Orientation in Birds

341 Pages·1991·7.758 MB·English

by  K. Schmidt-Koenig, J. U. Ganzhorn, R. Ranvaud (auth.), Prof. P. Berthold (eds.)

Orientation in Birds
Edited by P. Berthold
With a foreword by Riidiger Wehner
Birkhauser Verlag
Basel . Boston . Berlin
Editor's address:
Prof. P. Berthold
Max Planck Institut fUr Verhaltensphysiologie
Vogelwarte Radolfzell
Schloss Miiggingen
D-7760 Radolfzell/Germany
CIP-Titelaufnahme der Deutschen Bibliothek
Orientation in birds/ed. by P. Berthold.-Basel; Boston; Berlin: Birkhauser, 1991
(Experientia: Supplementum; Vol. 60)
ISBN-13: 978-3-0348-7210-2 e-ISBN-13: 978-3-0348-7208-9
DOl: 10.1007/978-3-0348-7208-9
NE: Berthold, Peter [Hrsg.J; Experientia/Supplementum
Library of Congress Cataloging-in-Publication Data
Orientation in birds/edited by P. Berthold.
p. cm.-(Experientia. Supplementum: v. 60)
Includes bibliographical references and index.
ISBN-13: 978-3-0348-7210-2 (U.S.: alk. paper)
I. Birds-Orientation. 2. Bird navigation. 3. Birds-Migration.
I. Berthold. P. (Peter). 1939- . II. Series.
QL698.85.075 1991
598.252'5-dc20
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Contents
Foreword. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. VII
K. Schmidt-Koenig, J. U. Ganzhorn and R. Ranvaud
The sun compass ........................................ .
W. Wiltschko and R. Wiltschko
Magnetic orientation and celestial cues in migratory orientation. . 16
C. Walcott
Magnetic maps in pigeons. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
F. Papi
Olfactory navigation. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
P. Berthold
Spatiotemporal programmes and genetics of orientation. . . . . . . . 86
R. C. Beason and P. Semm
Neuroethological aspects of avian orientation. . . . . . . . . . . . . . . .. 106
H. G. Wallraff
Conceptual approaches to avian navigation systems. . . . . . . . . .. 128
K. P. Able
The development of migratory orientation mechanisms. . . . . . . .. 166
S. B. Terrill
Evolutionary aspects of orientation and migration in birds . . . .. 180
T. Alerstam
Ecological causes and consequences of bird orientation. . . . . . .. 202
W. J. Richardson
Wind and orientation of migrating birds: a review. . . . . . . . . . . .. 226
R. Wiltschko
The role of experience in avian navigation and homing. . . . . . .. 250
A. J. Helbig
Experimental and analytical techniques used in bird orientation
research. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 270
E. NOirak and P. Berthold
Satellite tracking: a new method in orientation research. . . . . . .. 307
P. Berthold
Orientation in birds: A final consideration. . . . . . . . . . . . . . . . . . .. 322
Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 328
Foreword
If it is true that science proceeds from a romantic through a scientific to
a technological stage, then research on bird orientation is certainly on
its move from its first to its second grade, and recent developments in
radiotelemetry and satellite tracking of migrating birds might already
indicate the advent of the third stage. At this juncture, Orientation in
Birds is a timely account. Even though the study of animal migration in
general, and bird navigation in particular, has produced a literature of
impressive proportions, the threads provided by the plethora of research
papers, review articles and symposiums volumes have not yet been
knitted into a theoretical fabric.
This is partly due to our still incomplete understanding of fundamen
tal topics in avian navigation. The answer to the most intriguing
question of how a bird displaced to "unknown" territory finds its way
back home is as obscure now as it was a few decades ago. Whether and
how birds solve this problem by using far ranging grid-maps or more
local familiar-area maps, as has been proposed off and on, is still a
matter of heated debates. These debates frequently center around
provocative hypotheses - let alone the question about the physical
(topographic, magnetic, infrasonic, olfactory) parameters which might
constitute such maps. Furthermore, the number, structure and function
of the various (sunlight, skylight or magnetic) compasses by which the
bird determines and maintains its steering courses, are far from being
understood. In this context, it might come as a real surprise that even in
what is considered to be a well-established, almost classic field of
research - the study of the bird's sun-compass system - nearly nothing
is known about how the birds use this compass during their migratory
journeys.
In this state of affairs Orientation in Birds does not only present a
concise and lucid up-to-date review of the field, but takes an encourag
ingly fresh look at long standing puzzles like the ones mentioned above.
Certainly, there is no real need for collecting and presenting more data
within the conceptual framework entertained so far. This framework
was based on the ideal of perfect design. However, as natural selection
is principally opportunistic, it favours navigational systems not for their
mathematical elegance and aesthetic appeal, but simply because they
work. For example, during their evolutionary history birds using the
sun as a compass might have encountered neither the need nor the
opportunity to acquire a complete set of solar ephemeris functions
comprising all seasonal and geographical variations of the sun's daily
movements. Instead, for the avian navigator it might well suffice to refer
to particular aspects of the all-inclusive problem envisaged by the
human investigator. Consequently, what looks like awe-inspiring com-
plexity in the navigational behaviour the bird displays might, in the end,
turn out to be the interplay of a number of quite simple subroutines -
even though "in the end" may be some time off.
In learning about such subroutines, natural selection must be our
guide. There is no a priori way to decide what the peculiarities of a
navigational problem are to which the bird responds, and what compu
