun may have been observed together with the landscape, so that a certain
n Fig. 8.5. Angular deviations of mean single-release vectors of vanishing bearings of pigeons
whose circadian clock was shifted 6 h forward, from corresponding vectors of non-shifted controls. The deviation (abscissa) is given as a percentage of the varying angle between actually observed sun azimuth and expected sun azimuth according to the phase-shifted time scale of the
experimental birds (depending on date and site of release, this angle azimuth varied between 88
and 144; Sect. 5.1). A is from normal pigeons in a largely unfamiliar area (F), B from normal pigeons in a familiar area (F+) and C from pigeons deprived of olfactory access to natural air (O)
released in a familiar area (F+). Symbols refer to different series of experiments whose conditions
in detail were not standardized (circles in A and B+C from different series). (Wallraff et al. 1999;
see therein for references to data sources)
8.2 The Landscape as a Disturbing Factor
157
compass course was associated with the view and the alignment of a certain
landscape. For a bird with its circadian clock shifted by 6 h, the sun is in a
wrong position with respect to the landscape. Which of the two sources of information should the bird trust? Its confusion should be particularly strong if
the determination of position is based exclusively on visual landmarks, i.e. if
olfactory signals are excluded. In fact, under this condition, the effect of a 6-h
clock shift is quite variable and inconsistent (Fig. 8.5C). It is, however, not abolished (otherwise the vectors would be symmetrically distributed around the
control mean). The fact that a clock-shift effect does exist without olfaction at
all suggests that pigeons presumably using solely visual cues do not pilot home
by looking at the landscape alone but by looking at the sun as well. With both
olfactory and familiar visual signals available, reduced deflections, but not so
widely scattered bearings, have been observed (Fig. 8.5B).
Variable effects of clock shifts similar to those seen in olfactorily deprived
birds in familiar areas were observed in normal pigeons released only 13 km
away from the loft (Graue 1963; Keeton 1974a; Schmidt-Koenig 1979; Holland
et al. 2000). These birds were in a comparable situation. Odours could not be
used in the close proximity of the loft and the sun did not fit in the expected
way with the landscape. Splits between following the sun and following the
landscape were sometimes observed among bearings of individual pigeons at
the same site and sometimes among the majority of birds at one site and another. Such splits can be interpreted to reflect the dichotomous situation in
which sun and landscape, usually used in accordance with each other, are at
variance (see Wallraff 1991a, Fig. 6DF). The birds have no choice but to neglect one kind of signal or to make a compromise. Variable effects of clock
shifts on complete homing routes were also observed from more distant release sites in familiar areas (Papi et al. 1991; Bonadonna et al. 2000). They suggest that the landscape tends to dominate over the sun particularly at sites or in
areas where conspicuous features such as coastlines or mountain ridges act as
guidelines and/or barriers. It is likely that varying local topographical features
also contributed to the great variability of the vectors shown in Fig. 8.5C.
8.2
The Landscape as a Disturbing Factor
Landscape features not only may be helpful for homing when familiar, but also
may distract the pigeons from the course they would have chosen in a homogeneous environment. This negative influence of the landscape has already been
discussed and illustrated in another context (Sect. 3.4.3).
Although it is mostly impossible to verify topographical influences at a particular site with certainty, some site-specific patterns of initial bearings leave
hardly any doubt that they are substantially determined by such influences.
Figure 8.6 gives an example in which distributions of bearings show three distinct peaks. In the direction of each of these peaks is a village 12 km distant.
As usual, villages more than 3 km distant have no force of attraction within the
158
2-km radius of observation (cf. Fig. 3.21). The three peaks dominate both diagrams, but which of them is highest depends on whether or not the birds had
olfactory access to environmental air.
This example suggests three interacting tendencies: (1) a tendency to fly towards attractive features of the landscape; (2) a tendency to fly northwest, which
is the general PCD of pigeons of this loft (e.g. Figs. 3.4 and 7.2A) and here the
mean bearing of birds knowing nothing about home (Fig. 8.6C); and (3) a tendency to fly homeward in a southern direction, which is evident only in birds
that are able to smell the local air (Fig. 8.6B).
In this example, it is particularly obvious that a tendency to fly homewards is
only a modifier of stronger directional tendencies that characterize initial orientation at a given site, but have nothing to do with the home-finding mechanism
(Sect. 3.7.1). It seems very likely that trivial attraction and distraction by topographical peculiarities are common components of pigeon homing behaviour.
Thus, site-specific bearing patterns, and mean directions calculated from them,
n Fig. 8.6. Initial orientation of first-flight pi-
geons in 18 releases at a site 30 km north of the

home loft near Wrzburg. Distribution of vanishing bearings is smoothed by calculation of
running means. Filled arrows give directions of
resulting mean vectors. Symbols in A indicate
the approximate angular and distance ranges of
villages within a 5-km radius around the release
site. B Control pigeons allowed to smell natural
air before and after release. C Experimental pigeons breathing filtered air before release and
then departing under nasal anaesthesia. Distributions in B and C are significantly different
with P<0.0001. (Modified from Kiepenheuer et
al. 1993)
159
must not be taken as undisturbed indicators of navigational decisions based

solely on map information.
Figure 8.6 additionally shows that site-specific orientation is not exhaustively
described by a mean vector alone, which itself does not point towards a village.
The whole angular pattern of bearings is typical for this particular site (for more
such patterns, see Fig. 8 in Wallraff 1959b). Other patterns co-determined by the
local landscape are typical for other sites (cf. Kiepenheuer et al. 1993; see also
Fig. 1 in von Hnerbein et al. 2000). Temporal fluctuations of navigationally preferred directions and effects of clock shifts can be modulated by such discontinuous channelling of bearings towards attracting (or away from repelling) features of the surrounding landscape. Angular differences between navigationally
intended directions may thereby become over- or underexpressed in the resulting mean vectors (cf. Fig. 7 in Wallraff 1994b).
A mixture of the two kinds of landscape influence is also conceivable but so far
purely theoretical. Pigeons might respond to features they actually see in accordance with their similarities to other features they have memorized. They might
deduce from them a seemingly home-guiding direction. The effect would also
be distraction from a course deduced solely from true navigational cues.
8.3
More important than the deflections caused by topographical features is the
helpful role of a familiar landscape for way-finding. Many arthropods, vertebrates and other animals make use of visual landmarks to find their way about
(e.g. Papi 1992a; Wehner et al. 1996), so why should homing pigeons be an exception? Available experimental evidence is clearly in favour of the most likely
and most trivial hypothesis that non-olfactory homing from familiar sites is
based on visual features of the landscape. Also, the role of the hippocampal formation in pigeon homing appears largely equivalent to its otherwise known
role in vertebrate landmark orientation (Sects. 9.1 and 9.5).
There is a difference between many birds and many other animals which
makes the investigation of the role of landmarks in avian homing more difficult. Pigeons are able to home by using a second kind of location-bound signals which provide positional information also in unfamiliar areas where the
birds have never been before. These signals remain operational when an area
has become familiar to the birds. Also distracting factors (Sects. 3.7.1 and 8.2)
are not, or need not be, eliminated by familiarization. Thus, from the observation that pigeons released at a given site behave quite similar, independently of
whether or not they had previously been released at that site, it cannot be concluded that the birds do not pay attention to the surrounding landscape. In
many cases, landscape orientation may be redundant; in some cases, when olfactory signals are weak or ambiguous, it may be helpful; and in cases when
such signals are not available at all (whether naturally or experimentally),
homing may still be possible by means of landmarks alone. Pure landmark ori-
160
entation may also be applied if neither of the two compasses operate adequately, the sun compass because of overcast and the magnetic compass because of attached magnets or coils (Sect. 6.1.1). Conflicting situations, and
hence unclear, inconsistent or compromising results, may occur in experiments implying clock shifts, deflector aviaries or olfactory site simulations, if
they are conducted in a familiar area. Retrospectively, we must state that many
results described in the literature must be viewed with some reservation because, at least potentially, they might have been co-determined by influences
coming from the familiar landscape. Such influences are likely to occur when
pigeons experienced in homing from many sites around are tested at short distances from home. It is often difficult or impossible to retrospectively estimate
whether, or to what degree, one or another published result might have been affected by such interfering conditions.
If we accept that non-olfactory homing is based on visual cues, the findings
shown in Fig. 8.1 suggest that landscape orientation does not, or not only and
always, mean that the birds follow a chain of individually learned small-scale
landmarks such as houses, villages, forests, rivers, roads etc., connecting the release site with the home site. However, an alternative familiar-area map model
so far lacks clear contours (cf. Holland 2003). It is unknown whether, also in
these dimensions, the pigeons apply a geometric multiple-bearings system
(Kamil and Cheng 2001) or something similar, referring to large individual
landmarks such as mountains, lakes, rivers or towns, by which they triangulate
or multi-angulate home, or whether, perhaps more likely, the birds make use
of a large-scale aerial panoramic view over a wide area without focusing particularly on single landmarks. In either case, the system would allow evaluation of a landscape from viewpoints with varying parallax, including points at
which the birds had not before been. Otherwise, previous experience not with
the current release site itself but after releases at least 10 km away (Figs. 8.1 and
8.2) would not be sufficient to make olfactory inputs largely superfluous. An
assumed visual topographical map of an area would not provide more accurate positional information than the assumed olfactory gradient map, so that
familiarity with a site or an area does not express itself in remarkably improved homeward orientation. The wide bi-monocular visual field of a pigeon,
which covers slightly less than 360 in the horizontal plane (Donovan 1978),
should enable a survey over a wide aerial panorama of the landscape below.
Referring to previously experienced large-scale characteristics of an area
rather than to small-scale landmarks would facilitate vision-based homing
from sites that themselves have not yet been visited. However, this kind of
sketchy topographical mapping within a moderately familiarized area might
provide merely a rough idea of the direction towards home, and may not be
better than the estimate deduced from olfactory signals. A pigeons visual view
of the world differs considerably from a humans view (Waldvogel 1990). As in
the case of olfaction (Sect. 7.7), we should not extrapolate too narrowly from
our own sensory capabilities and restraints (Sect. 12.3.2).
The Neural Bases of Pigeon Homing
So far, we have seen the pigeon as an entity that interacts with the environment.
However, an organism is internally structured, and so we may ask which parts of
the neural machinery interact with which parts of the environment and in what
way. Allocating certain components of the homing process to certain regions of
the brain not only can shed some light on the organization of the biological
mechanism, but also may help to distinguish and elucidate the specific roles of
different external signals.Considerable efforts in this field have been made in recent years mainly by the experimental approaches of Verner Bingman, Anna
Gagliardo and colleagues (for review see Bingman et al. 1995, 1998b; Bingman
1998; Bingman and Able 2002).
The basic methodological tools used in these studies are lesions to particular
parts of the brain of pigeons at varying stages of their ontogenetic development
and/or after varying learning experiences. Orientation performances of birds
with and without such lesions are then compared under varying conditions (familiarity or non-familiarity of a release site, directional training in a cage, clock
shift etc.). Thereby, specific roles in navigation processes could be attributed to
particular regions of the telencephalon. Most clearly separated appear cerebral
representations primarily used in visual landscape orientation within a familiar
area and others used in olfaction-based homing from unfamiliar areas.
9.1
The Hippocampus and the Familiar-Landscape Map
The mammalian hippocampus is known to be a crucial centre in the neural organization of spatial cognition, in particular of orientation using familiar
landmarks represented in a so-called cognitive map (e.g. OKeefe and Nadel
1978; Gallistel 1990; Jacobs and Schenk 2003). An apparently homologous region in the avian brain (Fig. 9.1) turned out to fulfil comparable functions (cf.
Colombo and Broadbent 2000), as has been shown, in a first step, by the following experiment (Bingman et al. 1987). Having performed eight training releases from each of two sites 40 and 55 km away from home in opposite directions, two-thirds of a group of pigeons underwent ablation of the hippocampal
formation in the dorsomedial forebrain. Subsequently released at the two familiarized sites, these birds were disoriented, provided that their sense of smell
was also impaired (Fig. 9.2A, OH). Obviously, only familiar-landscape orientation was disrupted, whereas homeward orientation based on olfaction was
162
9 The Neural Bases of Pigeon Homing
n Fig. 9.1. A transverse section of the pigeon telencephalon (stereotactic coordinate A 5.75 according to the atlas of Karten and Hodos 1967), showing locations of hippocampal formation
(Hp hippocampus; APH area parahippocampalis), piriform cortex (Cpi) and caudolateral
neostriatum (NCL). Ad Archistriatum, pars dorsalis; TeO tectum opticum; V ventriculus. (Modified from Bingman et al. 1998b)
still functional (O+H). OH+ control pigeons, on the other hand, demonstrated that birds with an intact hippocampus were well capable of orienting
homewards when equally deprived of olfaction as were the OH pigeons.
Corresponding to initial orientation, also homing performances (bar graphs
in Fig. 9.2A) were drastically reduced (P<0.001) in the OH birds, but to a
lesser degree (P<0.05) also in the O+H group. This minor reduction might be
due to the necessity of visual landmark orientation during the end phase of
homing when olfactory navigation approached its short-distance limit
(Sect. 7.4.1).
Results differed when training took place post-operatively (five training releases at each of the same two sites). Now the hippocampal-ablated pigeons
were remarkably well-oriented homeward even if olfactorily deprived (OH
in Fig. 9.2B). The most straightforward interpretation of these findings would
be that the birds normally store representations of spatial signals in the hippocampus and lose them when this brain region is lost, but also that acquisition,
storage and activation of relevant information are possible in other brain areas
when the hippocampus is not available. At this stage, however, it was not certain whether the observed behavioural similarities of OH+ and OH in
Fig. 9.2B reflected an analogous underlying mechanism.
It should be noted that these experiments did not strictly test the application
of some kind of map (familiar-area map, topographical map, visual-landscape
map, cognitive map). As training was restricted to homing flights exclusively
from the two locations at which the pigeons were afterwards tested, two-dimensional map-building was not necessarily involved. In a dual-choice task
the birds had only to decide at which of two expected sites they were (referring,
as a minimum, to some simple local marker such as a house, a group of trees, a
pond, etc.) and could then follow the entrained compass course leading home
9.1 The Hippocampus and the Familiar-Landscape Map
163
n Fig. 9.2. A, B Pooled vanishing bearings obtained at two familiar training sites 40 km NNW
and 55 km SSE, respectively, of their loft near Pisa (home directions superimposed upwards) and
means of the two corresponding medians of homing speed. O+ Olfaction intact; O olfaction
abolished or reduced by nasal anaesthesia; H+ hippocampus uninjured; H hippocampus ablated. Hippocampal ablation A after familiarization of release sites and B before familiarization.
C The pigeons used in B were subsequently released at an unfamiliar site 45 km ESE of home.
(Data from Bingman et al. 1987, 1988)
164
release sites with circadian clocks shifted 6 h

