J Comp Physiol A (2015) 201:983–1001
DOI 10.1007/s00359-015-1026-3
ORIGINAL PAPER
Releases of surgically deafened homing pigeons indicate
that aural cues play a significant role in their navigational system
Jonathan T. Hagstrum1   · Geoffrey A. Manley2 
Received: 12 February 2015 / Revised: 11 June 2015 / Accepted: 14 June 2015 / Published online: 2 July 2015
© Springer-Verlag Berlin Heidelberg (outside the USA) 2015
Abstract  Experienced homing pigeons with extirpated
cochleae and lagenae were released from six sites in
upstate New York and western Pennsylvania on 17 days
between 1973 and 1975 by William T. Keeton and his co-
workers at Cornell University. The previously unpublished
data indicate that departure directions of the operated birds
were significantly different from those of sham-operated
control birds (314 total), indicating that aural cues play an
important part in the pigeon’s navigational system. Moreo-
ver, propagation modeling of infrasonic waves using mete-
orological data for the release days supports the possibil-
ity that control birds used infrasonic signals to determine
their homeward direction. Local acoustic ‘shadow’ zones,
therefore, could have caused initial disorientation of con-
trol birds at release sites where they were normally well
oriented. Experimental birds plausibly employed an alter-
nate ‘route-reversal’ strategy to return home perhaps using
their ocular-based magnetic compass. We suggest, based
on Keeton’s results from another site of long-term disori-
entation, that experienced pigeons depend predominantly
on infrasonic cues for initial orientation, and that surgical
removal of their aural sense compelled them to switch to a
secondary navigational strategy.
Electronic supplementary material  The online version of this
article (doi:10.1007/s00359-015-1026-3) contains supplementary
material, which is available to authorized users.
* Jonathan T. Hagstrum
jhag@usgs.gov
1 waves called infrasound (below the range of human hearing)
US Geological Survey, 345 Middlefield Road, Menlo Park,
CA 94025, USA
2 atmosphere (Bedard and Georges 2000).
Cluster of Excellence “Hearing4all”, Research Center
Neurosensory Science and Department of Neuroscience,
Oldenburg University, Oldenburg, Germany
Keywords  Cochlea · Lagena · Columba livia ·
Infrasound · Avian navigation
Introduction
Birds navigate over distances of hundreds to thousands of
km with pinpoint accuracy, and this phenomenal ability is
thought to involve a two-step process (Kramer 1959). Solar,
magnetic, and stellar compasses (the ‘compass’ step) have
been shown to orient avian flight (Keeton 1974; Wiltschko
and Wiltschko 2009), but how birds first determine the direc-
tion to their destination (the ‘map’ step) has been an endur-
ing mystery. Bi-coordinate ‘maps’, based on two or more
gradients in geomagnetic field parameters (Wiltschko and
Wiltschko 2009) or in ratios of atmospheric odors (Wallraff
2013), have been proposed, but each hypothesis has signifi-
cant difficulties (e.g., Wallraff 2005; Schmidt-Koenig 1987)
and no consensus has yet been reached on which naviga-
tional cue, or combination of cues, is used. Moreover, results
from experimental releases of homing pigeons (Columba
livia) by varying groups in different countries often appear
inconsistent, contradictory, or confusing (Walcott 1996), and
a number of observed behaviors remain unexplained (Wall-
raff 2001). Although such confusion could indicate an under-
lying navigational system of extreme complexity and vari-
ability, it might also mean that one or more key elements has
either been overlooked or misunderstood. One avian sense
that has received relatively little attention in navigational
terms is hearing (Quine 1982), and low-frequency acoustic
are known to travel undiminished for thousands of km in the
In the late 1960s, William von Arx, then at the Woods
Hole Oceanographic Institution, suggested to Donald R.
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984
Griffin that infrasonic signals might provide navigational
cues to flying birds (Griffin 1969). Subsequent laboratory
experiments demonstrated that homing pigeons are capable
of detecting atmospheric infrasound at frequencies below
10 Hz (Yodlowski et al. 1977), and, using a quieter test
chamber, at frequencies as low as 0.05 Hz (Kreithen and
Quine 1979). Similar experiments also showed that birds
could possibly localize infrasonic sources by evaluating
Doppler shifts during flight (Quine and Kreithen 1981; see
Appendix: Box 1 for details). Pigeons tested after bilat-
eral extirpation of their cochleae and lagenae were deaf
to infrasound (~2 Hz at 114 dB SPL), indicating that their
low-frequency acoustic receptors lie within the inner ear
(Yodlowski et al. 1977). Moreover, Schermuly and Klinke
(1990) demonstrated that, within the pigeon cochlea, affer-
ent fibers innervating hair cells near the apical end of the
basilar papilla respond to infrasound at levels comparable
to the behavioral thresholds determined by Kreithen and
Quine (1979). There is no doubt that pigeons, and possi-
bly many other birds, are sensitive to infrasound and thus
possess an additional sense of potential use in spatial
orientation.
Experimental releases of young pigeons confined to
aviaries since fledging had shown the importance of unob-
structed interactions with a nonvisual “dynamic factor
of the atmosphere” in establishing their homing capabili-
ties (Kramer and von St. Paul 1954; Kramer 1959; Wall-
raff 1966, 1970). Birds kept in shielded aviaries, with only
the sky visible (Kramer 1959; Wallraff 1979), or in ones
designed to deflect local winds (Baldaccini et al. 1975;
Kiepenheuer 1978; Waldvogel et al. 1978) were disori-
ented or flew off from release sites with correspondingly
deflected departure bearings, respectively. Wallraff (1972)
conducted several tests to determine whether the “atmos-
pheric factor” might be infrasound, but found that deafened
pigeons, released with intact controls at two sites in Ger-
many (N  = 52 total), apparently suffered no detriment in
their homing performance. Although both cohorts released
at one site 151 km SSW of the loft departed homeward
in nonrandom directions (Rayleigh test: N  = 13 in both
cases; operated, r  = 0.78, P < 0.001; controls, r  = 0.66,
P  = 0.002; Batschelet 1981), deafened birds released at
the second site 159 km NNE of the loft were homeward
oriented (N  = 13, r  = 0.63, P  = 0.004) while the paired
control birds departed randomly (N  = 13, r  = 0.30,
P = 0.316). No explanation was given for this finding, indi-
cating that, although both operated and control birds could
return to the loft, the true nature of their homing process
had not been fully understood.
In his investigation of release-site biases (consistent dif-
ferences between mean departure and homeward bearings)
at Castor Hill in upstate NY, Keeton (1973) also noted the
atypical random departure (N = 11, r = 0.068, P = 0.954)
13
J Comp Physiol A (2015) 201:983–1001
Castor Hill
LAKE
NY
ONTARIO Pulaski
PA
30 km
Weedsport
Auburn
Cornell loft
Jersey Hill
Caroline I
Orwell, PA
Fig. 1  Digital elevation model (DEM) of the Finger Lakes region of
upstate New York, and western Pennsylvania, USA (see inset), show-
ing locations of the Cornell loft (42.4360°N, 76.4319°W) just E of
Ithaca, NY, near the southern tip of Lake Cayuga, and pigeon release
sites at Auburn; Caroline I; Castor Hill; Orwell, PA; Pulaski; and
Weedsport (Table 1). Jersey Hill is a site of perennial disorientation
for Cornell pigeons (see text)
of untreated birds on 22 July 1970, but he, too, was unable
to recognize anything unusual about this particular day.
Sites of long-term disorientation have also been of keen
interest to those studying avian navigation (e.g., Kramer
1959; see Brown 1971), and one such well-studied site is
Jersey Hill located 120 km west of the Cornell loft (see
Brown 1971; Keeton 1974; Fig. 1). Although forewarned
by pigeon breeders in the Ithaca area to avoid releasing
his birds at Jersey Hill, over the next 19 years Keeton and
his co-workers, in attempting to understand the cause(s)
of disorientation, released thousands of pigeons at Jersey
Hill. Untreated Cornell birds typically departed randomly
from Jersey Hill (N = 984, r = 0.007, P > 0.900; Walcott
and Brown 1989) with high attrition rates (only ~10 %
returned), but pigeons from other lofts, located at various
directions and distances from Jersey Hill, were able to ori-
ent and return home from there on the same days the Cor-
nell birds were lost (Walcott and Brown 1989).
Recently, Hagstrum (2013) provided an explanation for
these observations based on acoustic propagation modeling,
using recorded atmospheric profiles, of infrasonic signals
(~0.2 Hz) possibly generated by coupling of microseismic
J Comp Physiol A (2015) 201:983–1001 985
waves, which pigeons might use as navigational cues (see
Box 1). Jersey Hill is situated within an acoustic ‘shadow’
zone relative to the Cornell loft, and homeward infrasonic
signals were mostly unavailable at Jersey Hill and the sur-
rounding region over the 19 years of the Keeton group’s
pigeon releases. Other lofts in upstate NY would have had
their own shadow zones of similar geometry that would not
have included the Jersey Hill location. Moreover, no evi-
dence has been presented showing that other navigational
cues were absent or somehow confusing at Jersey Hill,
indicating that infrasound is perhaps the predominant long-
range cue used by homing pigeons. However, if this is true,
why did deafening pigeons (Wallraff 1972) have no appar-
ent effect on their homing ability? And, if, on the other
hand, birds do have a multimodal navigational system (e.g.,
Walcott 2005), why did not intact Cornell pigeons released
at Jersey Hill resort to one or more alternate cues to orient
themselves?
William T. Keeton and his co-workers at Cornell Uni-
versity also conducted experimental releases with surgi-
cally deafened pigeons: releases of 314 operated and con-
trol birds were made on 17 days at 6 sites in upstate NY
and western PA (Fig. 1). Working with Keeton’s group, one
of us (GAM, then at McGill University, Montreal) surgi-
cally deafened a large number of pigeons in the early 1970s
that were subsequently released with similarly experienced
cohorts of sham-operated birds. Here again, a number of
the sham-operated (control) birds departed in directions
statistically indistinguishable from random, suggesting that
such results are caused by some recurring phenomenon.
These unpublished data, made available to all researchers
after Keeton’s premature death in 1980 (Brown et al. 1984),
are analyzed herein using standard statistical techniques
(Batschelet 1981) and acoustic ray-trace modeling (Jones
et al. 1986).
Methods
Cornell pigeon releases
Keeton’s published and unpublished Cornell pigeon release
data (Brown et al. 1984), collected between 1967 and 1980,
are presently available from C. Walcott at Cornell Univer-
sity (Department of Neurobiology and Behavior, Ithaca,
NY, USA; cw38@cornell.edu), and a program which sorts
and displays the database (‘Keeton’) is available from T.
Larkin (Abstract Tools, Brooktondale, NY, USA; tsl1@cor-
nell.edu).
After weaning at approximately 4 weeks, young Cornell
pigeons were allowed to develop their flying abilities during
flights around the loft area. Once deemed proficient, train-
ing flights were made for birds of a single pen or test group
by releasing them as a single flock at sites in all directions
and at increasing distances from the loft. The basic flock
tosses were at distances of up to ~24 km and were fol-
lowed by two or three short-distance, single-flight releases
to eliminate any birds unable to home independently. Two
or three training flights under overcast conditions were also
included in the training program. All of the birds utilized in
the experiments of this study were thus considered experi-
enced birds, and it can be generally assumed that a bird’s
homing experience increases with age. Young birds (YB;
Table 1) were those tested in their natal year, prior to sexual
maturity; yearlings (YL) were tested in their second year
after becoming reproductively active; and old birds (OB)
were aged two or more years.
The experimental pigeons (E) were operated on employ-
ing a fast technique developed by Schwarzkopff (1949) to
bilaterally extirpate their cochleae and lagenae with Equith-
esin used as an anesthetic. The eardrum was resected and
the columella footplate was drawn out of the oval win-
dow: a fine hook constructed from a tungsten electrode
was inserted just inside the oval window to gain purchase
on the cochlear limbus, surrounding the basilar membrane,
and was slowly withdrawn. This procedure reliably pulled
out the entire cochlear duct including the lagena. In each
case, the removed object was examined under a microscope
to confirm successful cochlear extraction. All birds recov-
ered and showed no evidence of vestibular disturbances,
with one exception, which showed balance problems and
was removed from its cohort. In addition, a second group
of experimental birds (ME) had their columellae bisected,
leaving the cochleae intact, thus eliminating direct sound
conduction between the tympanum and oval window. Con-
trol birds (C) were randomly selected pen mates of the
experimental birds that were similarly anesthetized, but not
subjected to surgical interventions. In three pilot releases
from Auburn, Caroline I, and Weedsport (Fig. 1; Table 1),
conducted during the autumn of 1973, the surgically deaf-
ened cohort flew well in all cases and showed no locomo-
tory or general behavioral deficits. A larger number of birds
underwent surgery in the spring of 1974, followed by eight
releases of these birds later that year also from Auburn and
Weedsport. Five more releases were carried out in 1975
from Castor Hill, Orwell, PA, and Pulaski. ME birds were
also released at Castor Hill and Pulaski in 1975.
Between 1973 and 1975, as the importance of outward-
journey influences became known, transportation of birds
to the release sites was progressively shifted from closed
vehicles to open pick-up trucks. Upon release, birds were
hand tossed in random directions, and two observers fol-
lowed each with 10 × 50 binoculars until it vanished
from view. A compass bearing for the vanishing point was
recorded to the nearest 5°, and the time interval between
toss and vanishing was timed with a stopwatch. During
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986 J Comp Physiol A (2015) 201:983–1001