tational system it has evolved to cope with these particulars. By shifting
our attention to the ecological and physiological constraints encoun
tered by the migrating bird we might finally be able to raise the proper
questions rather than try to solve problems ill-conceived for the avian
navigator. It is exactly here that the potential and strength of Orienta
tion in Birds come into play. Rather than contributing a mere summary
of what has already been documented, it extends the scope of research
on bird navigation to a wide range of fields including genetics and
evolution, ecology and physiology, and thus bears witness to the fact
that research on bird navigation can no longer be a self-contained
discipline. Investigations in the fields mentioned above must be estab
lished as equal partners of orientation research in the study of bird
migration. Beyond that, the message running as a thread through all the
chapters of the book is, at least in my eyes, "back to basics" - back, in
the first place, to the study of the spatial and temporal fine structure of
individual migratory routes, back to an understanding of the genetic
blueprint of the bird's vectorial route of migration, back to recordings
of the environmental conditions the migrant encounters on its way, and
back, finally, to basic studies on the bioenergetic machinery and neural
hardware mediating orientation behaviour.
Seen in this light, Orientation in Birds is a prolegomenon rather than
a finished treatise, but it is exactly in this open-minded heuristic ap
proach that Berthold's multi-author review excels among the vast litera
ture on avian migration and homing.
Zurich, August 1991 Rudiger Wehner
Orientation in Birds, ed, by P. Berthold
© 1991 Birkhauser Verlag Basel/Swi1zerland
The Compass
SUD
K. Schmidt-Koenig1,2, J, U, Ganzhorn2 and R, Ranvaud3
1 Department of Zoology, Duke University, Durham (North Carolina, USA)
2 Abt. Verhaltensphysiologie, Universitiit Tiibingen, D-7400 Tiibingen
(Federal Republic of Germany)
31nstituto de Ciencias Biomi>dicas, USP, Sao Paulo (SP, Brasil)
Summary. The sun compass was discovered by G. Kramer in caged birds showing migratory
restlessness. Subsequent experiments with caged birds employing directional training and
clock shifts, carried out by Hoffman and Schmidt-Koenig, showed that the sun azimuth is
used, and the sun altitude ignored. In the laboratory, McDonald found the accuracy to be
± J ~ ± 5c. According to Hoffmann and Schmidt-Koenig, caged birds trained at medium
northern latitudes were able to allow for the sun's apparent movement north of the arctic
circle, but not in equatorial and trans-equatorial latitudes.
In homing experiments, and employing clock shifts, Schmidt-Koenig demonstrated that the
sun compass is used by homing pigeons during initial orientation. This finding is the principal
evidence for the existence of a map-and-compass navigational system. Pigeons living in
equatorial latitudes utilize the sun compass even under the extreme solar conditions of
equinox, achieving angular resolution of about 30 in homing experiments. According to
preliminary analyses, the homing pigeons' ephemerides are retarded by several weeks (Ran
vaud, Schmidt-Koenig, Ganzhorn et al.l.
Experiments with Caged Birds
Discovery of the sun compass
In the late 1940s, Gustav Kramer (1950) discovered that the migra
tory restlessness (Zugunruhe) of caged migrants was directed rather than
random. In a series of pioneering studies with Starlings (Sturnus vul
garis) Kramer (1951) was able to show that the sun is the directional
cue used by the birds during daytime. Following Santschi's mirror
experiment (Santschi, 1911), mirrors attached to the testing cage shifted
the bird's directionality as predicted (Figure 1). These initial investiga
tions were dependent upon the bird's migratory restlessness, which is
highly seasonal and restricted to certain hours of the day. In order to
overcome these restrictions, Kramer and St. Paul (1950) and Kramer
(1951) succeeded in training Starlings in a circular cage to look for food
in a certain compass direction. This technique made experimenting
independent of migration, season, and time of day.
With only sky and sun visible to the birds in the training apparatus,
the birds maintained their training direction in the course of the day.
2
~" J••
.:'
B
Figure I. Gustav Kramer's (195 I) mirror experiment with a Starling showing migratory
restlessness in a circular arena with 6 windows. Each dot inside the arena symbolizes one
average direction observed in a IO-s interval. A Control condition, B with mirrors attached
clockwise and C attached counter-clockwise. Arrows indicate the direction of incoming
sunlight.
This result strongly supported the conclusion of the preceding mirror
experiment: Kramer had discovered the sun compass in birds. The
Starlings allowed for the sun's apparent movement. For this task the
bird needs to keep track of the time of day, i.e. it must possess a
chronometer, also known as an internal or biological clock, and it must
have knowledge of the sun's apparent movement.
The interaction between chronometry and the sun's apparent move
ment was particularly illuminated in experiments subsequently carried
out by Hoffmann (1954) who demonstrated that the Starling's biologi
cal clock could be reset experimentally. The number of hours of
resetting - 6 h counterclockwise in Hoffmann's original experiment -
resulted in a predictable shift of the bird's directional orientation in the
training apparatus. The prediction is roughly 15° of directional shift for
every hour of time or clock shift. The sun's rate of change of azimuth
is on the average 15°/h.
The bird's clock is easily reset if the bird is confined in a light-proof
room with artificial light regimes, e.g. shifted 6 h clockwise (i.e. the day
for the pigeons starts 6 h before the beginning of the natural day) or
counterclockwise (i.e. daylight for the pigeons starts 6 h after the