n Fig. 9.3. Vanishing bearings at two familiarized
forward. All birds were rendered anosmic (O ) by application of zinc sulphate (Sect. 7.1.2). H
and H as in Fig. 9.2. Filled symbols refer to OH+, open symbols to OH birds. Peripheral symbols indicate individual bearings of clock-shifted pigeons in two experiments at each site (distinguished by shape), heavy arrows the resulting mean vectors. Small-headed arrows indicate mean
vectors obtained from bearings of the same birds in the last training release before clock-shift (in
O+ condition). The one-headed peripheral arrow points to the approximate pseudo-home direction of the clock-shifted pigeons. (Data from Gagliardo et al. 1999)
from that site and/or could follow an entrained sequence of landmarks. The
moderately reduced homing speeds of OH pigeons in Fig. 9.2B may suggest
that final landmark-based home-finding was impaired. Further, more specific
investigations support the conclusion that, at least in the close vicinity of the
loft, some map mechanism is actually used (Bingman and Mench 1990;
Strasser et al. 1998).
A next experimental step confirmed that the structural basis of the good
performances after post-operative training (OH in Fig. 9.2B) was not fully
equivalent to the normal case with an intact hippocampus (OH+). The two
sorts of birds behaved differently when released at familiarized sites (eight
training releases) in a clock-shifted state (Fig. 9.3). The anosmic control pigeons (OH+) behaved as they usually do when exposed to conflicting information coming from the landscape and the sun (cf. Fig. 8.5C): They showed
considerable angular scatter and made, on average, a compromise between following the landscape pattern and following the shifted sun. The hippocampal-ablated birds (OH), in contrast, followed the sun entirely. A final interpretation of these results is not yet possible. The OH pigeons might have
applied a simple strategy as explained in the last paragraph (identifying one of
the two sites and then following an appropriate compass course without paying further attention to the landscape), whereas the OH+ birds apparently felt
the conflict raised by the unexpected position of the sun relative to the pattern
of familiar landmarks. Alternatively, both groups may have recognized this
pattern in a similar way but may have given different weights to the conflicting
cues: substantial dominance of the landscape over the sun in OH+ and 100%
dominance of the sun in OH (however, increased scatter compared with preceding control releases suggests that also the OH birds were in some way in-
9.2 The Hippocampus and the Olfactory Map
165
fluenced by the non-fit among the spatial signals). This latter interpretation
appears most appropriate with respect to recent results obtained with unilaterally hippocampal-ablated pigeons (Gagliardo et al. 2002). Both left- and rightside ablated birds, as compared to controls, produced initial orientation data
similar to that shown in Fig. 9.3. Homing performances were only moderately
reduced in these long-term anosmic birds, however, so that landscape-based
homing could not have been severely obstructed.
Also these clock-shift experiments did not definitely elucidate the map and
compass relationships. In order to produce and activate a topographical (cognitive) map, it would be necessary to conduct training releases from many sites
in varying directions, and finally to test the pigeons at an unexpected novel site
that is located within the familiarized area but that itself has not been used as a
release site before (cf. Fig. 8.1; Wallraff et al. 1994). Would post-operatively
trained OH birds orient homewards also under this condition which presents the landscape in a novel perspective and hence actually requires a map? So
far, one release has been reported suggesting that they would not (Bingman et
al. 1989). Anyway, Fig. 9.3 shows that learning and use of landmarks with and
without the hippocampus are not fully equivalent, although, at familiar sites,
non-shifted pigeons behaved indistinguishably (OH+ and OH in Fig. 9.2B).
It seems very likely, at least, that an intact hippocampus is necessary to establish
and to apply a fully functional map based on learned visual features of the landscape. Without a hippocampus, the sun compass is still operating but apparently
not spatially linked to the landscape pattern. It should be noted that the over-deflection by clock shift in Fig. 9.3 is possibly not merely a manifestation of usual
inaccuracy. For whatever reason, hippocampal-ablated pigeons also showed
larger clock-shift deflections than controls in several experiments at unfamiliar
sites (Bingman et al. 1996; Ioal et al. 2000a).
9.2
The Hippocampus and the Olfactory Map
The O+H diagrams in Fig. 9.2A have already shown that pigeons hippocampal-ablated after training were able to orient homeward if they were able to
smell. Thus, their pre-operatively developed olfactory map, which is supposed
to be a landscape-independent gradient map (Sect. 7.9), appeared unimpaired.
This map could apparently be applied by the hippocampal-ablated pigeons if
they were released at an unfamiliar site, provided that they were allowed to
smell (Fig. 9.2C, O+H versus OH+ and OH; see also other experiments,
e.g. Ioal et al. 2000a). The conclusion that not operation but acquisition of this
map by young pigeons requires an intact hippocampus (Bingman et al. 1990)
turned out to be valid only for pigeons that were, for a few months after fledging, confined in an aviary, but not for those that were allowed to fly freely
around their loft (Ioal et al. 2000b).Thus, under natural conditions, the hippocampal formation is apparently not necessary for operation as well as for development of the olfactory map.
166
9.3
The Piriform Cortex and the Olfactory Map
The piriform cortex (Fig. 9.1) receives a large projection from the olfactory bulb
and sends projections to numerous regions of the brain (Bingman et al. 1994). It
is, therefore, a candidate for the processing of olfactory signals that might be
used in navigation based on atmospheric odours. Consequently, Papi and Casini
(1990) ablated this brain area bilaterally in a group of pigeons at an age of about
3 months. In flock releases including also untreated control birds, the pigeons
were then trained to home, over two intermediate sites, from a site 40 km NNW
of the loft (five releases there). In a subsequent test release at this training site,
now with the birds flying individually, initial orientation as well as homing performances of the experimental pigeons were only slightly poorer than those of
the controls (Fig. 9.4A). At two unfamiliar locations, however, the ablated birds
maintained their training course southwards, whereas the controls shifted towards home (Fig. 9.4B,C). [The finding that the shift was not complete can easily
be explained by an effect of directional training which has often been observed
in untreated pigeons that hitherto had been homed from only one direction
(Sect. 3.2.2).] Also homing performance was significantly impaired by the surgical treatment.
These results support the hypothesis that the piriform cortex is involved in olfactory navigation. Further support comes from even more drastic differences
between untrained experimental and control pigeons at unfamiliar sites, which
additionally suggest that not only operation of a previously established olfactory
map but also its acquisition at an age between 1 and 3 months is prevented or, at
least, impaired in pigeons lacking the piriform cortex (Gagliardo et al. 1997).
n Fig. 9.4. Vanishing bearings and homing success (percent birds returned) of pigeons with ablated piriform cortex (open symbols) and of control pigeons (filled symbols) in three releases
(first A and then either B or C) as described in the text. (Data from Papi and Casini 1990)
9.4 Roles of Other Brain Regions and Hemispheric Lateralization
167
9.4
Roles of Other Brain Regions and Hemispheric Lateralization
A functional caudolateral neostriatum (Fig. 9.1), which shares some similarities with portions of the prefrontal cortex in mammals, also appears to be necessary for olfactory navigation, but not for homing based on familiar landmarks over short distances (Gagliardo and Divac 1993; Bingman et al. 1998b).
In contrast, lesions to the dorsal wulst (an anterior forebrain region considered
to be involved in processing of sensory, particularly visual stimuli) had no effect on homing from either unfamiliar or familiar release sites (Bingman et al.
1984). Nevertheless, according to Bingman et al. (1998b), there is some hint
that the dorsal wulst might be involved in the acquisition of familiar-landmark
homing abilities.
A number of investigations dealt with the question of whether, or to what degree, visual inputs or hippocampal functions during homing are lateralized to
one or the other cerebral hemisphere (Ulrich et al. 1999; Gagliardo et al. 2001c;
Prior et al. 2001; Diekamp et al. 2002). The results actually indicate some asymmetries. They may shed some light on the structure of brain functions, but so
far I do not see that they substantially contribute to the understanding of homing mechanisms.
9.5
The conclusion drawn elsewhere that pigeons make use of two independent
maps, one based on visual familiar-landscape features and the other based on
atmospheric odours (Sects. 8.1 and 7.9; Chap. 12), is well compatible with, and
supported by, the finding that these two functions appear to be largely under
the control of different parts of the brain, the hippocampus and piriform cortex. However, neither allocation nor support is as comprehensive as they could
possibly be. In almost all experiments conducted so far, only one or two
well-known training sites were used as familiar test sites, so that the birds were
not forced to develop and apply a two-dimensional familiar-area map which
should be applicable, within its range limits, also at previously not yet visited
sites from which the landscape is seen under a novel perspective (Sects. 9.1 and
9.3; cf. Sect. 8.3). Even if, in the close vicinity of the loft, configurations of individual landmarks are used in a hippocampus-controlled manner as shown in
studies with rats, food-caching birds, pigeons in laboratory tests etc. (e.g. Healy
1998; White et al. 2002), it is not certain that the same basic mechanisms are applied, or at least brain regions activated, by birds flying over an area they had
never overflown before, but from which they have a wide panorama-like aerial
view over a landscape they had previously seen from in-flight positions several
kilometres away (cf. Figs. 8.1 and 8.2). It remains a challenge for future investigations to determine whether also under these conditions visual way-finding
is controlled by the hippocampal formation.
168
It may depend largely on this question as to what degree training experiments with caged pigeons in a small arena can resemble free-flight conditions
in homing experiments over a range of some 20 or 50 km. Examples are tests
after directional training (1) in a circular arena where the birds could see the
sun as well as directionally fixed colour beacons (landmarks) along the arena
wall and (2) at a remote site from which they flew homeward. When the angular relationships between landmarks and the sun were uncoupled by means of
clock shift (Sect. 8.1.4), control pigeons in the arena tended to maintain the appropriate angle to the sun and to neglect the landmarks, whereas hippocampal-ablated pigeons neglected the sun and relied on the landmark beacons
alone (Gagliardo et al. 1996). In the homing experiments, however, the preferences were exactly reversed (Fig. 9.3). Thus, a generalization from one to the
other situation would have led to erroneous conclusions. Therefore, I refrain
from considering investigations in laboratory environments in more detail
(for further reading regarding this aspect, see Bingman et al. 1995, 1998b; Bingman and Able 2002).
10
Homing in Other Birds
Many ornithologists find it difficult to look upon domestic pigeons as real birds.
Consequently, they hesitate to accept that results obtained with homing pigeons
might be relevant also to starlings, warblers, swallows, hawks, gulls and albatrosses. In fact, we must not automatically presuppose that pigeon navigation
can be generalized to bird navigation. At least some crucial experiments conducted with pigeons should be repeated with other species of various avian orders. Unfortunately, because this is easier said than done, such repetitions are so
far scarce. We have seen that in order to answer a single question, it is usually
necessary to release large numbers of pigeons at different sites in order to obtain
statistical significance and clearly conclusive results. With wild birds, it is not
only much more difficult and time consuming to achieve comparable sample
sizes, but often impossible to achieve analogous data at all. Many birds do not
immediately fly away upon release and hence do not provide initial bearings, or
if they do, the bearings are initially not yet homeward-oriented (Sect. 10.1.2). In
order to collect useful numbers of recoveries, it is necessary to release manifold
more wild birds than homing pigeons (see, e.g., Perdeck 1958). Observation of
returners at the home site is also much more arduous and less reliable. There is
reason to expect, however, that things will become better in the future. Modern
satellite telemetry (Sect. 2.3) may make it possible to obtain more, and more
comprehensive, homing data from various species than is possible with conventional techniques. In the following text, with one exception (Fig. 10.6), I give the
current status of the pre-satellite age.
10.1
Experimental Approaches
10.1.1
Brief Overview
It is not the aim of this book to enumerate and discuss all the homing experiments that have been conducted, a great deal in the 1930s, 1940s and 1950s,
with birds of dozens of species belonging to a variety of taxa (e.g. petrels, gulls
and terns, swifts, swallows and starlings). Most of these experiments are listed,
with references to the literature, and summarized by Griffin (1944), Matthews
(1955, 1968), Wiltschko (1992) and kesson (2003). Distances of displacement
with subsequent homing varied from a few kilometres to more than 1,000 and
170
10 Homing in Other Birds
n Fig. 10.1. A Examples of long-distance homing flights of wild birds in Europe after experimental
displacement. A Alpine swift (Apus melba); B black-headed gull (Larus ridibundus); C red-backed
shrike (Lanius collurio); D wryneck (Jynx torquilla); E lesser black-backed gull (Larus fuscus); F
Manx shearwater (Puffinus puffinus); G Leachs petrel (Oceanodroma leucorhoa); H swallow
(Hirundo rustica); I starling (Sturnus vulgaris); J Arctic tern (Sterna paradisea); K white stork
(Ciconia ciconia). (Modified from Matthews 1968). B Homing experiments with Laysan albatrosses
(Diomedea immutabilis) that were displaced from their home island Midway Atoll in the northern
Pacific over long distances to various sites. Indicated are distances as well as numbers of displaced/returned birds and the duration until re-observation at home. (Data from Kenyon and Rice
1958)
10.1 Experimental Approaches
171
even 6,000 km. Return rates and homing speeds were extremely variable. Over
greater distances, they were often not very impressive (rate <50% and speed
<50 km/day), but, for instance in petrels, gulls, terns and swallows, remarkable
speeds in the range of 200400 km beeline distance per day were observed.
Some successful homing flights over longer distances are shown in Fig. 10.1A
and most spectacular performances in Fig. 10.1B.
Taken together, numerous experiments with wild birds displaced to far-distant areas, where they had hardly ever been before, demonstrate that goal-oriented navigation over unfamiliar terrain is a very widespread ability among
birds. The fact that performances are often (not always!) lower in wild birds
than those observed in homing pigeons need, and probably does, not indicate
lower navigational capacities. It is much more likely that the differences concern primarily the level of motivation to home at all and to home rapidly
rather than the ability to home (Sect. 1.2).
10.1.2
Compass Orientation and Preferred Compass Directions
The time-compensating sun azimuth compass (Chap. 5) was first detected and
investigated in a wild bird, the starling, Sturnus vulgaris (Kramer 1950, 1951;
Hoffmann 1954). It was only afterwards that it was shown to be used by homing pigeons as well (Kramer and Riese 1952; Schmidt-Koenig 1958). The magnetic compass (Sect. 6.3.2), first detected in robins, Erithacus rubecula (Merkel
and Wiltschko 1965; Wiltschko 1968), has up till now been more clearly demonstrated and more thoroughly investigated in migratory birds than in pigeons (e.g. Able 1994; Wiltschko and Wiltschko 1995). There is no indication
that compass mechanisms used by different avian species might be basically
different.
More remarkable than the apparent homology of mechanisms is the fact that
an unexplained kind of application of compass orientation has also been observed in various bird groups and under various circumstances in a similar
way. The very first starling, in which the existence of a sun compass has been
proved, already exhibited what later on was called nonsense orientation
(Matthews 1961) or preferred compass direction (PCD) (Wallraff 1978a;
Sects. 3.1.1, 3.7.1 and 7.5.3). With the sun being visible, this bird was, in spring,
consistently oriented towards west-northwest (Fig. 10.2), while its normal direction of migration would have been east-northeast. Under overcast skies it
was randomly oriented, thus showing that it determined the west-northwest
direction by means of a sun compass. In later years, many such cases were observed in which migratory restlessness of caged birds was not oriented towards the direction in which free-flying conspecifics would actually migrate,
but in a distinct nonsense direction. Such observations were made in birds belonging to various species, during daytime and night-time, under clear and
clouded skies, and in differently designed cages (e.g. Wiltschko 1980; Sandberg
et al. 1991; kesson 1993; Muheim et al. 1999; for more references see kesson
172
n Fig. 10.2. Spring migratory restlessness of a starling in a circular cage under a densely overcast
sky (A) and later on the same day under sun (B). Each dot represents the birds oriented activity
within a 10-s period. The bird maintained the northwest tendency in a number of experiments
quite consistently over weeks. (Modified from Kramer 1951)
2001). Individuals within a sample tested under the same conditions usually
preferred similar directions. Such nonsense PCDs often interfere with meaningful migratory orientation and thus render appropriate interpretation of results difficult (as they do in pigeon homing experiments; Sects. 3.7.1 and 7.5.3).
Even more comparable to pigeon homing than these cage experiments are
releases of wild birds displaced from their living area to distant, probably unfamiliar sites. Initial flights of terns, ducks and swallows were often clearly oriented towards a distinct PCD (e.g. Griffin and Goldsmith 1955; Bellrose 1958;
Matthews 1961, 1984; Baldaccini et al. 1999; Giunchi and Baldaccini 2001).
They did not reveal, however, as pigeon releases usually do (Sect. 3.7.1), an additional tendency towards home. Bellrose (1958) and Matthews (1963b, 1984)
showed that mallards (Anas platyrhynchos) use a sun compass during daytime
(as did the starling in Fig. 10.2) and a stellar compass at night to determine the
preferred direction. Clock shifts cause directional shifts under sun (Fig. 10.3)
as they do in pigeons (Sect. 5.1). The particular direction preferred by mallards
as well as by sand martins (Riparia riparia) is population-specific (Matthews
1963c; Giunchi and Baldaccini 2001), as it is loft-specific in pigeons (e.g.
Fig. 3.20).
At least in a descriptive sense,the PCDs observed in displaced homing pigeons
and wild birds, as well as in caged passerine migrants, appear analogous. Also
analogous is the fact that the PCD points most often, but not exclusively, in westerly (northwesterly, southwesterly) directions in all three categories of investigation. Moreover, analogous in all three categories is our complete lack of knowledge about origin and function of the PCD which, for us, still makes no sense
(Sect. 3.7.1). The assumption that the PCD is an immediate default response to a
strange or stressful situation may be reasonable. Thus it may help to keep a flock
together in a case of sudden disturbance by a predator (e.g. Matthews 1962;
173
n Fig. 10.3. Vanishing bearings of mallards (Anas platyrhynchos) displaced from their living site
over 45 km to the northwest. Control birds flew towards NNW, which was their usual PCD. Birds
with circadian clocks shifted 6 h forwards deviated more than 90 to the left, those with clocks
shifted backwards were correspondingly deflected to the right. (Redrawn from Matthews 1963b)
Thake 1981). Stress responses are even more likely to occur in freshly caught
wild birds rather than in domestic pigeons. Giunchi et al. (2003) have in fact
shown that sand martins switch from homeward to PCD-ward initial-flight orientation after stressful handling conditions, as has been found in pigeons
(Sect. 6.3.1). However, declaration of the PCD as an escape response would not
imply an explanation of the way in which a certain group of birds is triggered to
prefer just this and not any other direction. In some way the PCD reminds us of
the flight responses perpendicularly to a shoreline (y-axis orientation) as described for many shore-living animals (e.g. Ugolini and Macchi 1988), but a
shoreline determining the PCD of a given bird population has so far not been
recognized. Some observations suggesting a bipolar axis character of the pigeons PCD (Wallraff 1986, 1991a) would also fit into this category.
On the whole, a number of common features and no discrepancies have been
found in compass mechanisms as well as in particular types of compass orientation among domestic pigeons, on the one hand, and various wild avian species, on the other hand.
10.1.3
Goal-Oriented Navigation
More critical than compass orientation is the determination of a position relative to home. Can the conclusions drawn for homing pigeons be generalized?
In particular, do all birds, which are able to home from unfamiliar remote areas, determine their current position in such areas by means of airborne olfactory signals? Do all these birds (and possibly some other animals as well) have
a navigational map based on atmospheric trace gases? It is premature to decide
this question. So far, available experimental evidence, obtained with two species, is in favour of an affirmative answer.
The first of these species is the swift, Apus apus. In 43 specimens, caught at
night in their nests in Italy, Fiaschi et al. (1974) bisected one olfactory nerve and
plugged up one nostril with wax. As in corresponding pigeon experiments
174
n Fig. 10.4. Homing success of starlings displaced over varying distances east or west from their
breeding site in southern Bavaria. Columns (left ordinate) Percentage birds returned from birds
released: white columns starlings made anosmic by bilateral nerve section; black sham-operated
control birds. Line-connected dots (right ordinate) Return rate of anosmic starlings as a percentage of the rate of controls (respective black column = 100%). Numbers give sample sizes of displaced starlings. (Data from Wallraff et al. 1995)
(Sect. 7.1.1, Fig. 7.1), the two treatments were applied either contralaterally (experimental birds) or ipsilaterally (control birds). After release at 4766 km distance, 15 of 20 control birds returned to their nest, but only 3 of 23 experimental
birds. All of the latter three birds had lost their nose plugs when recaptured. The
difference in homing success, depending on ability to smell, is statistically highly
significant. Initial bearings were not homeward-oriented in either group.
Also in the second species investigated, the starling, only homing performances provided useful data (Wallraff and Hund 1982; Wallraff et al. 1995).
Anosmia caused by nerve section clearly reduced the return rates from release
sites beyond 100 km, but not from sites only 30 or 60 km distant (Fig. 10.4). In
its tendency, the dependence on the distance of displacement was equivalent to
the results obtained with pigeons (Fig. 7.2), but the range of no-effect was, with
at least 60 km, considerably larger in the starlings. However, no-effect within
this range concerned only the final homing success. The time required for
homing over 30 and 60 km was prolonged by anosmia (P=0.02); the median of
the observed-as-returned time was 15 days versus 9 days in the controls. (Medians of real arrivals were certainly earlier, but chances to be observed were the
same in both groups.)
On the one hand, and most importantly, these results strongly suggest that
olfactory inputs are utilized for home-finding not only by carrier pigeons but
also by other birds, which belong to different taxonomic groups. On the other
hand, the results give rise to the assumption that the hierarchical significance
of airborne odours within the set of spatial signals used for homing may vary
among avian groups. In swifts, olfactory deprivation prevented homing over
about 60 km (almost?) completely, whereas over this distance anosmic starlings returned at the same rate as the controls. This rate was, however, lower
than in swifts. The effect of anosmia in pigeons ranged somewhere in between
(Fig. 7.2). It is impossible to interpret the results in an appropriate manner, be-
175
n Fig. 10.5. 123 species belonging to 23 orders of birds according to their olfactory bulb ratio, i.e.
longest diameter of the olfactory bulb in percent of longest diameter of the brain. The graph
ranges from petrels and kiwi (left) to finches and chickadee (right). (After Bang 1971 from
Wallraff and Hund 1982)
cause the ranges are unknown within which the wild birds might have been
able to use familiar olfactory or non-olfactory cues.
We may, nevertheless, speculate about differences in olfactory sensitivity
and in the role that airborne odours play in the birds way of life. The relative
diameter of the olfactory bulb is about twice as large in swifts as in starlings
(Fig. 10.5), the relative volume, accordingly, about eight times larger. Thus,
starlings may be less sensitive to small changes of odours occurring over short
distances. Within a radius of 60 km, they may use odours as good as possible,
but not very effectively, and may use other, probably visual signals as well. After some searching around, which could last 12 weeks or more, about half of
the anosmic starlings eventually found their way back to the breeding site.
When released 120 km distant from home, olfactory signals may be reliable
enough to guide the birds back to the 60-km range, from which they find their
way back at the same rate as the starlings released there. Swifts, whose sense of
smell is probably better developed and which are permanent fliers in the atmosphere, may be very familiar with the olfactory landscape around them and
able to make effective use of it. Remaining in the air even during the night
(Weitnauer 1960; Bruderer and Weitnauer 1972; Bckman and Alerstam 2001),
they may be accustomed to rely on the always available olfactory signals and
not accustomed to searching around using alternative cues.
176
A more profound, less speculative discussion on the role of chemosignals in