Table 1  Cornell releases of cochlea-extirpated and sham-operated homing pigeons

Release number Date N/N0 φ r hc Δ ΔCE Age Treatment Site Exp Sky

Auburn (43.0216°N, 76.5714°W) 66 km N of loft; homeward bearing = 170°

 1067 11/20/73 6/9 168 0.936 *** 0.935 *** 2 YB C N S
 1067 8/12 149 0.882 *** 0.823 ** 21 19 YB E N S
 1073 05/01/74 3/6 120 0.967§ 0.622§ 50 YL E N S
 1076 05/02/74 4/5 96 0.676§ 0.186§ 74 YL C N S
 1076 9/14 160 0.585 ** 0.576 ** 10 64 YL E N S
†
 1082 05/10/74 13/13 260 0.137 0.000† 90 YL C N O
 1082 10/10 145 0.580 * 0.526 ** 25 115 YL E N O
 n/a 06/25/74 1/1 55 1.000§ −0.423§ 115 n/a C N O
 n/a 3/3 164 0.641§ 0.637§ 6 109 n/a E N O
‡
 C-all 24/28 163 0.252 0.250‡ 7b
 E-all 33/45 157 0.673 *** 0.656 *** 13b 6b
Caroline I (42.3647°N, 76.2705°W) 15 km SE of loft; homeward bearing = 301°
 1059 10/31/73 6/8 204 0.942 *** −0.115† 97 YB C N S
‡
 1059 5/5 241 0.738 0.369† 60 37 YB E N S
Castor Hill (43.6492°N, 75.8422°W) 143 km NE of loft; homeward bearing = 200°
 1385 06/02/75 6/7 191 0.700 * 0.691 * 9 OB C N S
 1385 7/9 220 0.898 *** 0.844 ** 20 OB CC F S
 1385 11/15 232 0.458‡ 0.388‡ 32 41 OB E N S
 1434 07/23/75 10/11 234 0.506‡ 0.419‡ 34 YL C n/a S
 1434 12/13 210 0.883 *** 0.870 *** 10 OB F0 F S
 1434 7/9 259 0.736 * 0.379‡ 59 25 YL E n/a S
 1434 11/11 270 0.864 *** 0.296‡ 70 YL ME N S
 C-all 35/40 214 0.727 *** 0.705 *** 14b
 E-all 29/35 257 0.654 *** 0.356 ** 57a 43a
Orwell, PA (41.8599°N, 76.2686°W) 66 km SSE of loft; homeward bearing = 348°
 1367 04/25/75 8/8 140 0.135† −0.119† 152 OB CB N O
 1367 8/10 220 0.483† −0.297† 128 80 OB EB N O
†
 1367 8/9 228 0.281 −0.141† 120 OB EM N O
‡
 1367 EB + EM 16/19 223 0.381 −0.219‡ 125
 1384 05/30/75 6/7 304 0.807 * 0.581‡ 44 OB CB N O
‡
 1384 7/8 312 0.511 0.413‡ 36 8 OB EB N O
 1384 7/8 208 0.237‡ −0.182† 140 OB EM N O
‡
 1384 EB + EM 12/14 247 0.229 −0.044† 101
‡
 C-all 14/15 309 0.273 0.212† 39b
‡
 E-all 13/16 269 0.374 0.071† 79b 40b
Pulaski (43.5657°N, 76.1801°W) 127 km NNE of loft; homeward bearing = 189°
 1501 09/30/75 11/11 198 0.817 *** 0.807 *** 9 OB C N S
‡
 1501 9/9 142 0.310 0.211† 47 56 OB E N S
‡
 1501 6/7 206 0.588 0.562‡ 17 OB ME N S
Weedsport (43.0965°N, 76.5469°W) 74 km N of loft; homeward bearing = 173°
 1063 11/8,12/73 11/15 182 0.792 *** 0.782 *** 9 YB C N O
 1063 12/15 211 0.484‡ 0.381‡ 38 29 YB E N O
 1184 07/26/74 7/7 197 0.610‡ 0.557‡ 24 YL C F/N O
 1184 8/8 203 0.969 *** 0.839 ** 30 YL CC F O
 1184 10/11 197 0.757 *** 0.692 ** 24 0 YL E N O
 1264 08/29/74 4/5 206 0.379§ 0.318§ 33 YL C N O
†
 1264 7/9 155 0.286 0.272† 18 51 YL EB N O

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J Comp Physiol A (2015) 201:983–1001 987

Table 1  continued
Release number Date N/N0 φ r hc Δ ΔCE Age Treatment Site Exp Sky
‡ †
 1264 5/6 216 0.396 0.290 43 YL EM N O
 1264 EB + EM 12/15 185 0.287‡ 0.281‡ 12
 1348 11/04/74 1/1 60 1.000§ −0.391§ 113 YL CB N O
 1348 1/1 180 1.000§ 0.993§ 7 YL CM N O
 1348 2/3 122 0.924§ 0.581§ 51 62 YL EB N O
 1348 3/3 180 0.700§ 0.695§ 7 YL EM N O
 1349 11/05/74 1/1 240 1.000§ 0.391§ 67 YL CB N O
 1349 1/1 335 1.000§ −0.951§ 162 95 YL EB N O
 1349 1/1 285 1.000§ −0.375§ 112 YL EM N O
 C-all 32/37 201 0.661 *** 0.584 *** 28a
 E-all 32/39 204 0.455 ** 0.390 ** 31b 3b

Release data were collected by WT Keeton and his co-workers at Cornell University and are mostly available in the Keeton database (Brown
et al. 1984; see also Fig. 3, Online Resource 1). C-all, E-all mean of all C and E bearings for each release site. n/a not available. N number
of vanishing bearings recorded and N0 number of birds released; φ mean vanishing bearing for the release, in degrees; r mean vector length
with critical level for the Rayleigh test (P) as follows, *** P < 0.001, ** P < 0.01, * P < 0.05, ‡ P > 0.05, † P > 0.500, and § not available; hc
homeward component with critical level (P) as above; Δ difference between φ and homeward bearings, for comparisons of the overall C and E
means, a they differ significantly, and b their 95 % confidence intervals overlap (Batschelet 1981); and ΔCE difference between C and E bird’s
mean directions, significance for overall mean comparisons as before. Age of birds: YB young birds, YL yearlings, OB old birds (see text). Treat-
ment: C sham-operated control birds, CB control birds with attached brass bars, CC second control cohort, E experimental birds, EB experi-
mental birds with brass bars, EM experimental birds with magnets, EB + EM experimental birds combined, F0 untreated OB (see text), and ME
experimental birds with bisected columellae. Training: homing training, and all birds are experienced except those unknown (n/a) for releases at
Auburn on 06/25/74. Site Exp site experience: experience of birds at site, N new to site, F familiar with site, and F/N familiar with and new to
site. Sky: weather conditions, S sunny, and O overcast

each release, an attempt was made to release the E and C
birds alternately, but this was not always strictly adhered
to (see Online Resource 1). A second cohort of C birds
(CC) was sometimes released as well (Table 1). In addi-
tion, a collaborator back at the loft recorded the time of
each bird’s return. Because these experiments were initially
designed to test the cochlea as magnetoreceptor, 10 of the
17 release days were overcast (Table 1) requiring the birds
to use their magnetic compass, which is subordinate to
their solar compass. Under sunny conditions, the Sun disk
was visible to the human eye throughout the test, even with
haze, but under overcast conditions the Sun disk was nei-
ther visible nor localizable.
In addition, to test if attached magnets had any effect on
the departure directions of the released birds under over-
cast, as part of three releases at Weedsport and Orwell,
PA, either small bar magnets or equivalent brass bars were
glued at the site to E, and sometimes C, birds dorsally at
the base of the neck using veterinary cement (see Keeton
1971). The EM and CM abbreviations (Table 1) thus indi-
cate E and C birds with magnets, and EB and CB indicate
E and C birds with brass bars. F0 birds were a subset of C
birds that tended to exhibit lower within-release scatter and
smaller release-mean biases at Castor Hill. Their offspring
retained this enhanced ability indicating that it was likely
an inherited trait (Walcott 2005).
The mean vanishing bearing (φ) for each treatment was
calculated by vector analysis, and the distribution of indi-
vidual bearings was evaluated by calculating the uniform
probability (P) under the Rayleigh test (Table 1). The bear-
ings of the two treatments in each experimental release
were compared under Watson’s U2 test. Comparisons of
mean bearings with κ > 2 and r > 0.750 were made with the
more restrictive Watson–Williams test. A nonparametric
test for dispersion (Batschelet 1981) was also used to com-
pare C and E data from the relatively few pigeon releases
made at Caroline I.
Atmospheric propagation modeling
To characterize the transmission of acoustic signals
between loft area and release site (Hagstrum 2013), and
thereby test the feasibility of an acoustic ‘shadow zone’
explanation for the random departures of control birds,
atmospheric propagation modeling incorporating daily
meteorological profiles was carried out using a Hamilto-
nian Acoustic Ray-tracing Program for the Atmosphere
(HARPA; Jones et al. 1986; available at http://cires.colo-
rado.edu/~mjones/raytracing/). Meteorological (rawin-
sonde) data from weather stations near Albany (WMO
#72518; 42.45°N, 73.48°W) and Buffalo, NY (WMO
#72528; 42.56°N, 78.44°W), for the time period covered