spatial orientation of different avian species, families or orders would require
much more empirical data than are presently available. It is remarkable that
even the starling, a passerine bird whose olfactory system ranges morphologically, and certainly also functionally, at a rather low level (but which is evidently able to smell: Clark and Mason 1987; Clark and Smerasky 1990), can apparently deduce positional information from atmospheric trace gases at all.
This finding can be interpreted as an indication that olfactory navigation does
not require a high capacity to discriminate among many compounds. It may be
sufficient to evaluate ratios among a small set of specific substances, which are
particularly suitable for navigational purposes.
This assumption does not exclude a possible correlation between efficiencies of the olfactory and the navigational system. Most famous for their highly
developed sense of smell are the procellariiform seabirds, i.e. petrels, shearwaters and albatrosses, which rank at the highest left end in Fig. 10.5 (see also
Bang and Wenzel 1985). Some of these tubenoses are known to be very effective in smelling and localizing foraging zones in the seemingly featureless
open ocean (Nevitt 1999, 2000) and also in finding, by means of olfaction, their
individual breeding burrows after passive displacement over short distances
of less than 2 km (Grubb 1974; Benvenuti et al. 1993; Bonadonna et al. 2003b,
2004). These performances concern the localization of an odour source, usually by moving against the wind, and not navigation over hundreds or even
thousands of kilometres towards a distinct geographical goal, whose individual odour can hardly be identified over such distances and which can be
reached by seabirds from various directions (cf. Fig. 10.1B). It remains to be
shown whether also long-distance goal-finding in procellariiforms is based on
olfaction. As probably more extended and more regular gradients of ratios
among atmospheric trace gases exist over the oceans than over less homogeneous continental regions, and as tube-nosed birds have a particularly powerful sense of smell, it appears worthwhile to test whether their spectacular homing abilities are largely based on atmospheric (and perhaps oceanic) trace
substances (Sects. 10.2.1 and 12.3.3).
10.2
Homing in Natural Life
Up until now, the whole book has dealt with quite unnatural phenomena. Birds
were displaced by man to distant areas and then asked whether they are willing
and able, under varying conditions, to return to their home site or territory. In
the course of such experimental interferences, pigeons and other birds revealed remarkable navigational capabilities. We can postulate, however, that
evolution has not developed these capabilities in order to satisfy human experimenters. Thus, we have to look for activities in the birds natural life that may
involve and require spatial orientation over long distances beyond the range
within which previously learned familiar landmarks can be used.
10.2 Homing in Natural Life
177
10.2.1
Non-Migratory Excursions
We stay with the seabirds in order to meet the most impressive applications of
avian navigational abilities in natural life. In recent years, satellite telemetry
has made it possible to observe foraging flights of albatrosses, lasting several
days or even a few weeks and covering thousands of kilometres, sometimes
more than 10,000 km, over the open ocean (e.g. Jouventin and Weimerskirch
1990; Prince et al. 1992; Fernndez et al. 2001). Finally, the birds return to their
breeding island in the middle of the ocean. Some of these excursions are round
trips including smooth and sharp directional changes, sometimes loops, without indicating a distinct destination. In other cases, the albatrosses fly fairly
straight ahead to a feeding zone some 1,000 km or more distant either in oceanic waters, lacking any recognizable visible guide-posts, or in a continental
shelf area. After staying there for several days they return, again fairly straight
on, to their home island (examples in Fig. 10.6). In these latter cases the birds
commute, obviously by means of goal-oriented navigation, between two geographical positions.
n Fig. 10.6. Spontaneous excursions of three wandering albatrosses (Diomedea exulans) ob-
served by satellite telemetry over the southwestern Indian Ocean (A, B) and southern Atlantic
(C). The home island is marked by a circle. Dots along the route indicate localizations (in C: filled
dots night-time; open dots daytime). Distance scales are approximate (owing to map distortions
by projection). (A, B After Jouventin and Weimerskirch 1990 from Papi and Luschi 1996; C modified from Prince et al. 1992)
178
It is unknown on which environmental cues these navigational performances are based. According to kesson and Alerstam (1998), geomagnetic
parameters in some of the oceanic areas where such trips were observed are
unsuited to allow purely magnetic bi-coordinate navigation. Disturbance of
potential magnetic information in petrels and albatrosses did not disturb
homing flights (Benhamou et al. 2003; Bonadonna et al. 2003a; Mouritsen et al.
2003). Presently there are no explicit indications that oceanic seabirds might
find their far-distant goals by means of olfactory signals, but there are several
arguments in support of the hypothesis that they do so:
1. If it is true that pigeons and some other birds find their way home from unfamiliar areas by deducing positional information from atmospheric trace
gases (Chap. 7), why should oceanic seabirds not do so as well?
2. Ratio gradients of trace gases, basically suitable for navigational purposes,
exist not only in meso-scale dimensions over continental areas (Sect. 7.6.2),
but also in large-scale dimensions over and inside the oceans (see Fig. 10.7
as an example) and can even be expected to be more regular there than over
the less homogenous European continent.
3. Anatomically, procellariiform birds have a particularly large and well-developed olfactory system (Bang 1971; Bang and Wenzel 1985).
4. Functionally, procellariiform birds are known to apply this system for orientation over considerable distances towards sources of food in seemingly
featureless oceanic landscapes (Nevitt et al. 1995; Nevitt 1999, 2000).
5. During their excursions covering thousands of miles in an environment
lacking visual landmarks, seabirds have an opportunity to develop an olfactory topographical (mosaic) map of a very extended familiar area.
n Fig. 10.7. Concentrations of two atmospheric trace gases measured on board a ship over the
Atlantic during the same cruise in October/November 1996 roughly along the meridian 30W
from north to south. (Data from Fischer et al. 2000 and Weller et al. 2000)
10.2 Homing in Natural Life
179
Considering the previous five issues, it would not be too surprising if seabirds had a particularly sophisticated olfactory navigation system comprising
learned local or regional signals as well as signals allowing, due to their inclusion of large-range gradients, extrapolation into unfamiliar areas. It might be a
complex system within which a clear distinction between mosaic map and
gradient map (Fig. 4.1) makes little sense, as it may consist of a mosaic of overlapping gradient fields with varying spatial extension. Owing to the complete
lack of empirical data, it would be premature to extend speculations into this
field.
In overland foraging flights of other birds over shorter distances, olfactory
navigation may also be involved in some cases. However, as such flights usually
occur within a familiar area, it is not immediately evident to what degree olfactory and/or visual cues are used for orientation. An example where usage of olfactory signals would appear profitable are bad-weather movements of swifts
(Apus apus) during which the birds apparently fly away over hundreds of kilometres, partly over sea and at night, and later return to their home territory
(e.g. Lack 1958; Weitnauer and Scherner 1980; Offringa 1996). It may not be by
chance that this airborne species appears to rely more on olfactory rather than
on visual cues (Sect. 10.1.3).
10.2.2
Homing as a Constituent of Bird Migration
Apart from the seabird voyages, bird migration is certainly the context within
which homing mechanisms are most substantially applied. Perdecks (1958)
famous displacement experiments demonstrate that starlings during migration actually make use of such a mechanism. Considering the results shown
above in Fig. 10.4 and in Chapter 7, it seems reasonable to assume that his displaced adult starlings re-oriented their courses towards the earlier experienced wintering grounds by evaluating airborne olfactory inputs. There is one
experimental hint supporting the assumption that olfaction is indeed involved
in migratory navigation (Wallraff et al. 1995): The percentage of anosmic starlings that were observed at their breeding site in spring of the following year
was very low (3 out of 40 = 7.5%), significantly lower than the percentage of
sham-operated starlings with an intact sense of smell (18 out of 63 = 28.6%). It
is not proved but, against the background of other findings, not unlikely that
the cause of the reduction was a deficit in navigational capability.
Even if we sympathize with the hypothesis that olfactory navigation is used
and necessary to reach the previously experienced breeding and wintering territories, we need not presume that long-distance migrants hold uninterrupted
olfactory contact with their home area. The atmospheric gradients, which they
may use, need not extend from Scandinavia to South Africa or from Canada to
Argentina. Navigational ranges depending on such gradients may be as limited
as they appear to be in pigeons (Sects. 7.4.1 and 7.6.5). As a basic skeleton of
migration, a genetically encoded time-and-direction programme, leading ju-
180
n Fig. 10.8. Schema explaining the hypothetical role of olfactory navigation in long-distance mi-
gration. The area thought to be visually familiar is shown in black, the range within which olfactory navigation is thought possible is grey. Dashed line with arrowhead indicates the genetically
programmed compass direction, which is actively intended by the bird (thick arrows), but from
which it is drifted away by side winds or deflected by an ecological barrier (dotted area). A First
autumn migration; B spring migration. In A, the bird reaches, by following its population-specific time and direction programme, a roughly determined wintering region within which it
looks for an ecologically suitable habitat. In B, to reach directly the small familiar breeding area,
the bird would have to stay within a very narrow angular range (thin dashed arrow rays). This is
impossible, and unnecessary, if the bird only needs to reach a much larger target area, within
which it is able to home by using olfactory gradients. (Wallraff 2003)
venile birds to their population-specific winter quarters (e.g. Berthold 1996,