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988
by the Keeton database, are available online from the
NOAA/ESRL Radiosonde Database (http://www.esrl.
noaa.gov/raobs/). At the time of the pigeon releases, instru-
mented balloons were released daily from both stations at
00:00 and 12:00 GMT. The 12:00 GMT data (08:00 EDT)
were collected closest in time to the pigeon releases and
are used here. Results based on the Buffalo weather data,
however, are also preferentially used because the Buffalo
station is closer both to the loft (~165 km versus ~242 km)
and to most of the release sites.
HARPA computations using meteorological data from
either the Buffalo or Albany weather stations generally
show similar results, indicating that atmospheric conditions
across upstate NY were often uniform on a broad scale,
but were in some cases significantly different. The pro-
gram uses one set of rawinsonde profiles to represent the
entire atmosphere, and any local and/or short-term (<12 h)
changes in conditions between weather station and transit
path could not be determined for these historical data. Ide-
ally, for this type of modeling, detailed meteorological data
would be available along the acoustic transit paths between
transmitter and receiver sites, contemporaneous with the
pigeon releases.
HARPA traces three-dimensional paths of acoustic rays
(narrow beams representing a wave’s travel path that are
perpendicular to the wave front) through inhomogeneous
atmospheres and accounts for both vertical and horizon-
tal refraction, as well as advection; the propagation effects
of diffraction and scattering, however, are not determined
(Jones et al. 1986; Norris et al. 2010). Acoustic rays were
launched from a virtual transmitter located at the Cornell
loft (42.4360°N, 76.4319°W) along the direct azimuth to a
given receiver (release site) and at several azimuths either
clockwise (CW) or counterclockwise (CCW) within ±45°
of the direct bearing. At each azimuth, rays were launched
from 0° to 45° in elevation at 0.5° steps; rays were not
launched at higher elevations because the ray paths are not
in the flight domain.
Model atmospheres are created by HARPA from avail-
able rawinsonde profiles of temperature, wind speed, and
wind direction that usually extended to ~16 km above
upstate NY (http://www.esrl.noaa.gov/raobs/), and upper
atmospheric conditions to 120 km were approximated with
the HWM-07 wind (Hedin et al. 1996; Drob et al. 2008)
and NRLMSIS-00 temperature (Picone et al. 2002) models.
Topographic profiles between the virtual transmitter and
release site, defining the reflecting terrain’s height (Jones
1982), were also entered into the program. HARPA accepts
only terrain models having smooth surfaces with continu-
ous slope and curvature, and both weather and topographic
profiles were represented in HARPA by fitted sequences
of linear segments, joined with hyperbolic functions, that
effectively smoothed the datasets (Jones et al. 1986).
13
J Comp Physiol A (2015) 201:983–1001
Results
In order to establish the normal departure behavior over
the longer term of untreated Cornell birds at the six release
sites (Fig. 1; Table 1), data for C birds of similar age and
experience from other releases at these sites, under similar
weather conditions, were selected from the Keeton database
(Fig.  2). For sites with fewer releases, these criteria were
relaxed somewhat so that a larger number of data points
could be used to characterize the pigeon release behavior
(Online Resource 2). These data also include days the C
birds departed randomly (Auburn releases 0730, 1403, 1811,
2023; Weedsport releases 0480, 0765, 1236; Pulaski release
1646; and Orwell releases 1311, 1554). In addition, release
data were also assembled for C birds flown under sunny and
overcast weather conditions at Auburn and Weedsport to
determine whether or not such atmospheric (or weather) dif-
ferences had any discernable effect on the birds’ behavior.
The mean departure bearings for each site show that
untreated C birds, for the most part, were homeward oriented
at Auburn (Fig. 2a, b), Caroline I (Fig. 2g), Pulaski (Fig. 2h),
and Weedsport (Fig. 2c, d; Online Resource 2). The mean
departure bearing for C birds released at Castor Hill, how-
ever, was ~70° CW of the homeward bearing (Fig. 2e), and
was ~33° CCW of the homeward bearing at Orwell, PA
(Fig.  2f). Recent propagation modeling of infrasonic sig-
nals from the Cornell loft area to a number of sites in upstate
NY (Hagstrum 2013) indicates that such release-site biases
could result from long-term deflections of infrasonic signals
by topographic features under prevailing weather conditions,
while short-term changes could result from rapid shifts in
atmospheric wind and temperature fields.
Auburn
Fifty-seven pigeons were released 66 km N of the Cornell
loft at Auburn on 5 days (Fig. 1). The release begun on 1
May 1974 was aborted due to many landings, attributed
to high winds by the releasers (Online Resource 1), and
was continued on 2 May; the last release at Auburn, begun
under total overcast on 25 June 1974 was also aborted due
to clearing of the cloud cover (Table 1). The E birds were
consistently homeward oriented on each of the three main
release days at Auburn (Fig. 3a–c), whereas the equivalent
C birds were first oriented homeward under sunny skies on
20 November 1973, then poorly oriented eastward with the
Sun visible on 2 May 1974, and finally randomly oriented
under overcast on 10 May 1974. The 20 November and 10
May releases at Auburn (Fig. 3a, c) essentially reproduce
the results obtained by Wallraff (1972) in Germany with
the C birds being homeward oriented one day and ran-
domly oriented the next while the E birds were consistently
oriented homeward on both days.
J Comp Physiol A (2015) 201:983–1001 989
a b
Auburn Auburn
SUN O/C
e f
Castor Hill SUN
SUN Orwell, PA
Fig. 2  Departure bearings (dots) for untreated (control) pigeons
from other experimental releases at the 6 sites in upstate NY (Fig. 1).
Each dot represents the bearing of an individually released expe-
rienced bird (Online Resource 3). Geographic N is at the top of the
circle, and dashed lines indicate homeward directions. The mean
vectors (arrows) have bearings (φ) and length (r) that are 0 for ran-
domly oriented birds and 1 (radius of circle) for those perfectly
grouped (Table 1). All birds were released under sunny conditions,
except where noted. a Young and yearling birds released on 20 days
Clearly, there is no significant difference between
the E and C samples at Auburn on 20 November 1973
(Fig. 3a; Watson U2 test, P > 0.50; Batschelet 1981), but
on the remaining 4 days (Fig. 3b, c) the samples appear to
have behaved differently with the E birds oriented home-
ward and the C birds poorly oriented. Two of these days
were sunny (1 and 2 May 1974) and two were overcast
(10 May, 25 June 1974), but comparisons of other C-bird
releases at Auburn and Weedsport (Fig. 2a, b, c, d) show
that different weather conditions had no effect on their
mean departure bearings (Watson–Williams test: Auburn,
F = 0.008; Weedsport, F = 0.265; Baschelet 1981). Thus,
by pooling the results for Auburn, except those from 20
November 1973, we find that the sample of E departure
bearings were significantly different than those of the
C birds (Watson U2 test, P < 0.02). This result is robust
enough that divergence between the two samples is still
significant (P < 0.02) even when all the release data at
Auburn are pooled, including those of 20 November 1973
(Fig. 3a).
Caroline I
Peculiarly, C birds released just 15 km SE of the loft at Car-
oline I on 31 October 1973 under sunny conditions departed
c d
Weedsport Weedsport
SUN O/C
g h
Caroline I Pulaski
SUN SUN
at Auburn. b Similar birds released on 5 days under overcast con-
ditions (O/C) at Auburn. c Birds of all ages released on 26 days at
Weedsport. d Young and yearling birds released on 8 days at Weed-
sport under overcast conditions. (e) Yearling and old birds released
on 12 days at Castor Hill. f Birds of all ages released on 16 days at
Orwell, PA. g Birds of all ages released on 10 days at Caroline I; dot-
ted line indicates bearing to nearby farm buildings (see text). h Year-
ling and old birds released on 6 days at Pulaski
well oriented (Table 1), but 97° CCW from the homeward
bearing (Fig. 3d). Three of the E birds also headed in this
general direction, but the other two were oriented along the
homeward bearing. A subset of C birds from other releases
at Caroline I (Fig. 2g) also had departure directions toward
the WSW that are similar to those of the C birds observed
on 31 October 1973 (Fig. 3d). Apparently, departure direc-
tions for the E birds released at Caroline I are bi-modally
distributed. The E and C samples, however, are not signifi-
cantly different using Watson’s U2 test (P > 0.50), but are
significantly different when compared using a nonparamet-
ric test for dispersion (P < 0.05; Batschelet 1981). Unfortu-
nately, so few birds of each cohort were released at Caro-
line I (N = 11 total), that a more robust statistical analysis
cannot be made.
Weedsport
Although E birds released at Weedsport, 74 km N of the
loft (Fig. 1), under overcast on 8 and 12 November 1973
were poorly oriented (Table 1), C birds were well oriented
homeward (Fig. 3e). Watson’s U2 test (P > 0.10), however,
shows that the two samples were not significantly different
on these two days. On 26 July 1974 (Fig. 3f) both C and
E samples and a third of non-sham operated control (CC)
13

990 J Comp Physiol A (2015) 201:983–1001

a b c
Auburn Auburn Auburn
20 NOV 1973 1, 2 MAY; 25 10 MAY 1974
JUNE 1974
1067 1073 1082
SUN SUN O/C
1076

E birds 01 MAY 1974 02 MAY 1974

C birds 25 JUN 1974

d e f
Caroline I Weedsport Weedsport
31 OCT 1973 8, 12 NOV 1973 26 JUL 1974

1059 1063 1184

SUN O/C O/C
CC

08 NOV 1973 CC birds ( SUN)

g h i
Weedsport Castor Hill Castor Hill
29 AUG; 4, 5 2 JUN 1975 23 JUL 1975
NOV 1974 ME
1264 1348 1385 1434
O/C 1349 CC SUN SUN
F0

EB birds 04 NOV 1974 ME birds

CC birds
EM birds 05 NOV 1974 F0 birds

j k l
Pulaski Orwell, PA
30 SEP 1975 25 APR 1975 1384
O/C
1501
1367 Orwell, PA
ME SUN
O/C 30 MAY 1975

ME birds EB birds CB birds EB birds CB birds

EM birds EM birds

Fig. 3  Departure bearings for cochlea-extirpated (open dots) and
control (solid dots) pigeons released individually at the 6 sites in
upstate NY (Fig. 1). All birds are experienced and new to the release
site, and the diagrams are explained in the Fig. 2 caption. Home-
ward directions (dashed lines), release data, and distances are given
in Table 1 and Online Resource 2. Mean vectors are coded to the
released cohorts as shown below the diagrams. a Young birds released
at Auburn on 20 November 1973 under sunny conditions (SUN). b
Yearling birds released at Auburn on 1 and 2 May 1974 with Sun, and
on 25 June 1974 under overcast conditions (O/C). c Yearling birds
released at Auburn on 10 May 1974. d Young birds released at Car-
oline I on 31 October 1973; dotted line indicates bearing to nearby
farm buildings (see text). e Young birds released at Weedsport on 8
and 12 November 1973. f Yearling birds released at Weedsport on
26 July 1974. g Yearling birds (experimentals with either brass bars
(EB, gray symbols) or magnets (EM, open symbols) attached to them)
released at Weedsport on 29 August 1974 and 4 and 5 November
1974. h Old birds released at Castor Hill on 2 June 1975. i Yearling
and old (F0) birds released at Castor Hill on 23 July 1975; columel-
lae-bisected birds (ME) were also released. j Old birds released at
Pulaski on 30 September 1975; ME birds were also released. k Old
birds (controls also with brass bars, CB) released at Orwell, PA, on
25 April 1975. l Old birds released at Orwell, PA, on 30 May 1975

13
J Comp Physiol A (2015) 201:983–1001 991
birds were similarly well oriented (Table 1), and a compari-
son of the C and CC cohorts shows they are not statisti-
cally different (Watson U2 test, P > 0.20). All three cohorts
(C, EB, EM) were poorly oriented on 29 August 1974
(Fig. 3g).
Two more minor releases of E and C birds at Weedsport
on 4 and 5 November 1974 also included pigeons wearing
brass bars and magnets (Table 1). Because of the differ-
ences observed between released birds carrying magnets
and those with brass bars on overcast days (Keeton 1971),
we have omitted the birds with magnets (EM, CM) from
further consideration. We find that the E (Release 1063;
Fig.  3e) and EB (Release 1264; Fig. 3g) samples are not
significantly different (Watson U2 test, P > 0.10), and
conclude that the brass bars had little to no effect on the
pigeons’ behavior; we expect that carrying brass bars (~3 g;
Keeton 1971) would be potentially less disruptive than
carrying miniaturized GPS units (23–35 g; Schiffner et al.
2013). Also, the C sample is not significantly different from
the CC sample (Release 1184; Watson U2 test, P > 0.50).
If we pool all Weedsport data for the 5 release days into E
(including EB) and C (including CC) cohorts, the two sam-
ples are significantly different (Watson U2 test, P < 0.05),
most likely because of the more highly dispersed E bird
departure directions.
Castor hill
Sham-operated C birds released 143 km NNE of the loft at
Castor Hill on two sunny days (Fig. 3h, i), were well ori-
ented on 2 June 1975 at the same time the E birds were
more poorly oriented (Table 1). On 23 July 1975 the oppo-
site was true with C birds being less well oriented to the
WSW than the E birds. For both days, however, differences
between the two samples are not significant (Watson’s U2
test: 2 June, P > 0.20; 23 July, P > 0.50). Untreated CC
birds were also released on 2 June (Fig. 3h), and again had
similar departure directions to those of the C birds released
the same day (Watson’s U2 test: P > 0.10). If we pool the
C and CC cohorts and compare them to the E birds direc-
tion there is also no statistical difference (Watson’s U2 test:
P > 0.50). On 23 July 1975, ME and F0 birds were also
released at Castor Hill (Fig. 3; Table 1). Here again the
C and F0 samples are indistinguishable from one another
(Watson’s U2 test: P > 0.50), and the E and ME samples
are also statistically the same (Watson’s U2 test: P > 0.10).
If we pool the four cohorts released on 23 July 1975 into
operated and C samples, however, they are significantly
different (Watson’s U2 test: P < 0.01). A combination of
all release data at Castor Hill into operated and C samples
shows that they are significantly different as well (Watson’s
U2 test: P < 0.02).
Pulaski
On the one day, 30 September 1975, that E, ME, and C
birds were released under sunny conditions at Pulaski
(Fig. 3j), 127 km N of the loft, C birds were better oriented
than the E and ME birds (Table 1), and Watson’s U2 test
indicates that the difference between the E and C samples
is significant (P < 0.05). The E and ME samples are not
significantly different (Watson’s U2 test: P > 0.20), but if
we combine the E and ME directions into a single sample
and compare it with the C sample, the two samples become
statistically indistinguishable (Watson’s U2 test: P > 0.05).
The ME sample apparently has a strong homeward compo-
nent (Table 1) and these birds are statistically indistinguish-
able from the C birds released on that day as well (Wat-
son’s U2 test: P > 0.50).
Orwell, PA
For the releases conducted under overcast skies at Orwell,
PA, 66 km SSE of the loft, brass bars were attached to the
C birds and a randomly selected cohort of E birds, and
magnets were attached to a second cohort of E birds. On 25
April 1975, all three cohorts of birds (CB, EB, EM; Fig. 3k)
were randomly oriented (Table 1) and on 30 May 1975,
although both cohorts of E birds carrying brass and mag-
net bars were poorly oriented, the CB birds carrying brass
bars were well oriented to the NE (φ  = 304°; Fig. 3l) in
close agreement with the mean departure bearing for other
untreated birds released at Orwell, PA (φ = 306°; Fig. 2f;
Table  1). As before, because all releases were conducted
under overcast conditions, we will omit considering the
birds carrying magnets from our comparisons of the oper-
ated and C samples. The C and EB cohorts are statistically
indistinguishable (Watson’s U2 test: 25 April, P > 0.20; 30
May, P > 0.50), and a difference between the oriented and
random birds on the two days cannot be established (Wat-
son’s U2 test: P > 0.10). Again, too few E and C birds were
released at Orwell, PA, for a robust statistical analysis.
In summary, it is unfortunate that many individual test
releases at the six sites (Fig. 1; Table 1) contained too few
birds for reliable statistical comparisons to be made. By
pooling the available data for each site, however, such com-
parisons showed significant differences between the depar-
ture directions of C and E birds at all but two of the release
sites (Caroline I; Orwell, PA). At Caroline I, the departure
directions of the E birds were bimodal, and thus more dis-
persed, than those of the C birds (Fig. 3d), but only 11 birds
were released in total. At Orwell, PA, again few birds were
released in each cohort (≤8), but on 30 May 1975 the C
birds showed better homeward orientation than the E birds.
Overall, the C birds were better homeward oriented than
13