2000; Mouritsen 2003), certainly persists also in adult birds. It includes an intended compass direction (or a sequence of directions) roughly connecting
the breeding area with the wintering area and its reversal. However, by only
keeping the genetically intended compass course over thousands of kilometres, a returning migrant would hardly reach its familiar breeding area with
a diameter of at most 50100 km. During its long, possibly nocturnal journey,
wind drift and ecological barriers, such as coastlines or mountain ridges, deflect the bird from a direct route, and it is questionable that it is able to compensate for the enforced deviations without referring to position-indicating
geographical signposts. The problem can be solved if the target area, which
must be reached by keeping an intended compass course as good as possible, is
considerably larger than the familiar home area (Fig. 10.8). If the target area is
the range within which olfactory navigation exploiting atmospheric gradients
is possible, it may be sufficient for a warbler coming from Africa to arrive, by
181
means of compass orientation, somewhere in central Europe. The bird may

then reach its familiar home area by using olfactory signals and finally find its
last-years territory by remembering visual landmarks.
Very little is known about the involvement of olfactory navigation in bird
migration. The areas overflown between summer and winter territories need
not remain so olfactorily blank for the birds as drawn in Fig. 10.8. Even if there
are no monotonic atmospheric gradients covering the whole spatio-temporal
living range of a population, the birds may consider olfactory conditions experienced en route and possibly recognize and evaluate the composition of
large-scale olfactory landscapes. The scheme in Fig. 10.8 is probably too simple. For instance, also stopover sites may be visually and olfactorily characterized and embedded in a navigational system. The field is still open for speculation; our knowledge about possibly relevant atmospheric conditions is very
limited.
10.3
So far, there is no indication that different taxa of birds, including homing pigeons, apply basically different mechanisms for long-distance navigation.
Therefore, it seems justified to use the homing pigeon as a model bird in which
these mechanisms can most easily and effectively be investigated. Generalizations from pigeon homing to bird homing appear valid at least to a considerable degree. Nevertheless, two critical issues should be kept in mind.
First, although there is no contra-indication against generalizations, experiments testing their actual validity are still restricted to only a very few species
of wild-living birds (Sect. 10.1.3). A definitely positive conclusion would require a larger sample of comparative investigations on a broader taxonomic
and ecological basis. So far, we cannot exclude, for instance, that some species
determine positions not (only) by means of olfactory but (also) by means of
magnetic inputs. Actual available indications, however, suggest merely largerange determination of lines of position, namely latitude, on a magnetic basis
(Sect. 6.3.3).
Second, even if there is little variance in the basic principles of navigational
mechanisms, different components are variably developed in different species
and have a variable hierarchical position. Many non-migrant species, for instance, appear to be unable or unwilling to home over longer distances from
outside their familiar area (e.g. Matthews 1955, 1968; Wiltschko 1992). No wonder they do not need such an ability in their common life. However, even
among species apparently able to navigate by means of olfaction, the hierarchical role and efficiency of visual and olfactory orientation appear to differ
(Sect. 10.1.3). Oceanic seabirds, which have little opportunity to use visual
landmarks, appear to have developed a particularly effective non-visual navigation system (Fig. 10.6). It may be, nevertheless, an olfactory system basically
equivalent to that of pigeons, but more sophisticated using a more powerful
182
sense of smell, which alone is responsible for navigation over very long as well
as over short distances. Of course, as yet, it cannot be excluded that seabirds
use (also) non-equivalent methods.
It is apparent that birds of many species are able to navigate to a geographical
home position over wide unfamiliar territories. It is less apparent whether the
mechanisms behind this ability are largely the same in all species. Thus, it is necessary to extend comparative investigations to a variety of species living in varying ecological environments. Our knowledge of pigeon homing, especially of olfactory navigation (Chap. 7), can serve as a template for such studies into which
findings obtained with other species may fit or not fit. It is of particular interest
to investigate the degree and the manner in which olfactory navigation might be
integrated in the control system of bird migration. Modern techniques of telemetry and route recording should facilitate related experiments with wild-living
birds which hitherto were impossible (Sect. 2.3).
11
Research History:
Blind Alleys and an Unexpected Passage
Viguier (1882) was probably the first to contemplate a magnetic map, using
geomagnetic inclination and intensity as coordinates, as a possible basis of animal homing, including pigeon homing. A few years earlier, Darwin (1873)
speculated that animals might find their way home from a long distance by
dead reckoning (path integration). More concrete experimental research
started at the beginning of the 20th century in whose first half displacements
of birds of many species revealed that homing over considerably long distances is a widespread ability among birds (for review see Griffin 1944;
Matthews 1955). Thus, it became clear that there is an extraordinary problem
to be solved. The more cautious and conservative researchers, hesitating to
speculate on enigmatic unknown senses, tried to explain home-finding by
means of visual landscape knowledge combined with some more or less systematic search strategies (e.g. Schneider 1906; Riviere 1923; Heinroth and
Heinroth 1941; Griffin 1944). Riviere (1929), referring to pigeon races along a
trained line and to related experiments, speculated on a sense of direction and
even on the possibility that it might rely on the position of the sun. Realizing
that pigeons appear to find home also over long distances from untrained directions, he furthermore speculated on a sense of geographical position. Others refreshed the idea that such senses might use (electro-)magnetic inputs
(e.g. Thauzis 1904, 1913; Casamajor 1927; Daanje 1936), but without providing empirical evidence supporting such assumptions. A number of experimental approaches have been made in order to test hypotheses proposing kinaesthetic sensations, i.e. to test whether pigeons or other birds might record
their movements during transport to the site of release by measuring angular
accelerations with their semicircular canals (e.g. Exner 1893; Hachet-Souplet
1911; Huizinga 1935; Kluijver 1935; Rppell 1936). Neither of these attempts
included an indication that birds might apply some kind of inertial navigation
(as later theoretically outlined by Barlow 1964, as a particular type of path integration). A comprehensive historical survey until about 1950 is given by Griffin (1944, 1952).
Rigorous experimental research on mechanisms of bird homing, in practice
of pigeon homing, did not begin before the middle of the last century and
hence embraces little more than 50 years. The starting point of this period is
marked by a very specific geophysical grid hypothesis and attempts to verify
its validity by displacement experiments with pigeons (Yeagley 1947, 1951).
The theoretical basis is grounded on the fact that the geomagnetic poles di-
184
11 Research History: Blind Alleys and an Unexpected Passage
verge from the geographic poles and that, therefore, magnetic latitudes are not
fully identical to geographic latitudes. If both could be measured independently, a grid of two coordinates would be available which, however, intersect in
most regions at small angles, i.e. are much closer to parallel than perpendicular. Yeagley proposed the two parameters as being the magnetic vertical component and the (mechanical) Coriolis force which is caused by the earths rotation and which could, if at all, be measured only during flight. The physical
basis of the hypothesis as well as its claimed experimental support were severely criticized over the following few years (Griffin 1952; Matthews 1955,
1968, and references cited therein). Thus, debates and experiments dealing
with the Coriolis force/magnetic field hypothesis have quickly become an historical episode.
The real pioneers initiating an enduring period of intense research on pigeon homing were, independently of each other, Gustav Kramer in Germany
(Kramer and von Saint Paul 1950, 1952) and G.V.T. Matthews in England
(Matthews 1951). Kramer (1950, 1951) contributed the newly detected avian
sun compass and demonstrated that also homing pigeons can make use of it
(Kramer and Riese 1952). Matthews (1951, 1953, 1955) made the sun, theoretically, the key cue considered to be used not only for compass orientation but
even for the whole process of home-finding from unfamiliar areas. The sun
navigation hypothesis proposes that pigeons dispose (1) of the ability to observe very precisely not only the suns altitude, but also the rate of change of altitude, (2) of a very precise chronometer conserving local home time even after
eastwest displacements, and (3) of memorized knowledge of the suns position at home at any time of day. Using these data, determination of the direction towards home from any distant site is, in principle, possible (for details of
an improved theoretical framework, see Pennycuick 1960). The sun navigation
hypothesis remained in the focus of debate and experimental research until
Kramers untimely death in 1959; its spirit faded out during the following decade. Growing experimental evidence revealed that displaced pigeons do not
use the sun to determine their current position (e.g. Kramer 1957, 1961;
Schmidt-Koenig 1965, 1979; Keeton 1969, 1974a; Wallraff 1974a).
Instead, experimental evidence made it clear that the use of the sun is restricted to the function of a compass (Chap. 5). The fact that the sun compass
could be identified as being a separate element of the homing process
(Schmidt-Koenig 1958, 1961) supported the map-and-compass concept suggested by Kramer (1953b) several years earlier (Sect. 5.1; Fig. 5.5). Yet this concept left one of its two parts unlabelled. While compass could be complemented with the prefix sun, and later alternatively also with magnetic
(Sect. 6.3.2), no physically defined attribute could be associated with map.
The magnetic field as a possible (part of an) indicator of position appeared
outdated not only with respect to the specific hypothesis of Yeagley (1947), but
also more generally, because several attempts to interfere with homing by attaching magnets to the wings of pigeons remained without effect (Gordon
1948; Matthews 1951; Van Riper and Kalmbach 1952).
185
Research continued over about two decades (ca. 19551975) without being
guided by particular ideas on the physical basis of homing navigation. This period was, nevertheless, productive in that it accumulated lots of empirical data
revealing peculiarities of the pigeons homing behaviour (e.g. Kramer 1959b,
1961; Schmidt-Koenig 1965; Wallraff 1967, 1970a; Keeton 1974a). The broadened empirical basis (as described in Chap. 3) made it possible to ask specific
questions henceforth more adequately, to adapt experimental methods and to
avoid inappropriate interpretations. Theoretical approaches, however, were restricted to general considerations on possible properties of physically undetermined coordinates (gradients) and their possible evaluation for the purposes of home-finding (Wallraff 1974a; Wiltschko and Wiltschko 1978b).
During the 1970s and 1980s, after it had been detected that birds use the geomagnetic field for compass orientation (Merkel and Wiltschko 1965; Wiltschko
1968), interest in magnetism as a potential basis also of position determination
experienced some revival. It was not driven by new concepts or hypotheses
about the potential structure of a magnetic map, but several findings indicated
now an influence of magnets attached to flying pigeons or of exposure to artificial magnetic fields before release and by apparent disorientation in the area of
magnetic anomalies (Sects. 6.1 and 6.2). Some of these influences and correlations were not very consistent, however, and generally they were restricted to
disturbance or modification of initial orientation behaviour just after release
and did not prove impediment of homing performances (Sect. 6.3). Nevertheless, toying with the idea of a magnetic map used by pigeons is still alive and has
recently even led to a new (though unrealistic) specific hypothesis (Walker 1998,
1999; Walker et al. 2002; but see Wallraff 1999; Reilly 2002; Wiltschko and
Wiltschko 2003b). In the period of ca. 19701980, also other potential cues were
shown to be unimportant or, at least, non-essential for home-finding. Among
them were all conceivable kinds of sensory input picked up during the outward
journey to the site of release (Walcott and Schmidt-Koenig 1973; Wallraff 1980a).
Thus, any method of path integration operating during passive displacement
could practically be excluded (Sects. 4.2.1 and 6.3.4). Seeing the landscape at
home or during the homing flight was found to be non-essential in determining
a rough direction towards home (Figs. 7.19 and 8.3) or even to finding the vicinity of home (Schmidt-Koenig and Walcott 1978). The conclusion that visual
landmarks are not used by pigeons during homing flights, except in the vicinity
of the loft (Schmidt-Koenig 1979), however, obviously went too far (see below).
Also acoustic stimuli, including infrasounds, were shown to be unnecessary to
find the way home (Wallraff 1972; Sect. 4.2.2).
However, not only not necessary findings were obtained. Several series of
experiments, initiated by Kramer (1959a,b) and later continued (Wallraff
1966a, 1970b, 1979), with pigeons raised in differently shielded aviaries
(Sect. 7.5.1) were successful in that they were the first to show that it is at all
possible to deprive pigeons of their capability to find the way back to their
home from distant sites. In order to develop a navigational map, the birds apparently needed to be exposed to natural airflow, but did not require visual
186
contact with the surrounding environment. I stated the involvement of unidentified atmospheric factors, but I did not believe that they might be chemical substances dispersed in the atmosphere. It was obvious that the pigeons
need not smell a home loft odour because, first, this odour would not have
been shielded by walls surrounding the aviary and, second, home odours could
never reach a release site some 100 km upwind from home. I could not imagine
in what way other airborne chemicals might provide positional information
over such long distances. So I was closest to the deciding next step but did not
make it.
Fortunately, others were less burdened with preconceptions and made the
deciding step. Realizing that the sense of smell had not yet been considered in
the context of avian homing, Floriano Papi and colleagues in Pisa bisected the
olfactory nerves of ten homing pigeons and released them at some distance
from their loft (Papi et al. 1971). The deteriorating effect of this treatment was
obvious and could soon be confirmed by further experiments, adding not only
additional data but also support from differently designed approaches (e.g.
Papi et al. 1972; Papi 1976). It was not immediately obvious, however, that
olfaction-based homing is not simply a kind of extended landmark orientation, allowing some short-range extrapolation beyond an explored radius of
activity. Rather, it was several years more before it became apparent that airborne chemosignals, together with winds and the compass references to the
sun and magnetic field, are the only substantial sources of information that are
used for home-finding from unfamiliar areas even over hundreds of kilometres. The matter is broadly discussed in Chapter 7 and therefore need not be
repeated here. Looking retrospectively at this period with all its results and debates, I do not hesitate to state that the ten anosmic pigeons released in 1971
54 km west of Florence, Italy, opened an unexpected passage towards a solution of the mystery of avian navigation. Nevertheless, as long as a final solution and full understanding have not been achieved, I cannot definitively exclude that a future historical retrospect will merely state another blind alley. (If
I did not feel pretty sure that this will not be the case, I would not have written
this book.)
Once it was possible to eliminate one key sensory input used for home-finding, it was possible to ask again whether pigeons take the visual landscape
within a familiar area into account. As olfactory signals are also available there,
so that responses to visual and olfactory signals cannot be distinguished, only
anosmic pigeons could provide answers. It turned out that pigeons apparently
have two maps, a visual topographical map, applicable only in a familiar area
around home, and an olfactory gradient map, which can be applied also in
more distant unfamiliar surroundings (Sects. 8.1 and 8.3). The two maps were
shown to be represented in different parts of the brain, and particularly visual
landmark orientation, as localized in the hippocampal formation, has been investigated more closely under neuroethological aspects (Chap. 9).
12
Overall Synthesis and Perspective
This concluding section condenses the many particulars exhibited in the pages
before in a brief take-home message. Also, it tries to outline some problems
that should most urgently be tackled by future investigations.
12.1
Environmental Cues Involved in Home-Finding Processes
There is no reason to place the mystery of pigeon homing in a paraphysical
world of psi factors, dowsing rods, earth rays, morphogenetic fields (Sheldrake 1994) or something alike beyond the borders of traditional science.Pigeon
homing is well accessible to classical methods of experimentation which actually have revealed the nature of physical cues involved in the home-finding system. These are the cues that I consider as identified in principle:
1. The temporally variable azimuth of the sun as an omnipresent directional
(compass) reference.
2. The geomagnetic poleward direction as another directional reference available in parallel.
3. Wind directions at the home site in terms of angles to these compass references.
4. Atmospheric chemosignals correlated with wind direction at home and
evaluated in distant areas as indicators of radial position from home.
5. Visual landscape features in a familiar area around home including their relationship to the compass references.
So far, there is no indication that any other components of the environment act
as navigational signals. As long as the sun is visible, the directional references
(1) and (2) imply some redundancy, with the sun usually dominating over
magnetism. The signals mentioned under (4) and (5) are known as categorical
features only, whose concrete composition in detail has not yet been identified.
In a familiarized area around home, they constitute a largely redundant double
system, within which category (5) loses and category (4) gains relative weight
with increasing distance from home.
188
12 Overall Synthesis and Perspective
12.2
Two Homing Mechanisms
It is obvious that pigeons apply two different homing mechanisms. One of
them is trivial insofar as it basically bears the same problems of spatial orientation that all animals have to solve in order to find their way about in a largely
explored and familiarized area. In detail, these problems are all but trivial and
fill thousands of relevant publications which can, by and large and with pros
and cons, be subsumed under the heading cognitive map (e.g. OKeefe and
Nadel 1978; Gallistel 1990; Bennett 1996; Kamil and Cheng 2001; Mackintosh
2002; Jacobs and Schenk 2003). Pigeons apparently have such a map (familiar-area map, topographical map, mosaic map) which refers to the familiar visual landscape (Chap. 8). The spatial range of this map depends on the range of
the spatial experience of the individual bird, and its reliability and accuracy
certainly depend on the degree of experience (i.e. on repetitions and spatial
density of previous presence in a given area). Little is known about the peculiarities of a topographical map of a bird flying in a wide airspace as compared
with terrestrial animals (rats, mice, etc.) whose spatial orientation in the laboratory has been studied most intensely. I even have some doubts whether it is
heuristically productive to extrapolate from arena or maze experiments with
pigeons, using small nearby landmarks, to open-field conditions allowing the
birds an overview over a wide aerial panorama in which distinct small-scale
landmarks may play merely a minor role (Sects. 8.3 and 9.5).
Closer investigation of visual landscape orientation by homing pigeons is not
only difficult because it occurs in a wide space where landmarks can hardly be
manipulated. It is also complicated because the second homing mechanism, olfactory navigation, operates concurrently, so that responses to visual and olfactory signals cannot be separated. Thus, visually guided homing alone can only
be analysed in pigeons that have been made anosmic (Chap. 8). Then it turns out
that the sun is usually included in the pattern of landmarks used by pigeons, i.e.
that they pilot home using their topographical map together with the sun compass. If the latter is shifted by clock shift, the birds are in a state of conflict, because the suns azimuth does not fit in the expected way in the familiar pattern of
visual cues (Sect. 8.1.4).
The problem of redundancy does not play a role in more distant areas lacking any familiar landscape features. Then homing is based on true navigation
and specifically, if the presently available data reveal the truth, on olfactory
navigation alone. So far, the environmental input signals of either mechanism
are identified in principle (visual landscape, atmospheric trace gases), while
the particular components that are actually processed, and the modes of processing, are largely unknown in either case. It is known, however, that different
brain areas are primarily responsible for the two homing systems (Chap. 9). In
the olfactory mechanism, directional and positional markers (compass and
map) seem to be so closely linked to a unit that clock shifts result in angular
shifts of that unit as a whole (Sect. 7.9).
12.3 Challenges for Future Research
189
12.3
Challenges for Future Research
Possibly, this monograph will later be seen as a compendium of classical pigeon homing research which in this form lasted for about half a century. Certainly, pigeon releases as described here will be conducted also in forthcoming
years, but the decisive open questions can only be solved by interdisciplinary
research applying other and more advanced methods as well. In the following,
I give some hints on possible aims of future investigations. Others may have
other, possibly more productive ideas.
12.3.1
Unsettled Basics of Pigeon Homing
Even the basic features of pigeon homing are not yet understood in full detail.
For instance,the complex including the roles of age,experience (learning what?),
selection and motivation (Sects. 3.1.2 and 3.2) has not yet been exhaustively
studied. As it involves many variables, however, its closer analysis requires many
carefully designed series of experiments and huge amounts of statistical data.
Considering the expected costbenefit ratio, I have some doubts whether such
analyses on a broader basis than have so far been carried out will ever be conducted.
Up to now, only few attempts have been made to correlate aperiodic fluctuations of pigeon orientation and homing (Sect. 3.3.2) with concurrently occurring fluctuations of meteorological conditions (Wallraff 1960; Dornfeldt 1977,
1996). Since we know that the atmosphere is directly involved in pigeon homing (Chap. 7), not only correlative but also causal interrelations should be expected. Related investigations are worthwhile doing, but also they require large
amounts of carefully collected data. Owing to the many interfering sources of
variability, it is more difficult to obtain reliable material than one might expect
(cf. Kiepenheuer et al. 1993).
Advanced techniques, most effectively recorders using satellite-based positioning systems (GPS) (Sect. 2.3), allow us to gain information not only on the
pigeons initial orientation and homing speed but also on their complete flight
routes. It should thus be possible to find out whether the routes of individual
birds from a given site towards home tend to follow similar detour paths. If yes,
one could ask to what degree these paths appear adapted to the regional topography (cf. Ioal et al. 1994) and to what degree they appear to follow invisible
rules (cf. Fig. 7.34B). Of course, these techniques can also be applied in combination with particular experimental treatments and thus in the direct context
of the two following sections.
Finally, there is the riddle of the preferred compass direction (PCD)
(Sects. 3.1.1, 3.7.1, 7.5.3 and 10.1.2). As outlined above, its existence is presumably not a necessary consequence of the home-finding process. Rather, it
seems to be invented to confuse human investigators not only of pigeon hom-
190
ing but of bird orientation in general (nonsense orientation: Sect. 10.1.2). Beyond that, one would like to know whether it has any other functional purpose
as well, either navigational or non-navigational. Not only its function but even
more its ontogenetic creation is unsettled. What makes pigeons from one loft
fly preferentially towards SW, those from another loft towards NW and those
from a neighbouring third loft towards ESE? What makes the directional preferences fluctuate in time (Sect. 7.4.2)? What are the interrelations between
winds, PCD, homeward orientation and olfactory signals (Sect. 7.5.3)? I do not
definitely exclude that this complex comprises a not yet deciphered riddle of
the homing system.
12.3.2
Problems of Visual Landscape Orientation
One of the unsettled problems of visual orientation in a familiar area is the
question of whether birds in flight refer to selected individual landmarks or
more to a holistic aerial panorama of the entire landscape below them
(Sect. 8.3). Also, it would be interesting to know whether birds exploit their
large bi-monocular visual field by considering all its parts at once or whether
they focus on the parts of a large field sequentially as we do. The finding that
visual orientation alone seems to provide a similarly vague idea of the direction towards home as olfactory orientation alone, but is effective even at sites
several kilometres away from previously visited sites (Fig. 8.1), may suggest
that a hypothesis favouring the use of the appearance of a wide panorama
rather than of a set of individual landmarks is more appropriate. Owing to
their wide visual field, pigeons might overview a large area at once, which
looks similar from aerial observation points several kilometres away from
each other.
An approach towards solving the at once problem could be made in the laboratory. By some method of conditioning in fixed position (see, e.g., Hahmann
and Gntrkn 1993), it could be tested whether a pigeon is able to pay simultaneous attention (short exposure!) to two or more objects (e.g. figures on
monitor screens) distributed at a few metres distance over an angular range of
some 90180. If initial results are positive, more detailed analyses could be
performed by varying number, positions, sizes and/or pattern elements of objects.
Another approach could tie in with homing experiments as described in Section 8.1.1, but now using modern techniques that allow analyses of complete
homing routes. Biro et al. (2002) and Guilford et al. (2004) have shown that GPS
tracking (Sect. 2.3) can provide elucidating data. If applied not only in the close
vicinity of the loft, as in those studies, but also in investigations analysing sequences of homing routes from repeatedly used distant sites as well as from sites
more or less far away from each other, and including pigeons made anosmic,
much more elaborate and specific results than those shown in Figs. 8.1 and 8.2
could be expected.
12.3 Challenges for Future Research
191
12.3.3
Problems of Olfactory Navigation
Most challenging, of course, is homing based on atmospheric chemosignals.
Obviously, it does occur, but the empirical basis showing its occurrence should
be broadened and, most importantly, we should try to achieve an understanding of how it can operate. There are several very different approaches that need
to be taken:
1. Extension of basic material. Although there is, in my view, sufficient experimental evidence justifying the conclusion that long-distance homing of pigeons, and apparently also of other birds, essentially depends on olfactory
inputs, additional data obtained by researchers working independently of
each other would still be welcome. According to reasonable scientific rules,
general acceptance of fundamental findings, especially if at variance with
commonsense expectations, requires multiple replications. Related experiments could be simple repetitions (conducted with sufficient care:
Sect. 7.8.3), but would have additional heuristic value if they would include
modern tracking techniques (Sects. 2.3 and 12.3.2) and/or were performed
with birds of varying species. In the latter case, possible involvement of olfactory navigation in the particular birds natural life (Sect. 10.2) should be kept
in mind. Most suitable for olfactory deprivation of wild birds in the field is
nasal irrigation with zinc sulphate (Sect. 7.1.2). In large birds, such as albatrosses or geese, its possible influence over several days after application
could most effectively be observed by using satellite telemetry (Sect. 10.2.1).
2. Analysis of the chemical atmosphere. The suns position in the sky can be exactly calculated for each time and place on the earth. Magnetic maps are
available for the whole globe; the geomagnetic vector can be exactly measured everywhere. Thus, as far as birds respond to such input data, we can
precisely determine the available signals and hence can focus on the birds responses. When dealing with olfactory navigation, however, we know no more
about the related signals than that they are in the air, moved with the wind
and can be perceived by the birds olfactory system. Our principal problem at
the current early stage is not so much the animal but its atmospheric environment. However, what shall we search for in the atmosphere? The first challenge is thinking about an alternative to the hypothesis proposed above
(Sect. 7.6.1). The second challenge, if no alternative comes to light, is verifying or falsifying that hypothesis by trying to investigate chemospheric ratio
gradients and their variations, as correlated with seasons, weather and winds,
much more intensively and extensively than has been done in the first pilot
approach (Sect. 7.6.2). Future investigations should pay particular attention
to rules according to which ratios varying with wind direction are correlated
with the directions of spatial ratio gradients. Also, they should try to include
more of the naturally emitted (non-anthropogenic) compounds.
192
3. Identification of chemical compounds used for navigation. It is certainly difficult to fill this hitherto completely blank sheet. One approach might be to
search for atmospheric trace compounds to which pigeons are particularly
sensitive. This could be tried by gas-chromatographic sequential sorting of
airborne compounds, leading them into a nostril of an immobilized pigeon
and investigating potentially correlated nerve or brain activities, the latter,
e.g., by using functional magnetic resonance imaging (fMRI) (cf. Sobel et al.
1999). Lacking fMRI responses, however, would not say very much, because
compounds might specifically stimulate the brain only when administered
in specific combinations. Also, as the system should be adapted to continuous stimulation, it might not appropriately respond to sudden events. Once
a particular compound is, for whatever reason, suspected to be a candidate
stimulus component, it could be added, at very low concentration, to the air
the pigeons breathe at the release site before they are released under nasal
anaesthesia. The site-specific natural pattern of ratios, and hence initial orientation, should then be disturbed, whereas compounds without specific
relevance should have no effect.
4. Analysis of olfactory signal processing. It is very likely that signal processing
in the context of olfactory navigation does not fully follow the rules otherwise known in the neurophysiology of the sense of smell, mainly because atmospheric concentrations are extremely low and because the system must
be permanently ready to respond without habituation (Sect. 7.7). At the
moment, I do not see any potentially useful method to tackling this problem, at least not before some progress is made in solving the question
treated in the preceding paragraph. Additional difficulties may arise from
the possibility that behavioural responses cannot be elicited in the laboratory but only while birds are motivated to home under field conditions.
12.4
Outlook
The present state of research on avian homing is to some degree satisfying,
provided that the basic conclusions drawn here on olfactory navigation are not
only confirmed but also more than so far generalized from homing pigeons to
wild-living bird species. Then we can be fairly confident of knowing at least the
basic categories of environmental signals that are used by birds to find their
way back to a previously established home site over distances of a few kilometres in a familiar area to several hundred or thousand kilometres over never
before experienced terrain. The categories of decisive cues can then be considered as identified: the visual landscape and atmospheric chemosignals (together with the sun and geomagnetic field as directional references and with
wind direction recorded at home).
The present state of research is not yet satisfying as regards our knowledge
of the nature of decisive signals within these categories. Especially the concrete
12.4 Outlook
193
nature of the actually used atmospheric trace compounds, their spatio-temporal distribution in the airspace and the mode of their utilization for navigational purposes are completely unknown. At least, however, some spatial
structures in the seemingly structureless atmosphere have been found which
theoretically could be used by birds for home-finding. Also, a preliminary hypothetical framework has been suggested which could be used by other investigators as a first guideline for continuing endeavours or as a stimulus to develop competing ideas.
Apart from a number of still unsettled details concerning pigeon homing
and an extension to homing experiments with wild birds, the most fundamental questions to be tackled in the future require interdisciplinary research
teams comprising not only behavioural biologists but also neurobiologists
and/or, most importantly, experts in atmospheric chemistry. This necessity as
well as methodological and financial problems will certainly complicate and
retard further progress. Otherwise I would be confident that we are on a good
way towards a solution to the homing problem. I hope that some enthusiastic
young scientists will be able to surmount the technical and organizational barriers.
Appendix:
Notes on Selected Particulars
These Notes were written for critical expert readers and also for those among the non-experts
who may have missed one or another result or aspect about which they had previously read or
heard and which appears to disagree with the picture drawn in the main text (for such disagreeing views in the literature, see, e.g., Gould 1982; Wiltschko and Wiltschko 1982, 1998, 1999b;
Schmidt-Koenig 1987, 1991, 2001; Wiltschko 1991, 1996). Readers who still remain unsatisfied by
these Notes may consult my earlier comments on various issues treated in the literature (Wallraff
1980b, 1983,1988a,b, 1990a,1991b, 2000b, 2001,2003; Wallraff and Sinsch 1988; Wallraff et al. 1982,
1994, 1999) and also contributions by Papi (e.g. 1986, 1989, 1991, 1995) and Able (1996). Some of
the Notes merely append some details which may be necessary to know for a critical assessment
of particular findings. The following remarks refer to sections of the main text as indicated.
Section 2.1:
Initial Orientation
For statistical analysis of orientation data including considerable angular dispersion, the conventional methods of linear statistics (arithmetic mean, standard deviation, etc.) are inapplicable,
because their results depend on the point on which the endless circle is numerically cut. A variety of methods applicable to circular statistics have been described (Batschelet 1981; Fisher 1993;
Mardia and Jupp 2001). It is necessary to consider which of the computational algorithms should
be used for what kind of question to be answered.For a strong test of whether pigeons are actively
oriented towards home and/or towards a commonly preferred compass direction (Sect. 3.1.1),for
instance, a second-order test (cf. Fig. 2.4) should be applied in which mean vectors or mean bearings obtained at a number of symmetrically arranged release sites serve as statistical units (and
not individual bearings of pigeons). Namely, as results at a given site are usually similar in repeated releases, an increase in sample size per site would not change the overall outcome very
much. With only few sites, apparent preference for the homeward (or any other) direction, even if
significantly exhibited by large numbers of birds, could still be a product of chance resulting
from the selection of sites. On the other hand, if the effect of an experimental treatment is investigated in a two-sample test comparing experimental birds with control birds, also first-order tests
are applicable using single bearings as statistical units, because this comparison concerns individuals flying independently under corresponding conditions. If within-release numbers per
treatment group are equal or similar, bearings observed at different sites can be pooled (for
pseudo-pooling considering unequal sample sizes see Wallraff 1979). Second-order tests applied also in this case are more rigorous (due to considerably reduced sample size n), but not
obligatory to demonstrate potential statistical significance.
Section 3.1:
Homing of Inexperienced Pigeons
Wiltschko and Wiltschko (e.g. 1982, 1998, 1999b; Wiltschko 1991) show moderately clear data
from which they conclude that between 2 and 4 months of age accuracy of initial homeward orientation degrades. In a hardly comprehensible way they incorporate this conclusion in their hypothesis of a change in navigational strategy during ontogeny (Wiltschko and Wiltschko 1985)
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Appendix
from transitional early path integration to later map-based navigation (Sects. 4.2 and 6.3.4).
However, releases of younger and older pigeons were conducted separately at variable sites, days
and years, so that the many other sources of variability (Sects. 3.1.1, 3.3, 3.7.1 and 3.7.3) were not
equilibrated and could have produced pseudo-correlations. In simultaneous releases of different
age groups no significant differences in homeward orientation were found; on average, the older
birds were somewhat better; also 2- to 3-month-old pigeons apparently used map-based navigation using site-specific cues (Gagliardo et al. 1988; Wallraff and Sinsch 1988).
Section 3.1.3:
Limited Significance of Exercise Flights at Home
The importance of flying for the development of the navigational system is amply documented.
Pigeons will consider as their home the location where they flew around during the first months
of life (Wiltschko and Wiltschko 1998,p. 176).As support for this view, the authors refer to barely
relevant data on returns of passerine birds in their first spring after migration (e.g. Lhrl 1959),
whereas they persistently (e.g. Wiltschko and Wiltschko 1985, 1987, 1998, 1999b, 2000, 2003a) refrain from considering the really relevant findings obtained with aviary pigeons which amply
document the contrary (e.g. Kramer and von Saint Paul 1954; Kramer 1959a,b; Wallraff 1966a,
1970a, 1979; Gagliardo et al. 2001a). Ignoring these findings, Wiltschko and Wiltschko speculate
that extensive flights around the loft are necessary to unravel the alignment of (imaginary) gradients which are thought to constitute the grid of a navigational map (Sect. 4.2.2). It is obvious,
however, that pigeons displaced over some 100300 km are able as well as motivated to orient towards a home location where they could never fly around (cf. Figs. 3.7 and 3.8).
Section 3.2.1:
Number and Range of Previous Homing Flights
Figures 3.9 and 3.10: results from 11 series of four experiments, each series conducted at a symmetrically arranged quartet of release sites as shown in Fig. 3.4 (two series 30, 60, 120 and 180 km
cardinal, one series 150 km cardinal, one series 30 and 90 km diagonal). Experienced birds released at the four sites in each series had an equal or similar history of previous releases which
were, as good as possible, balanced in spatial symmetry. They were more than 1 year old (up to
8 years) and had been returned from at least 12 previous releases including distances of at least
90 km, partly from more than 20 or 30 releases including displacement distances of 180 km and
more. In most cases, the pigeons had not been released previously at the current release site; in
some cases, they had been released there once or twice before, but then they had afterwards made
a number of other flights from other directions. Inexperienced pigeons were usually 46 months
old. Groups of approximately 10 birds of either type were transported together and released singly in alternating sequence. In a second-order comparison, mean homeward components as well
as return rates per release (n = 44) were significantly different with P < 0.0001 (Wilcoxon
matched pairs test). In 31 of the 44 releases the homeward component was greater and in 40 releases homing success was better in the experienced birds (three times the return rate was 100%
in both groups, in one case the inexperienced birds were better).
Section 3.2.3:
Familiarity with the Release Site or Area
In a series of 14 releases at various sites 120200 km distant from home, Grter and Wiltschko
(1990) found improved initial homeward orientation and homing performance in a second release conducted at the same site several days later. If most of the pigeons had not homed from any
other site in the meantime (not reported), no distinction is possible between effects of local/directional training (Sect. 3.2.2) and effects of memorized site-specific signals. When a longer period of time, filled with homing flights from other sites, separated the second from the first release (experiments at 40 km distance), no consistent improvement was observed. The authors
Appendix
197
attribute the different outcomes to the different degrees of familiarity with navigational signals
inside and outside of a training range, but owing to the apparently different procedures the real
causal connection remains unsettled.
Section 3.7.1:
Home-Related and Home-Independent Components of Initial Orientation
Neglecting lots of empirical data, Wiltschko and Wiltschko (e.g. 1998, 1999b; Wiltschko 1993) accept only issue 1 in this section (navigational error resulting from non-fit between map and nature), deny issue 2 (PCD) as a phenomenon per se and ignore issues 3 and 4 (deflections by topographical features and preceding homing flights). A monofactorial explanation of release-site
biases is certainly most parsimonious (Wiltschko and Wiltschko 1998, p. 173), but does not
cope with reality and easily leads to inadequate interpretations (or to perplexing detection of a
paradox: Wiltschko and Wiltschko 1989). If homeward orientation (together with its possible
misguidance) is experimentally eliminated, a PCD persists and even tends to be strengthened
(Figs. 7.2, 7.3 and 7.8; see also Sect. 10.1.2). Also attracting or repelling influences of landscape
features persist in anosmic pigeons which fail to orient homewards (Sect. 8.2; Wallraff 1994b).
Section 5.2:
Linkage Between Sun Compass and Map
In their modern concept schema of ontogenetic development of the avian navigational system,
Wiltschko and Wiltschko (2003a, Fig. 7) failed to draw any connection between sun compass and
map. However, this connection is essential to make a map-and-compass system work. Without
this linkage, results as shown in Fig. 5.6 could not be explained because the map of coordinates in
Fig. 5.5 would be free to rotate in relation to the sun (cf. Wallraff 2004).
Section 6.1.1:
Magnets or Coils During Flight
The conditions in the magnet experiments described by Keeton (1971, 1972) and Moore (1988),
respectively, may have been different according to two aspects. First, even pigeons without magnets are not always well-oriented towards home under overcast skies. Their performances seem
to depend on previous exercise flights at home under such conditions (Sect. 5.3). The mean
homeward component of control pigeons in the later experiments listed by Moore is considerably
shorter than in Keetons earlier experiments. Against weak controls, experimental birds cannot
be very much weaker. Second, the experiments were conducted with experienced pigeons and
mostly at short distances (< 35 km). Thus, many birds may have been familiar with the area (even
if not with the particular site of release) and hence may have been able to low-level homeward
orientation on the basis of landscape knowledge without using a compass (Chap. 8). Since details
about the pigeons individual homing histories were not reported, it is impossible to draw firm
conclusions. Also, kind and attachment of magnets may have contributed to the diversity of results. According to Ioal (2000), the bar magnets and their attachment on the pigeons backs, as
used by Keeton (1971, 1972; Moore 1988), were less effective than the magnets attached to head
and wings, as used by Wallraff et al. (1986a) and Ioal (1984, 2000).
In a somewhat perplexing publication, Harada (2002) concludes that the lagena (an otolith in
the inner ear with so far unclear functions), which is reported to contain magnetic elements (Fe,
Mn and Zn), acts as a magnetic sensor in avian navigation. The conclusion is based on experiments with ten homing pigeons close to whose ear canal, in the neighbourhood of the lagena, tiny
magnets were (unilaterally?) attached inside the temporal bone (two different treatments not
clearly described). In another eight pigeons unilateral nerve-sectioning of the temporal bone
was performed, later described as the lagenal nerve was cut electrically. All the pigeons, together with untreated controls, had been trained >20 times over a radius of 30 km from home
before the experiments. The test releases were conducted at a site only 15 km from home. Initial
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Appendix
bearings were not observed, but homing performances were drastically reduced in the birds
treated in one or the other way. They either did not home at all or returned very slowly after
110 days, whereas all control birds returned within 30 min after release. This result is puzzling
for two reasons. First, over such a short distance in a familiar area, pigeons should be able to home
quickly by means of visual landmark orientation without requiring true navigation (Sect. 8.3).
Second, pigeons with their cochleae and lagenae bilaterally removed, were not in any way impaired in initial orientation and homing after displacements over more than 150 km into unfamiliar areas (Wallraff 1972). Harada (2002) quotes these experiments but does not comment on
the discrepancy of the results. It is unclear what actually made his experimental birds so weak in
short-distance homing within familiar terrain. Time intervals between surgery and release were
not reported. It is remarkable that, after removal of the magnets in returned pigeons, four out of
five birds still required more than 7 h to home again from the same site. Thus, it seems likely that
some unidentified non-specific disturbance was produced rather than specific interference with
navigation.
Section 6.1.2:
Magnetic Fields Before Release
Item 6: Long-term magnetic fields at the home site (Wiltschko et al. 1983). Young pigeons were
raised in a small loft and aviary in which the horizontal component of the magnetic field was
shifted clockwise against the natural one by 65 or 120,respectively.While sitting in the aviary,the
birds experienced both the natural sun and the shifted magnetic field. Exercise flights were allowed only under complete overcast when the sun was obscured by clouds. Upon displacement
and release at varying sites under sun, these birds deviated by about half the shift angle clockwise
from control birds in the first release and counterclockwise in a second release after they had experienced the sun together with the natural magnetic field. It is difficult to interpret these results,
because potentially interfering additional variables were not sufficiently controlled. Most importantly, during that phase of life the pigeons have to establish their map-and-compass system by
associating wind directions with olfactory sensations (Sects. 7.5 and 7.9.2). If inside the aviary
wind direction was fully accessible at all,it was natural with respect to the sun and shifted with respect to magnetic north, whereas outside the aviary winds were natural with respect to magnetic
north and could not be related to the (invisible) sun. Moreover, different winds may have been
differently frequent under sun and under overcast. It is unclear in what way the pigeons might
have interpreted the varying conditions to which they had been exposed before release.
Section 6.3.1:
The Interference with Stress Issue
It is undisputed that artificial magnetic fields as well as complete darkness during transport can,
under certain circumstances, disturb subsequent initial orientation of pigeons (e.g. Wiltschko and
Wiltschko 1985; Papi 1995). Two hypotheses have been presented to explain this analogy: (1) both
treatments interfere with magnetoreception, which is thought to depend on the presence of light
(e.g. Wiltschko and Wiltschko 1985; Sects. 6.3.4 and 6.3.5); and (2) magnetic treatments inhibit
compensation of stress caused by usual handling conditions; darkness induces additional stress
(Fig. 6.5). The finding that application of a tranquillizer suspends the effect of either treatment
(Fig. 6.4) is compatible with the second hypothesis, but hardly with the first.
Section 6.3.3:
The Magnetic Map Issue
The strongest objections (among others) against Walkers (1998, 1999) model are the following
(Wallraff 1999; Reilly 2002): (1) The model proposes measurement of extremely subtle changes of
magnetic intensity over distances in the 1- to 2-km range, together with precise directions over
which such changes occur, and even comparison of these measurements with such data memo-
Appendix
199
rized from the home site. Not only is the necessary precision of such data acquisition and processing outside of any conceivable capability of an organism but also are the subtle variations to
be measured inside the range of natural spatio-temporal noise of geomagnetic parameters.
(2) The model proposes in-flight scanning of magnetic parameters over some distance in varying
directions around the home loft as well as near the release site. However, pigeons do not need free
flight at home to develop their homing ability (Sect. 3.1.3). Upon release, they are homeward oriented already after 20 s of flight (Fig. 2.4) and thus need no sampling of data while circling around
in order to determine the direction of a magnetic gradient.
Section 7.3.1:
Spatial Separation of Sites of Air Smelling and Release
In the area around Tbingen, Germany, many experiments were conducted in which five to seven
pigeons were kept and transported in large airbags, i.e. in 750-l plastic containers, mostly filled
with air collected at different sites. In the first series of such experiments (Kiepenheuer 1985) the
results corresponded with those achieved elsewhere with a somewhat different method (see
Fig. 7.12). In later airbag experiments, including several modifications, the results were less
clear-cut and, therefore, not very elucidating (Kiepenheuer 1986; Ganzhorn 1990; Ganzhorn and
Burkhardt 1991). Three reasons made the conditions unclear: (1) Most of the releases were conducted at very short distances from the loft, ranging between 9 and 25 km, and with pigeons that
were more or less experienced in homing and hence could have been more or less familiar with
the area on a visual basis (Sect. 8.3). The familiar landscape most probably provided the other
release-site information in Kiepenheuers (1986) experiments and contributed also to variable
blurring of other results. (2) Also owing to the short distances, great differences between air samples collected at different sites could hardly be expected (Sect. 7.4.1). (3) The sort of anaesthetic
used (Gingicain), and its application at varying times before release certainly made the depth of
nasal anaesthesia in the critical moment variable (cf. Wallraff 1988b). Nevertheless, if no additional undetected shortcomings were involved, it is difficult to see possible causes for the difference between control and experimental pigeons in the pooled data of experiment 2 in Ganzhorn
and Burkhardt (1991), which, however, did not compare different olfactory conditions.
Section 7.5.1:
Shielding, Deflecting and Reversing Winds
On the basis of several series of deflector loft experiments, Phillips and Waldvogel (1982, 1988),
Waldvogel and Phillips (1982,1991) and Waldvogel et al. (1988) questioned the decisive role of olfactory inputs and speculated, instead, on assumed effects of disturbing light cues reflected from
the transparent deflector panels. Experiments and associated discussions (overloaded with
sharp-witted interpretations) were more confusing than elucidating. Distances of displacement
were quite short and the pigeons were often more or less (when more? when less?) familiar with
visual landscape features. Information on the birds experiences has been poorly communicated,
so that it is impossible to assess which factors might have determined which results to what degree. Nevertheless, the overall results of the 48 tests obtained over 7 years with pigeons permanently living in aviaries with either original or optically altered deflector panels agree quite well
with the results described in Section 7.5.1 and Fig. 7.22 (Waldvogel and Phillips 1991). For a detailed discussion of the short-term (ca. 1-week) deflector experiments see Wallraff (1990a,p. 108)
and for comments on deflector cages additionally equipped with anti-cheating slats (Waldvogel
and Phillips 1991) see Wallraff (2001, p. 38). These experiments hardly contribute any important
aspect to the debate on olfactory navigation, in no case to the question of whether pigeons make
use of olfactory signals to navigate at all (the experiments interfered neither with olfactory sensing nor with the composition of the inhaled air).
200
Appendix
Section 7.6.4:
Assumed Dependence on Geographical Peculiarities
Some details on measurements of airborne substances as shown in Fig. 7.35: Data acquisition as
well as gas-chromatographic and statistical analyses were performed as described by Wallraff and
Andreae (2000). From the data collected in the course of that study, included here were those 48
samples that had been obtained at the 24 sites located in a northsouth corridor 50 km eastwest
of the meridian of the Andechs loft (= 50150 km east of the longitude of Wrzburg). In addition,
during each of two transalpine trips, two samples were taken at each of 10 sites, one on the southward trip and one on the northward, in total 40 samples at 20 different sites. Thirty compounds observed in 95% of the 88 chromatograms,combined to 435 possible pairs,included up-and-down ratio gradients similar to those shown in Fig. 7.35C on a random probability level of P = 0.01.
Section 7.8:
Arguments Against Olfactory Navigation And Replies
In some publications olfactory navigation is rashly laid aside with a few words, so that no basis for
a substantial discussion is left. As a recent example, I mention Goulds (2004) sketchy gallop
through the wide field of animal navigation. The role of odours in pigeon homing is polished off
there by four facts that are thought to argue against it:
1. The rearing technique used in the studies does not permit the normal imprinting flight.
Gould probably refers to aviary experiments (Sect. 7.5), which are, however, used in only a
minor part of the relevant studies (for other parts see Sects. 7.17.3). Moreover, what is the
normal imprinting flight? Exercise flights at home are evidently not necessary to establish a
home site toward which pigeons are able to navigate (Sect. 3.1.3).
2. The scatter of pigeons using olfactory cues should increase with the distance of the release
site from the loft, but exactly the opposite is observed. No reason for the expectation should
increase is given and no reference for observation of the opposite. For more thorough considerations of the problem, see Section 7.6.5.
3. These results have been very difficult to repeat (see Sect. 7.8.3).
4. Pigeons have a notoriously poor olfactory sensitivity (see Sect. 7.7).
Such cursory statements and verdicts disclosing little familiarity with the relevant literature are
not very helpful for a serious scientific discourse. No publication is cited.
Section 7.8.2:
Potential Non-Specific Side Effects
There is one construct of possibilities against which it is difficult to argue. One might hypothesize
that pigeons need to smell a specific compound, or set of compounds, as a precondition to be either able or motivated to activate a non-olfactory mechanism of position determination. This
mechanism would then operate only while the birds can smell natural open-field air
(Figs. 7.117.13); it would operate slowly (Fig. 7.9c); and air within forests would contain little of
the specific compound(s) (Fig. 7.10). The physical basis of the assumed non-olfactory mechanism would remain enigmatic; magnetism could be excluded (Sects. 6.1 and 6.3.3). Its apparent
dependence on simultaneous olfactory perception of particular airborne substance(s) would
burden the mechanism with an additional enigmatic property. I do not consider this game of
thoughts heuristically promising.
Appendix
201
Section 7.8.3:
Inconsistent Results
Some details to Fig. 7.37. Criteria for inclusion of published experiments in the figure were: double method of nostril occlusion and nasal anaesthesia as described; analogous release means
available from four sites fairly symmetrically arranged around home (a few further releases
omitted); release sites (not always surrounding areas) unfamiliar to the birds. Conditions at the
home loft and preceding homing experience of pigeons were uniform within a quartet of releases, but varying between quartets (see original publications). Homing speeds could not be included for all releases either because less than half of the birds returned on the day of release or
because data were not reported.
On the whole, even the results of declared opponents of olfactory navigation or of their
co-workers, respectively, involve much more data supporting rather than questioning the important role of olfactory inputs, especially if not only initial bearings but also returns to the loft are
considered (quartets #34 and #922; Tbingen in Fig. 7.4). With temporally acting nasal anaesthesia homing speeds were consistently reduced (Fig. 7.37), and with long-term acting zinc sulphate also homing rates (Schlund 1992; Fig. 7.4B). Also in initial orientation, the differences between control and experimental pigeons in most series are statistically significant. This holds
true even in some series for which they were declared insignificant by the authors who applied
only second-order statistics using release means as units (e.g. Wiltschko et al. 1987b,d; quartets
#913 and #1922 in Fig. 7.37). In two-sample tests comparing different treatments, there is no
reason to disregard individual bearings and thus to reduce original sample sizes by a factor of 10
or more (ten or more bearings per release; see this Appendix under Sect. 2.1).
As long as we do not know all the environmental circumstances that influence pigeon homing, it
makes little sense to speculate in detail why analogous methods of incomplete olfactory deprivation
produced different results on different days, at different sites, in different countries or in pigeons
differently exposed to winds at their home loft. Such circumstances might involve, for instance,
varying overall levels of concentrations of decisive airborne substances; occlusion of nostrils might
then lead to sub-threshold levels in some cases but not in others.Or pigeons accustomed to sub-optimal air and wind conditions at a ground-level loft among buildings and trees might cope with reduced smelling better than pigeons from a wind-exposed loft for which the impairment makes a
larger difference. Or the relative strengths of competing directional tendencies [towards home, towards a PCD, towards or away from topographical features (Sects. 3.7.1 and 8.2)] might play some
role. With a strong PCD, for instance, a weakened homeward impulse might become invisible,
whereas with no or weak deflecting tendencies it may remain perceptible (cf.Wallraff 1990a,p. 104).
Only if the birds definitely cannot smell, before and upon release, should any homeward component at symmetrically arranged sites in reliably unfamiliar areas be eliminated in all circumstances.
Section 7.9.1:
Experimental Evidence of Olfactory Navigation
Merely relative comparison between olfactorily treated and untreated pigeon groups reveals often considerable similarities rather than differences. For instance, if the mean bearings of the experimental birds per site or release, as shown in Figs. 7.2A and 7.8A, B, are drawn as deviations
from the simultaneously obtained mean bearing of the controls, the controlward components of
the resulting mean vectors (0.63 and 0.49) are similar to the homeward components of the control
birds in those figures (0.46 and 0.56). In both cases, the median absolute angular difference between the two group means is only 38, significantly (P < 0.0001) lower than 90, the mean angle
expected in the case of complete independence. For other such comparisons see Kiepenheuer et
al. (1993), Fig. 7 in Wallraff (1990a) and Fig. 4 in Wallraff (2001). In order to study the role of
olfaction, the levels of homeward orientation must be compared.
202
Appendix
Section 8.1.3:
Homing with Olfaction and Vision Impaired
An experimental series by Benvenuti and Fiaschi (1983), preferentially quoted to demonstrate that
there must be a third (unknown) factor involved, beyond olfaction and vision (Schmidt-Koenig
1991; Wiltschko 1996), differs in its design so much from commonly performed pigeon releases,
that its results cannot be generalized (for detailed discussion see Streng and Wallraff 1992; Wallraff
2001).
Section 8.1.4:
Interrelations Between Landscape and Sun Compass
Full-scale deflections observed in clock-shift releases after more than 50 consecutive releases of
the same pigeons at always the same site (Fller et al.1983; Wiltschko et al.1994) are not comparable with clock-shift effects at varying sites. After so many repetitions of flying the same compass
course, the clock-shifted birds apparently continued to fly the entrained compass direction without feeling noteworthily disturbed by the wrong landscape features they were crossing. In contrast, pigeons released at a non-predictable site must first know where they are, and for this purpose they have to pay attention to the topographical geometry (and/or to olfactory signals). For
more detailed considerations about clock shifts and landscape, see Wallraff (1991a) and Wallraff
et al. (1994, 1999).
Wiltschko et al. (1994) and Wiltschko and Wiltschko (2001) tend to attribute the reduction of
deflections caused by clock shift with increasing age of the pigeons to an effect of age itself and
possibly of associated general homing experience rather than to increasing visual familiarity
with the area. Actually it cannot be excluded that, with increasing age and experience, the relative
hierarchical weights of sun compass and magnetic compass gradually shift from complete dominance of the first to a growing participation of the latter. It is not immediately obvious, however,
why such a shift should occur as long as the birds are not confronted with conflicting directional
signals (which they do not meet during normal homing flights). To clarify the question, it would
be necessary to release experienced veteran pigeons, clock-shifted for the first time in their lives,
at reliably unfamiliar sites far beyond the radius so far covered by homing flights. If they show reduced clock-shift deflections also at these far-distant sites, an influence of age and/or general experience per se may be stated and, consequently, an influence of familiar landscape features in
other experiments doubted. In the above-mentioned studies the release areas (not only sites!)
were not clearly defined as familiar or unfamiliar and it has not been reported how often the
same birds had previously been released with shifted clocks (which might have made them
sceptical about trusting the sun in a case of conflict).
Section 9.2:
The Hippocampus and the Olfactory Map and
Section 9.4:
Roles of Other Brain Regions and Hemispheric Lateralization
A more puzzling rather than elucidating result has been reported by Gagliardo et al. (2001c). Aviary pigeons hippocampal-lesioned on the right-hand side at an early age were homeward-oriented at unfamiliar release sites, but left-side lesioned pigeons were not. Less clear differences between the two groups were found in homing performance. One may ask, therefore, whether
hemispheric lateralization might have concerned motivation to orient homewards rather than
navigational ability. On this assumption, it may also appear conceivable that impairment of olfaction-based homing after bilateral hippocampal lesion in aviary pigeons, but not in free-flying
birds (Ioal et al. 2000b), was due to weakened motivation: Confinement in an aviary plus lack of
hippocampal functions might have reduced alertness to associative learning.
Appendix
203
Section 12.1:
Environmental Cues Involved in Home-Finding Processes
It has frequently been emphasized that pigeons may use a multiplicity of more or less redundant
cues for home-finding and may decide on one or the other depending on varying circumstances
under which different signals provide differently clear positional information (e.g. Keeton 1974a;
Wiltschko et al. 1987d; Schmidt-Koenig and Ganzhorn 1991; Ganzhorn 1992; Walcott 1996). It is
true that (1) position determination is not always and everywhere equally clear-cut and correct and
that (2) the pigeons orientation behaviour is extremely variable depending on a variety of parameters such as geographic region, relation between home site and release site, season, day of release,
previous sequence of homing flights of the individual birds, etc. However, observation (2) does not
necessarily imply that performance (1) is based on varying kinds of input signals.If olfactory information is differently distinct, for instance, home-independent directional preferences (Sects. 3.7.1
and 8.2) may determine the birds behaviour to a lesser or higher degree. I do not see more redundancies than those among sun compass and magnetic compass on the one hand, and among olfactory map and visual landscape map (in familiar areas) on the other hand. Anyhow, the number of
potential navigational cues is very limited unless we suspend any limitation by proposing usage of
hypothetical unknown physical cues yet to be detected. The multiple cue argument is often used as
an alibi if appropriate interpretation of (seemingly) inconsistent results proves difficult.
Section 12.3.2:
Problems of Visual Landscape Orientation and
Section 12.3.3:
Problems of Olfactory Navigation
Guilford et al. (2004) and Roberts et al. (2004) applied elaborate mathematical algorithms to analyse GPS-tracked homing routes of pigeons released at very familiar sites only 24 km distant
from their loft. Within this small range around home, their approach has hardly more than descriptive value and probably no or little relevance to navigation.After gaining height upon release
and some circling and looking around, the birds were most likely always aware where they were
with respect to the nearby home. I consider it at least unsettled that a high state of entropy (tortuous flight), as correlated with some topographical features underneath (e.g. human settlements
as opposed to little structured empty areas), indicates navigational uncertainty. Like schoolboys
returning from school, the pigeons had probably no problem of way-finding, but were not in a
hurry to home and hence readily followed distracting influences by this and that, as for instance
by other pigeons sitting on houses or flying around. In real homing experiments over longer distances, however, and in areas with different degrees of familiarity, track analyses of that kind may
well be expected to allow not only recognition of distracting properties of villages, forests, etc.
(Sect. 8.2), but also navigationally meaningful interpretations concerning mechanisms of homefinding using visual and/or olfactory signals.
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Subject Index
accuracy of orientation 32, 4246, 4951,