992
the E birds at all 5 sites with sufficient data (Caroline I
excluded; Table 1), but notably the C birds were randomly
or poorly oriented one day each at Auburn, Orwell, PA, and
Weedsport (Fig. 3c, g, k).
Ray‑trace simulations
From the acoustic explanation of Cornell birds’ disorien-
tation at Jersey Hill (Hagstrum 2013), the question arises
whether the C birds that departed randomly from Auburn on
10 May 1974 (Fig. 3c), and from Orwell, PA, on 25 April
1975 (Fig. 3k), did so because infrasonic signals from the loft
area were unavailable at the release sites on those two days.
Because rawinsonde data were collected at only two widely
separated weather stations in upstate NY during this time,
and HARPA runs incorporate only one set of daily profiles
per run, the propagation models do not account for local vari-
ations in atmospheric conditions. For this reason, Hagstrum
(2013) had previously selected extraordinary pigeon behav-
ior to see if it corresponded with exceptional regional atmos-
pheric variations resolvable with the available meteorologi-
cal data. Thus, to evaluate the possibility that the untreated C
birds in this study exploited ambient infrasonic cues, HARPA
computations were made of infrasound propagation for the
release days. It is presumed for these tests that well-oriented
C birds departing a release site had received clear homeward
signals, while those that were poorly oriented had not.
The calculated rays depicted in Fig. 4 show how infra-
sound waves virtually generated by HARPA at the Cornell
loft location (lower right corners of the diagrams), would
have traveled through the atmosphere above upstate NY on
each particular day, and show whether or not the infrasonic
signals would likely have reached the release site (lower left
corners of the diagrams). In these diagrams, the sound waves
(rays) are either refracted upward (Fig. 4b, c, f, h, k) by the
normal temperature structure of the troposphere (0–11 km
altitude) or were refracted back to the surface, and release
site, in ephemeral ducts caused by temperature inversions,
and/or wind shear, that had briefly formed at various eleva-
tions within the troposphere (Fig. 4a, d, e, g, i, j, l).
On 20 November 1973, C birds released at Auburn were
oriented homeward (Fig. 3a) and infrasonic signals appar-
ently traveled there from the loft vicinity (Fig. 4a), but on 10
May 1974 the C birds released at Auburn were randomly ori-
ented (Fig. 3c) and homeward infrasonic signals on that day
were refracted above the release site (Fig. 4b). Infrasonic sig-
nals from the loft area were also refracted upward over the
Caroline I release site on 31 October 1973 (Fig. 4c), and the
C birds headed off on bearings well away from the homeward
direction (Fig. 3d), although two E birds did orient along the
homeward bearing. Satellite photo imagery (Google Earth),
however, shows a conspicuous cluster of older farm buildings
(42.3556°N, 76.2834°W) situated ~1.5 km SW of Padlock
13
J Comp Physiol A (2015) 201:983–1001
tower (Caroline I release site, now a home site), which might
have visually attracted most of the released birds; homing
pigeons do not like to fly alone and often depart release sites
toward nearby buildings or villages in search of conspecifics
to flock with (e.g., Kiepenheuer 1993; Wiltschko et al. 2007).
Other C birds released at Caroline I (Fig. 2g) also followed
SW bearings perhaps caused by a similar attraction to the
farm buildings (Online Resource 2).
C birds were homeward oriented from Weedsport on
the first two release days (Fig. 3e, f) when infrasonic sig-
nals could have reached Weedsport along direct azimuths
(Fig.  4d, e), and were less well oriented from there on 29
August 1974 (Fig. 3g) when Weedsport was apparently
within a sound shadow (Fig. 4f). The ~15° CW release-
site bias documented at Weedsport by Keeton (1974) was
inferred to have resulted from diffraction by intervening ele-
vated terrain (Hagstrum 2013). Similarly, C birds departed
Castor Hill essentially without bias on 2 June 1975 when
infrasonic signals would have arrived there directly from the
loft (Fig. 4g). On 23 July 1975 at Castor Hill (Fig. 4h) and
30 September 1975 at Pulaski, infrasound transmitted along
direct azimuths would have been refracted above these
release sites. Acoustic signals traveling northward along the
Lake Cayuga valley (Fig. 4i, j), however, would have trave-
led within a strong ephemeral duct allowing the signals to
possibly pass around the intervening terrain and arrive at
their respective sites from directions consistent with the
expected release-site biases (Hagstrum 2013). In addition,
C birds (with brass bars) released at Orwell, PA, were ran-
domly oriented the day acoustic signals from Cornell were
refracted up and over the release site (Fig. 4k), and well ori-
ented along the site’s bias direction on the day the signals
were apparently received there (Fig. 4l); their deflection by
topographic features possibly caused them to arrive from
the 315° azimuth (Fig. 2f; Online Resource 2).
Discussion
The significant differences between the C and E birds of this
study (Table 1), and at the site NNE of the loft in Germany
(Wallraff 1972), indicate that the E birds were using a dif-
ferent navigational strategy than the C birds. For this reason
results for the C and E birds will be discussed separately. By
employing separate strategies, however, both the disoriented
C and E birds were able to return home after similar time
intervals and with similar overall losses to those shown by
the initially oriented cohort of birds (Online Resource 1).
Control birds
Although propagation modeling shows that the avail-
ability of infrasonic signals from the loft area is a feasible
J Comp Physiol A (2015) 201:983–1001 993

a AUBURN November 20, 1973 8 AM EDT

b AUBURN May 10, 1974 8 AM EDT
10 10

8 1067 8 1082

6 6

4 4

345° 350°
2 2

0 0
60 50 40 30 20 10 0 60 50 40 30 20 10 0

c CAROLINE I October 31, 1973 8 AM EDT

d WEEDSPORT November 12, 1973 8 AM EDT
10 10

8 1059 8
1063
HEIGHT (km)

6 6

4 4

121° 350°
2 2

0 0
14 12 10 8 6 4 2 0 70 60 50 40 30 20 10 0

e WEEDSPORT July 26, 1974 8 AM EDT

f WEEDSPORT August 29, 1974 8 AM EDT
10 10

8 8
1184 1264

6 6

4 4

350° 353°
2 2

0 0
70 60 50 40 30 20 10 0 70 60 50 40 30 20 10 0
RANGE AT SEA LEVEL (km)

Fig. 4  HARPA runs for the days on which operated and control birds
were released at the six sites in upstate NY and western PA (Fig. 1).
For all diagrams the right edge is at the Cornell loft, the opposite
edge is at the release site, the bottom edge is at mean sea level, and
the top edge is at 10 km altitude. The diagrams are oriented along the
azimuth listed to the lower left in black; for those diagrams with topo-
graphic profiles in gray (a, d, e, i and j) the topography is that along
the direct azimuth. For diagrams i and j the launch azimuths are along
the Lake Cayuga valley (320°). Launch elevations are from 0.0° to
45.0° at step intervals of 0.5°. Insets show the pigeon release data for
the same day (Fig. 3; Table 1). The topographic profiles are from a
30-m DEM of NY and PA to which the topographic profiles used by
HAPRA have been fitted. In some cases acoustic rays pass through
topographic features because the topographic profile used by the pro-
gram is a smoothed version of the actual DEM

explanation for why C birds were oriented or not upon larger dataset (Online Resource 2). The actual propagation
release, better characterizations of the atmosphere would of acoustic signals through the atmosphere is far more com-
be necessary to provide statistically convincing results for a plex than that represented by our HARPA computations

13

994 J Comp Physiol A (2015) 201:983–1001

g CASTOR HILL June 2, 1975 8 AM EDT

h CASTOR HILL July 23, 1975 8 AM EDT
10 10

8 1385 8 1434

6 6

4 4

20° 20°
2 2

0 0
140 120 100 80 60 40 20 0 140 120 100 80 60 40 20 0

i CASTOR HILL July 23, 1975 8 AM EDT

j PULASKI September 30, 1975 8 AM EDT
10 10

8 1434 8 1501
HEIGHT (km)

6 6

4 4

320° 320°
2 2

0 0
140 120 100 80 60 40 20 0 120 100 80 60 40 20 0

k ORWELL, PA April 25, 1975 8 AM EDT

l ORWELL, PA May 30, 1975 8 AM EDT
10 10

8 8
1367 1384

6 6

4 4

168° 168°
2 2

0 0
60 50 40 30 20 10 0 60 50 40 30 20 10 0
RANGE AT SEA LEVEL (km)

Fig. 4  continued

(Fig.  4), and reception throughout a given region can be
quite variable as shown by the following example. A large
chemical explosion (1–2 kilotons TNT equivalent) occurred
at Oppau, Germany, on 21 September 1921, and the inves-
tigation of its audibility across the region provided an
early illustration of the effects of upper atmospheric tem-
perature and wind profiles on sound propagation (Fig. 5;
Mitra 1952). Although infrasound can travel for thousands
of km, audible sounds are usually absorbed within a few
tens of km, except when the pressure level of the source is
extremely high (e.g., chemical or volcanic explosions).
Both the infrasonic and audible sound waves released
at Oppau would have been refracted upward by the normal
tropospheric temperature profile (decreasing upward) and

13
J Comp Physiol A (2015) 201:983–1001 995

200 km
refracted down to the surface at range by the stratospheric
temperature profile (increasing upward) creating the inter-
vening ‘classic zone of silence’ at the surface (e.g., Fig. 6
GERMANY
SW). In addition, finer-scale changes in wind and/or tem-
100 km perature fields would also have had significant effects on
Frankfurt
the characteristics of infrasound propagation and its far-
field reception (Evers and Haak 2010) and could cause
Oppau Nuremberg
the changes in audibility of the Oppau blast observed over
short distances in SW Germany and Switzerland (Fig. 5);
such changes cannot be modeled by HARPA using weather
Metz profiles from a single weather station but could explain
why pigeons can be disoriented at release sites where they
FRANCE are usually oriented.
Munich The ‘zone of silence’ for the Oppau blast has similar
dimensions to that expected for a ‘standard’ atmosphere
(Donn 1978) and to that determined from propagation
modeling relative to the Cornell loft in upstate NY. Figure 6
AUSTRIA shows a series of HARPA runs launched from the Cornell
loft location depicting the full atmosphere (120 km height)
SWITZERLAND Zurich
at 45° intervals in azimuth for the arbitrarily selected day of
21 Sept 1921 21 October 1977. Between the NW and E azimuths, near-
surface ephemeral ducts formed, but otherwise a shadow
zone was formed between ~25 and ~50 km and ~200 to
Fig. 5  The audible blast of a large chemical explosion at Oppau,
Germany, (square) was heard (black dots), or not heard (open cir- >250 km of the loft where the acoustic waves returned to
cles), in alternating zones of audibility and silence (after Mitra 1952). the surface from stratospheric (Fig. 6, SW to NW) and/or
The explosion was generally heard up to ~100 km away (dashed cir- thermospheric reflections (N to SE, and W). Variation in
cle), not heard within the classic ‘zone of silence’ between about 100
the dimensions of these shadow zones would result from
and 200 km, and heard beyond ~200 km (dashed circle; see Fig. 6) by
most observers to the E and S. Other selected cities in the region are changes in the atmospheric wind and temperature profiles
indicated by triangles that are also color-coded to the blast’s audibil- with time and between locations.
ity Similar types of sound shadows might also explain
the observed ‘distance effect’ on pigeon homing. In a