56, 58, 188, 195
aerial view 27, 29, 160, 167, 188, 190
aerosol particles 98
age 18, 27, 30, 50, 76, 80, 85, 112, 166, 189,
195f, 202
airbag experiment 101, 199
airplane tracking 13
albatrosses 4, 170, 176178, 191
altitude of homing flight 13
anaesthesia (local / nasal) 93f, 96104, 106,
140142, 150154, 158, 163, 192, 199201
Anas platyrhynchos 172f
annual cycle / periodicity 3841, 44, 110
anosmia / anosmic 8896, 104106, 109,
120, 142, 153f, 164f, 174f, 179, 186, 188, 190,
197
Apus apus 173, 179
artificial
air 96, 107, 108
light-dark regime 60f, 65
magnetic field 6977, 79, 82, 185, 198
odour 118
photoperiod 38
wind 114, 117f
atmospheric
cues / odours / signals 93, 98, 105, 131,
133, 166, 167
gradients 128, 135, 146, 179181
compounds / substances / trace gases 57,
95, 120, 123, 125, 134, 136, 138, 144, 146,
173, 176, 178, 188, 192, 193
aviary experiments / pigeons 2729, 36, 47,
87, 111120, 122, 131, 144f, 165, 185f, 196,
198200, 202
bearing-and-distance programme (see also
time-and-direction p. 179) 1
benzaldehyde experiment 118f, 135
bi-coordinate
grid 57, 184
navigation 121, 178
brain 4f, 116, 138, 140, 145, 161f, 166f, 175,
186, 188, 192
Brazil 92
breeding
area / range / site / territory
179f
burrow 176
cycle 38
1, 4, 47, 174f,
cage (arena / maze) experiment 16, 73, 83,

154, 161, 168, 171f, 188
chemical atmosphere 120136, 191
chemosignal (see also odour) 57f, 98, 111,
121, 137f, 140, 143f, 151, 156, 176, 186f, 191f
circadian clock / rhythm 5966, 156f, 164,
173
climate 51, 110, 131
clock shift 37, 5968, 71f, 80, 116, 122, 145,
155157, 159161, 164f, 168, 172f, 188, 202
cochlea 58, 198
cognitive map 5, 54, 144f, 161f, 165, 188
Columba livia 3
Columba palumbus 3
compass 1f, 4f, 64, 66, 68, 69, 77, 171, 173, 197
alignment 65, 115, 128, 156
course 1, 37, 78, 84, 156, 162, 164, 180, 202
direction 1, 4, 13, 34f, 59, 68, 8285, 117,
144f, 180, 202
magnetic c. 53, 62, 6668, 7174, 77, 80,
82, 85, 155, 160, 171, 184, 187, 202f
orientation 16, 59, 69, 71, 78, 171, 173,
181, 184f
reference 143, 186, 187
sun c. 18, 37, 53, 5968, 71, 77, 80, 85, 122,
155157, 160, 165, 171173, 184, 186188,
197, 202f
coordinates 4, 47, 49, 5557, 65f, 69, 81, 121,
143, 146, 183185, 197
Coriolis force 57, 69, 184
darkness (effect of) 75f, 79f, 198
day-to-day fluctuations / variations 42, 111
dead reckoning, see path integration
deflector aviary / loft 114120, 142, 144, 160,
199
detour experiment 101, 103105
Diomedea exulans 177
Diomedea immutabilis 170
226
Subject Index
direction recorder 13
directional
reference 1, 53, 57f, 64, 143f, 187, 192
training 16, 36f, 47, 161, 166, 168, 196
distance (of displacement / from home)
8, 2325, 2932, 4248, 50, 53, 56, 82, 90f,
94, 103, 107111, 122, 133138, 143, 146,
152154, 157, 162, 169171, 174176, 179,
182f, 187, 192, 197200, 203
diurnal rhythm 41
ducks 172
electromagnetic sferics 58
England 37, 92, 110, 184
Erithacus rubecula 171
exercise flights 27, 29, 36, 65, 111f, 114, 117,
149, 196198, 200
experience 5, 17, 21, 26, 2937, 42, 49f, 54f,
67, 72, 8991, 114, 146, 155, 160f, 188f,
196199, 202
familiar / familiarity (area, site, landmarks,
landscape) 2, 4, 26f, 29, 36f, 46, 48, 55, 57,
67, 9096, 109, 116, 134, 142, 145f, 149167,
175f, 179181, 186188, 190, 196199, 202f
familiar-area map 5, 54, 160, 162, 167, 178,
188
filters / filtration / filtered air 96108, 150f,
153, 158
first-flight pigeon (definition) 8
flight path / route 3, 9, 1113, 44, 103, 129,
131, 189
frosted lenses /spectacles 152154
gas chromatography / chromatogram 97,
123125, 127f, 192, 200
geographical variability 43f, 47, 51, 66, 93,
107, 131, 133, 203
geomagnetism / geomagnetic see magnetism / magnetic
Germany 15, 21, 30, 32, 4042, 44, 47, 51, 67,
74, 76, 92, 96, 98, 103, 107f, 110, 123, 131,
133135, 141
goal-oriented / -related navigation 14, 143,
173186
GPS-tracking 13f, 26, 189f, 203
gradient 5, 27, 5558, 6466, 69, 81f, 120137,
143146, 176180, 185, 191, 196, 199
direction 56, 66, 68, 122, 129
field 55, 179
map 5, 55, 57, 64f, 120, 128, 144146, 160,
165, 179, 186
gravity 58
grid map (see also gradient map) 55, 145
guidance by adults 1
gulls 78, 169171
helicopter tracking 13
Helmholtz coils 69, 72, 7476
hippocampus 159, 161168, 186, 202
homeward
component (definition) 8
navigation 35f, 47, 49f, 105, 114
homing 4
ability 24, 199
drive 2f, 23, 25, 50
mechanism (see also navigational mechanism) 2, 64, 68, 74, 80, 145, 167, 179,
188
path / route 2, 1214, 46, 103f, 129, 131,
152f, 157, 190, 203
performance 11f, 26, 32f, 3644, 7376,
88, 89, 93, 96, 104, 107, 110, 136, 149f, 154,
162, 165f, 185, 196, 198, 202
rate (see also return rate) 11, 23, 70, 74,
77, 90, 93, 201
speed 3f, 11, 25, 30, 38, 46, 51, 58, 70f, 74,
77, 80f, 93, 141f, 154, 163f, 171
success 18, 23, 25, 30, 32, 89f, 166, 174,
196
image vision 152154
inertial navigation 54, 183
infrasound 57f, 185
interdisciplinary research 189, 193
Italy 21, 30, 40f, 44, 47, 51, 7376, 92f, 96, 98,
103, 107f, 110, 112, 133, 135, 141
lagena 58, 197f
landmarks / landscape 4, 7, 29, 37, 44f, 49,
5458, 68, 71, 92, 116, 142f, 145, 149168,
175f, 178, 181, 183, 185188, 190, 192,
197199, 202f
Laysan albatros 170
lateralization (hemispheric) 167, 202
learning 18, 27, 36, 47f, 50, 51, 54, 56, 6568,
111, 117, 121, 133, 145, 155, 160f, 165, 176,
179, 189, 202
lobsters 82
long-distance homing / navigation / orientation 16, 23, 27, 29, 133135, 137, 143, 170,
176, 179183, 191
magnetic
activity 58, 78
anomaly 77f, 80f, 185
compass 53, 62, 6668, 71f, 74, 77, 80,
8285, 155, 160, 171, 184, 187, 202f
domains 77
field 1, 54, 57f, 64, 66, 6985, 139, 143f,
183186, 192, 198
fluctuations / variations 76, 78, 80
gradient 57, 82, 199
Subject Index
inclination 57, 69, 73, 8184, 183
inclination compass 73f, 83f
information 74, 178
input 81, 85, 181, 183
intensity 57, 69, 8185, 183, 198
irregularities 69, 77, 80
latitude 69, 82, 85, 181, 184
map 5, 49, 78, 8185, 139, 183185, 191,
198
navigation 81, 178
noise 80, 81
oscillations 71, 75f, 79, 83
polar compass 83
pole 83, 183
sense 8285
sensor 197
signal 57, 71, 78, 81f
treatment 73, 74, 7678, 80f, 198
vector 53, 65, 72, 73, 83, 191
magnetism 58, 69, 79, 81, 139, 185, 187, 200
magnetite 77, 83f
magnetoreception 69, 78, 8385, 140, 198
mallard 172f
map 4f, 19, 36, 48f, 54, 64, 66, 69, 72, 74, 77,
82, 117, 120, 144146, 164, 167, 184, 196f
cognitive 5, 54, 144f, 161f, 165, 188
familiar-area 5, 54, 160, 162, 167, 178, 188
gradient 5, 55, 57, 64f, 120, 128, 144146,
160, 165, 179, 186
grid (see also gradient map) 55, 145
magnetic 5, 49, 78, 8185, 139, 183185,
191, 198
mosaic 5, 54f, 134, 178f, 188
navigational 5, 27, 37, 145, 173, 185, 196
olfactory 5, 18, 87, 115f, 119f, 139,
144146, 160, 165f, 186, 203
radial 146
topographical 5, 54f, 57, 145f, 160, 162,
165, 178, 186, 188
map and compass 6466, 68, 77, 145, 165,
184, 188, 197f
meteorological conditions / variations 58,
111, 189
migration 13, 171, 179182, 196
migratory
birds 3, 47, 73, 80, 83, 171
navigation 179
orientation 1f, 172
restlessness 16, 171f
mobile loft 48
Morocco 92
mosaic map 5, 54f, 134, 178f, 188
motions (recording of) 54, 82
motivation 3, 17, 2326, 32, 35, 38, 46, 50f,
79f, 90, 117, 140, 171, 189, 202
multiple cues 203
227
nasal tubes 96
navigation (see also navigational) 4
bi-coordinate 121, 178
goal-oriented / related 14, 143, 171,
173176
homeward 35f, 47, 49f, 105, 114
inertial 54, 183
map-based 196
olfactory 57, 87147, 152, 154, 162, 166f,
176, 179182, 188, 191f, 199201
sun 57, 59, 64, 81, 184
true 4f, 29, 48, 111, 188, 198
unicoordinate 82
navigational
ability / capability / capacity 3, 23, 29, 46,
50, 111, 171, 176f, 179, 202
cue / signal / information 3, 32, 35, 40,
46f, 49, 57, 80, 87, 110f, 133, 142, 159, 187,
197, 203
error 19, 22, 48, 197
map 5, 27, 37, 145, 173, 185, 196
mechanism (see also homing mechanism) 3, 7, 17, 49f, 73, 81, 85, 181
performance 4, 25f, 131, 178
process 4, 36, 49, 64, 79, 161
range 46f, 107110, 179
system 5, 29, 45, 81, 134f, 145, 147, 176,
179, 181, 196f
newts 82
noise 26, 42, 46, 51, 56, 80f, 123, 129, 131,
134, 137, 199
non-olfactory
behaviour 88
cues 110, 175
homing 149151, 153155, 159f
mechanism 200
side-effects 92f, 98, 140
nonsense orientation (see also preferred
compass direction) 49, 171f, 190
non-specific effects 58, 139, 154, 198,
200
North America / USA 42, 67, 73, 92, 94,
110, 141
occlusion of nostrils (see also plugging) 9395, 106f, 109, 113, 141f, 200f
odour (see also chemosignal) 76, 87147,
152, 157, 166f, 174176, 186, 200
olfaction 57, 87147, 151, 153f, 157, 161163,
176, 179, 181, 186, 201f
olfactory 74, 150, 158, 198
bulb 166, 175
condition 101, 135, 181, 199
cues / signals 71, 81, 87147, 149, 151,
157, 159f, 166, 173, 175, 178f, 181, 186,
188, 190, 192, 199203
228
Subject Index
deprivation 58, 8796, 110, 120, 139143,