Cornell Loft 21 October 1977

HEIGHT (120 km)

N NE E SE

S SW W NW

Zone of Silence

RANGE AT SEA LEVEL (250 km)

Fig. 6  HARPA runs showing acoustic rays launched from the Cor-
nell loft location (42.4360°N, 76.4319°W) at 45° intervals in azimuth
for 21 October 1977 under sunny conditions. Dashed lines indicate
divisions between the troposphere, stratosphere, mesosphere, and
thermosphere. Rays were ducted near the surface between the NW
and E azimuths, but otherwise the classic ‘zone of silence’ at the
surface was formed between the SE and W azimuths; it is indicated
below the SW panel. Thermospheric reflections also occurred along
the NW to SE azimuths, and stratospheric reflections along the SW
to NW azimuths. The loft is at the lower left corner in the upper four
panels, and at the lower right in the lower four; the height (120 km)
and range (250 km) values refer to the individual panels

13

996
number of studies, pigeons showed a high level of home-
ward-directed orientation nearer the loft, a zone of poor
homeward directedness farther out (>25 km), improved
homeward directedness beyond ~100 km, and declining
homeward directedness again by 250–300 km (Schmidt-
Koenig 1966, 1979). Whether this phenomenon is observed
or not would depend on the regional atmospheric condi-
tions as well as the homeward bearing: as shown in Fig. 6,
acoustic propagation along some azimuths produces zones
of silence at the surface (e.g., SW azimuth), whereas along
others near-surface ducting produces continuous signal
availability (e.g., NE azimuth).
At Jersey Hill, untreated Cornell birds were lost with
high attrition rates probably because the site lay within an
extensive acoustic shadow zone ranging from at least the
Prattsburg release site (74 km W of the loft) to the Spring-
ville II site (178 km W of the loft; Hagstrum 2013). Why
Cornell birds departing the Jersey Hill zone of silence
did not eventually reorient to available acoustic signals
is unclear, but perhaps after a lengthy period within the
shadow zone they had given up searching and selected a
bearing, albeit incorrect, for extended flight; one-banded
Cornell pigeon released at Jersey Hill was later found
in Cleveland, OH, and another in central Canada. On the
other hand, birds initially disoriented at sites where they
were normally oriented, and that returned home within
normal flight times, were perhaps released at sites within
local shadow zones (e.g., Fig. 5) caused by nearby topo-
graphic features and/or small-scale changes in atmospheric
temperature and wind fields. In such cases, random depar-
tures would be a useful strategy in more quickly exiting
the shadow zone and finding areas with better reception of
their homeward acoustic signal.
Experimental birds
If the C birds used infrasonic cues for initial orientation,
what other strategy could the E birds have used to return to
the Cornell loft? Cornell pigeons released at Jersey Hill did
not switch to alternate cues when other navigational fac-
tors (e.g., Sun’s azimuth, geomagnetic field, atmospheric
odors, visual landmarks) would have been available over
much, if not all, of the 19 years that birds were released
there. It is also seems unlikely that some selection of these
cues, being related to separate senses, could so completely
interfere with one another. Moreover, non-Cornell birds
released from Jersey Hill had no difficulty in returning to
their respective lofts (Walcott and Brown 1989), showing
that for these pigeons there was no conflict between navi-
gational cues. Such differences in behavior between birds
from different lofts have been attributed to the influence of
local variations in one or more navigational factors at their
home sites (e.g., magnetic fields, Walcott 1992). Some of
13
J Comp Physiol A (2015) 201:983–1001
the oriented birds released at Jersey Hill, however, were
from a loft only 25 km W of Ithaca (Walcott and Brown
1989), and in many studies of pigeon homing investigators
have only been able to assert that their birds were respond-
ing to the designated cue; the experiments were not, and in
most cases could not be, adequately controlled. Thus, sites
of disorientation (Schmidt-Koenig in Brown 1971) indicate
that pigeons, at least, consistently select one navigational
cue, rather than ‘checking’ for and selecting alternative
ones. From acoustic propagation modeling of the Jersey
Hill releases (Hagstrum 2013), the preferred cue appears to
be infrasound.
With the homing pigeons’ apparent reluctance to use
alternative non-acoustic cues, and the E birds’ aural sense
destroyed, we propose that pigeons have evolved to search
for an acoustic homeward signal, and, with their hearing
sense intact, would continue to do so ad infinitum. If, on
the other hand, they are made completely and permanently
deaf, then we expect, at some level, they would become
aware of this and realize that their only chance of return-
ing home would be switching to an alternate strategy. A
plausible explanation for the homing of E birds is that they
monitored a compass sense during transport from the loft,
and after integrating this information with time, were able
to reverse the overall outward route and determine their
homeward bearing (Wiltschko and Wiltschko 1978). This
task would be relatively straightforward if transport from
loft to release site was for the most part direct and without
significant detour (e.g., Papi et al. 1984). Young, inexperi-
enced birds (<12 weeks old) apparently use their magnetic
compass to return home from experimental displacements,
and this strategy is apparently used before they learn their
more sophisticated adult navigational system (Wiltschko
and Wiltschko 1985).
Transport of very young birds in a distorted magnetic
field showed significantly increased scatter in depar-
ture bearings compared to those for unexposed birds of
equivalent age (Wiltschko and Wiltschko 1982). Simi-
larly, transport of young birds in total darkness or under
red light (660 or 630 nm) also caused increased scatter in
their departure bearings upon release compared to C birds
transported in ‘white’ or green (565 nm) light, respectively
(Wiltschko and Wiltschko 1981, 1998). These results indi-
cate that either white or green light is necessary to activate
the putative cryptochrome-based avian magnetic compass
(Ritz et al. 2009; Zapka et al. 2009). Thus, deafened adult
birds could have resorted to a navigational strategy that
they had once used as younger birds. Notably, when young
pigeons shift from a route-reversal strategy to their adult
navigational system they appear to do so “completely”,
even though their homing performance would be better at
the outset had they continued to use directional informa-
tion acquired during the outward journey (Wiltschko and
J Comp Physiol A (2015) 201:983–1001 997
Wiltschko 1985). This observation is also consistent with
experienced pigeons’ dedication to a single long-range
navigational cue. Interestingly, proponents of the olfac-
tory hypothesis of pigeon homing have also concluded
that pigeons rely predominantly on a single “atmospheric
factor” for navigation from unfamiliar regions (e.g., Papi
1986; Wallraff 2005).
Although the E birds possibly used a light-dependent
magnetic compass situated within their retina as part of
a route-reversal strategy in returning home, the cochlea,
lagena and other organs of the vestibular system have also
been implicated as magnetic field receptors (Harada 2002;
Wu and Dickman 2011, 2012; Lauwers et al. 2013). Stud-
ies of chicken lagenae have shown that it is not a recep-
tor of normal sound signals, although infrasound was not
tested in that study (Manley et al. 1991). Iron-rich orga-
nelles have been found within the sensory hair cells of the
avian cochlea and vestibular system that might mediate
magnetoreception (Lauwers et al. 2013), and cells within
the pigeon’s brainstem vestibular nuclei have been shown
to encode magnetic field direction, intensity, and polarity
(Wu and Dickman 2012). Apparently, the ocular magnetic
compass supersedes an equivalent within the inner ear,
and perhaps the directional redundancy between these two
receptors is a developmental byproduct of the inner-ear
receptor’s functionality in detecting geomagnetic intensity
(e.g., Wiltschko and Wiltschko 1995).
The ME birds released at Castor Hill and Pulaski with
bisected columellae would certainly have been hearing
impaired, but were most likely not profoundly deaf to infra-
sonic signals. The mean departure directions for these birds
(Fig.  3i, j; Table 1) are the same as those for the other C
birds released at Castor Hill and Pulaski shown in Fig. 2e,
h (Watson–Williams test: Castor Hill, F = 0.278; Pulaski,
F  = 0.793; Online Resource 2). Unfortunately, only two
releases of ME birds were made, and more data would be
needed to determine if the behavior of these birds was gen-
erally consistent with that of C birds released at these same
sites. Still, the ME birds released at Castor Hill on 23 July
1975 departed in significantly different directions than the
C and F0 birds also released on that day (Watson’s U2 test:
P < 0.01).
The infrasound hypothesis of avian navigation is based
on known acoustic signals (see Box 1) and is therefore
directly testable, whereas the olfactory hypothesis, based
on unidentified odors from unknown sources (Wallraff
2005, 2013), is not. Moreover, the infrasound hypothesis
might explain observations attributed to atmospheric odors.
For example, winds favor reception of acoustic waves at
the surface in the direction they are blowing (Bedard and
Georges 2000), and young birds restricted to open aviar-
ies for weeks to months, and able to detect near-surface
wind direction, could determine the associated bearings of
N
W E
1m
S
Fig. 7  Plan view of a deflector aviary with flow lines showing
the simplified counterclockwise deflection of air currents entering
the aviary from the W (after Papi 1986). In this example, station-
ary pigeons cannot localize a western infrasonic source, enhanced
by a westerly wind, but could associate the infrasonic signal with a
deflected wind direction from the S (see text). Because fluids do not
follow the law of reflection, wind currents would not be deflected
at exactly 90° as shown. Baldaccini et al. (1975) observed variable
deflections, and turbulence, during wind tunnel experiments that aver-
aged ~70° in the expected sense within a 1:10 scale model of the
deflector aviary
proximal sound sources from relative changes in acoustic
amplitude as the wind direction shifts over time. Solid wall
structures surrounding open aviaries (Kramer 1959; Wall-
raff 1979) are small compared to infrasonic wavelengths
and would have minor effects on infrasonic signals, but
would block the wind preventing its directional information
from reaching the birds within. The slanted panels posi-
tioned around aviaries to deflect wind currents ~90° CW or
CCW (e.g., Baldaccini et al. 1975; Fig. 7) are also much
too small to influence the passage of infrasonic waves (and
likely large-scale odor gradients as well), but would change
the apparent direction of the wind. Thus, pigeons within a
CCW deflector aviary (Fig. 7) could interpret the unique
infrasonic signals or ‘voice prints’ from western sources,
enhanced by westerly winds, as coming from the S based
solely on the perceived wind direction. Moreover, Kie-
penhauer (1979) demonstrated that pigeons with plugged
nares or anesthetized olfactory epithelia still responded to
the ‘deflector effect’, indicating that it does not result from
olfactory stimuli.
In conclusion, release experiments carried out at Cornell
University between 1973 and 1975 with homing pigeons
having extirpated cochleae and lagenae show significant
differences between their departure bearings and those of
intact birds, and a hypothesis based on aural navigational
13