149, 153, 156, 174, 191, 201
gradient 131, 180
gradient hypothesis 120123, 144
information 87147, 151f, 203
input 93, 95, 105, 109, 135, 160, 174, 179,
181, 191, 199, 201
map 5, 18, 87, 115f, 119f, 139, 144146,
160, 165f, 186, 203
homing mechanism 188
mosaic hypothesis 133135
navigation 57, 87147, 152, 154, 162, 166f,
176, 179182, 188, 191f, 199201
nerve section 8892, 106, 109, 173f, 186
orientation 181, 190
sensitivity 93, 95, 136f, 141, 175, 200
site simulation 101104
system 138, 140, 176, 178, 181, 191
ontogeny / ontogenetic 18, 50, 68, 161, 190,
195, 197
opioid system 76, 79, 85
outward journey 47, 54, 70, 74, 76, 82, 104,
107, 109, 133, 185
overcast sky 37, 53, 6668, 7073, 85, 160,
171f, 197f
path integration 54, 74, 82, 85, 111, 183, 185,
196
PCD see preferred compass direction
(PCD)
petrels 169171, 176, 178
pigeon
loft 2, 4, 14, 23, 26, 29, 47f, 87
race 3, 46, 58, 78, 183
strain / stock 3, 30, 44, 76, 80, 85, 110
piloting 4f, 37, 48, 157, 188
piriform cortex 162, 166f
plugging of nostrils (see also occlusion) 88f,
9294, 116, 173f
position determination 46f, 49, 5360, 64,
6972, 78, 81f, 85, 121f, 139, 142f, 145, 157,
173, 181, 184f, 200, 203
position recorder 13
positional information 11, 16, 46, 57f, 68,
81, 93, 98100, 104f, 107, 109, 120, 123,
125, 134f, 139, 149, 155, 159f, 176, 178,
186, 202
preferred compass direction (PCD) 19, 21,
29f, 33, 35f, 42f, 49, 75, 80, 90, 101, 104,
116, 119f, 143, 158, 171173, 189f, 195, 197,
201
radial map 146
radio telemetry 13, 153
ratio gradient 121137, 144, 146, 176, 178,
191, 200
recovery sites / recoveries 2, 14f, 16, 18, 23,

25, 2729, 42, 47, 74, 90f, 106f, 109, 112f,
129132
release (definition) 7
release-site bias 18, 21, 37, 48f, 77, 149
return rate (see also homing rate) 12, 24,
2830, 32, 38, 42, 46, 89f, 92, 107, 171, 174,
196
Riparia riparia 172
robin (European) 171
rock pigeon 3
route recorder 103
route reversal 74, 82
salmon 87, 138, 143
sand martin 172f
satellite telemetry 13f, 169, 177, 191
seabirds 176179, 181f
season 2, 4, 17, 3841, 47, 51, 63, 191, 203
second-order analysis 10, 195
selection 3, 30, 50f, 189
semicircular canals 183
sense of direction 183
sense of geographical position 183
sensitive period 47, 117
sferics 58
shearwaters 170, 176
short-distance homing / navigation 23, 90,
110, 122, 133135, 137, 152, 167, 182, 198
side effect 88, 9294, 98, 116, 139f, 200
signal processing 136, 138, 146, 192
simulated position 82, 101104, 135, 160
smell, sense of (see also olfaction) 92f, 136,
139, 143, 175f, 179, 182, 186, 192
starling 4, 169172, 174176, 179
statistical analysis of orientation data 711,
195
stellar sky / compass 1, 172
stopover sites 181
stress 70, 71, 76, 79f, 85, 151, 172f, 198
Sturnus vulgaris 170f
sun 1, 37, 53f, 5772, 78, 80f, 143f, 155157,
164, 168, 171f, 183f, 187f, 191f, 198
altitude 60, 62, 184
azimuth 53, 6064, 68, 123, 155f, 171, 187f
compass 18, 37, 53, 5968, 71, 77, 80, 85,
122, 155157, 160, 165, 171173, 184,
186188, 197, 202f
navigation 81
navigation hypothesis 57, 59, 64, 184
spots 58, 78
swallows 169172
swift 169, 173175, 179
target area 29, 47, 146, 180
telencephalon 161f
Subject Index
temperature 3840, 43f, 110, 121
temporal fluctuations / variations 26, 3842,
44, 48f, 51, 78, 80f, 95, 107, 110f, 123, 126,
129, 137, 159
terns 169172
time-and-direction programme (see also
bearing-and-distance p. 1) 179
topographical
characters / features 4, 44f, 145, 152, 155,
157159, 197, 201, 203
geometry 202
influence 13, 157
map 5, 54f, 57, 145f, 160, 162, 165, 178,
186, 188
topography 13, 49, 189
trigeminal nerve 83
true navigation 4f, 29, 48, 111, 188, 198
turtles 82, 84
unfamiliar (area / site) 1f, 29, 46, 48, 50, 55,
57, 71, 90, 96, 110f, 114, 120, 134, 142146,
150, 152, 155f, 159, 163167, 171173, 179,
198, 201f
unicoordinate navigation 82
USA see North America
229
vanishing bearing (definition) 7

vanishing interval 8f
visual (cues / signals / features / landmarks
etc.) 37, 48, 54, 57f, 68, 71, 88, 90, 94,
113f, 116, 142, 149167, 175, 178192,
198f, 202f
flow 54
impairment / limitation 112f, 117, 152154
landscape map 162, 203
VOC (volatile organic compound) 123132,
135f
wandering albatros 177
weather 17, 66f, 121, 123, 127, 131, 179, 191
wind 29, 66, 87, 111123, 126131, 133138,
143145, 176, 186f, 190192, 198f, 201
deflection 114120, 142, 144, 160, 199
drift 1, 13, 180
reversal 116f, 144
winter quarter 1, 180
wintering area / grounds / region /
territory 1, 47, 179181
zinc sulphate experiment
201
92, 154, 164, 191,

Avian Navigation - Pigeon Homing As A Paradigm

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Avian Navigation - Pigeon Homing As A Paradigm

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Hans G.

Dr. Hans G. Wallraff

ISBN 3-540-22385-1 Springer-Verlag Berlin Heidelberg New York

Bird migration includes homing performances in almost global dimensions.

ical findings. Presentation of these findings, discussion of their significance

4 Potential Input Signals Exploitable for Home-Finding ............................. 53

7.5 Varying Home Site Conditions in Aviary Experiments ................... 111

10.1.2 Compass Orientation and Preferred Compass Directions ... 171

1.2 Why Investigate Domestic Homing Pigeons?

oculars over approximately 2 km and large enough to carry the weight of a

sense, an individual bird is not an independent unit. Moreover, the pigeons of a

1.3 On Terms and Definitions

Observation Data Used

2 Observation Data Used to Investigate Pigeon Homing

ularly when a number of symmetrically arranged releases are combined

2.1 Initial Orientation

2 Observation Data Used to Investigate Pigeon Homing

terval represents an indirect measure of the decidedness of initial orientation.

2.2 Homing Performance

2 Observation Data Used to Investigate Pigeon Homing

2.3 Homing Routes

1. Observation from airplane or helicopter. Using a slowly flying airplane

2 Observation Data Used to Investigate Pigeon Homing

n Fig. 2.6. Examples of homing flight routes of pigeons

2 Observation Data Used to Investigate Pigeon Homing

Basic Features of Pigeon Homing

In a first approach, before coming to possible explanations, it is necessary to

3 Basic Features of Pigeon Homing

3.1 Homing of Inexperienced Pigeons

3 Basic Features of Pigeon Homing

3.1 Homing of Inexperienced Pigeons

3 Basic Features of Pigeon Homing

3.1 Homing of Inexperienced Pigeons

3 Basic Features of Pigeon Homing

3.1 Homing of Inexperienced Pigeons

3 Basic Features of Pigeon Homing

3.1 Homing of Inexperienced Pigeons

n Fig. 3.7. Vanishing bearings (cir-

cular diagrams) and recoveries of

3 Basic Features of Pigeon Homing

3.2 Experience Gained by Homing Flights

ers landed soon afterwards nearby or disappeared low-flying behind trees or

3 Basic Features of Pigeon Homing

3.2 Experience Gained by Homing Flights

3 Basic Features of Pigeon Homing

3.2 Experience Gained by Homing Flights

3 Basic Features of Pigeon Homing

3.2 Experience Gained by Homing Flights

3 Basic Features of Pigeon Homing

3.2 Experience Gained by Homing Flights

3 Basic Features of Pigeon Homing

3.3 Temporal Variability

3 Basic Features of Pigeon Homing

3.3 Temporal Variability

Corresponding annual cycles in homing performance were also found at two

3 Basic Features of Pigeon Homing

3.4 Accuracy of Homeward Orientation

3 Basic Features of Pigeon Homing

cally distributed in 2031 km around six home sites at different latitudes

3.4 Accuracy of Homeward Orientation

3 Basic Features of Pigeon Homing

3.6 Multiple and Mobile Home Sites

3 Basic Features of Pigeon Homing