998
cues can explain all observations concerning both cohorts.
Although a greater quantity of release data would have
permitted more rigorous statistical analyses, our interpre-
tation illustrates the risk of conducting too few experimen-
tal releases (e.g., Wallraff 1972) with limited numbers of
birds. The behavior upon release of the C birds examined
here is inferred to have depended on the local availability
and strength of homeward infrasonic signals, as did the
occasional initial disorientation of C birds at the same sites
in other releases from the Keeton database (Fig. 2; Online
Resource 2). On the other hand, E birds are inferred to
have employed an alternative route-reversal navigational
strategy to return home, plausibly using their ocular-based
magnetic compass. Because pigeons released at sites of
disorientation do not switch to alternate navigational cues,
it could be complete destruction of the crucial aural sense
that forces experienced pigeons to shift from their primary
navigational strategy to a secondary one; this could be
tested by releasing E birds at Jersey Hill, or other sites of
perennial disorientation, to see if they orient homeward. We
agree with proponents of the olfactory navigational hypoth-
esis that birds depend predominantly on an “atmospheric
factor” for long-range navigational purposes (e.g., Papi
1986; Wallraff 2005), but think the dynamic propagation of
infrasound, rather than the passive transport of trace odors,
might better fit much of the experimental evidence as well
as our current understanding of atmospheric processes.
Acknowledgments  Foremost we wish to acknowledge Bill Keeton
and other collectors of pigeon-release data at Cornell University, and
in particular Irene Brown, Tim Larkin and Steven Emlen for assem-
bling the ‘Keeton’ database. Charlie Walcott kindly sent it to us, as
well as original correspondence and data sheets. Tim Larkin also
generously provided his program ‘Keeton’ and other insider infor-
mation concerning the database and Cornell pigeon releases. Many
thanks to Mike Jones for providing HARPA and guidance in its use,
and to Larry Baker for installing it on USGS computers and making
improvements to the code. We thank Al Bedard, Hugh McIsaac, Bruce
Thigpen, Doug Drob, and Rich Raspet for contributing insightful
comments. In addition, John Spritzer contributed the digital elevation
model and topographic profiles. Ingo Schiffner, Wolfgang and Roswi-
tha Wiltschko, Patrick Muffler, Bob Gill, and Hans-Peter Lipp help-
fully reviewed an early version of the manuscript. Funding for this
work, in part, came from a USAF SBIR grant (FA9302-12-C-0006)
through Technology International Incorporated of Virginia to the
USGS, and from grant A6368 to GAM from the Canadian National
Research Council.
Appendix: Box 1
The infrasound hypothesis of avian navigation
Because atmospheric absorption of acoustic waves
increases with frequency, infrasound can travel great
13
J Comp Physiol A (2015) 201:983–1001
distances in the atmosphere with relatively undiminished
intensity (Bedard and Georges 2000). Thus, an array of
low-frequency microphones placed at any point on Earth
will detect an incessant cacophony of infrasonic waves
from natural and man-made sources on a global scale. A
worldwide network of infrasonic and other sensors has
been established to monitor compliance with the Com-
prehensive Test Ban Treaty (Sullivan 1998), as well as
for research purposes. Since 1949 a considerable amount
of research and technological development has gone into
detecting and isolating the infrasonic signatures of large
explosive events from a wide variety of atmospheric
background ‘noise’ (Bedard and Georges 2000). A near-
continuous source of infrasound, nearly monochromatic
in frequency (~0.2 Hz peak) and with 5–7 s wave peri-
ods called microbaroms, is generated by interacting trains
of deep-ocean waves that produce local areas of verti-
cal motion or ‘standing’ waves (Rind 1980; Bedard and
Georges 2000). Such standing waves also cause pressure
changes on the seafloor that generate seismic waves in the
solid Earth of equivalent frequency called microseisms
(Rind 1980; Kedar et al. 2008). Microseismic waves
similarly are detected by seismometers worldwide, and
are commonly referred to by seismologists as the ‘6-sec
noise’.
Aside from interfering with earthquake signals, micro-
seisms also cause the ground surface to raise and fall
slightly (~0.5 to ~5 μm; Kedar et al. 2008), and ground-
to-air coupling of this movement creates acoustic waves.
As ‘proof of concept’, much higher amplitude ground
movements from earthquakes are regularly detected above
the ‘noise floor’ of infrasound arrays, and only those sig-
nals radiated from steep-sided terrain, having significant
horizontal components, travel over great distances (e.g.,
Young and Greene 1982; Greene et al. 2009). We suggest
that the land surface’s complex geometry in any particular
area (i.e., surrounding any loft) radiates unique infrasonic
signals, or ‘voice prints’, that pigeons are able to identify
from any direction as their homeward signal. An alternative
source of characteristic infrasound from the loft area could
result from the scattering of microbaroms, arriving directly
from the ocean surface, off of topographic features. Only
those features with dimensions similar to, or larger than,
the infrasonic wavelengths would cause significant scatter-
ing and/or radiation.
Microbaroms have wavelengths much too long
(~1.7 km) for birds to locate their sources with binaural
separations of only a few cm. Source azimuths could be
determined, however, using Doppler shifts of the home-
ward signal as birds circle or fly about the release site
prior to departing in the homeward direction (Quine and
Kreithen 1981). Such looping flight at low altitudes,
combined with optical flow of the ground surface (e.g.,
J Comp Physiol A (2015) 201:983–1001 999
Schiffner et al. 2014), might also serve to calibrate wind
drift. Not all pigeons circle upon release, and some-
times head off directly, but they could have tracked their
homeward signal’s direction with Doppler shifts during
changes in direction while being transported from the loft.
As discussed in the main text, compass senses might also
apply in keeping track of orientation during transport.
Thus, no gradient ‘map’ would be needed, and neither
bi-coordinate geophysical (i.e., magnetic; e.g., Wiltschko
and Wiltschko 1995) nor natural olfactory (e.g., Wall-
raff 2005) gradients are known, or likely, to persist with
adequate resolution over significant distances. Compass
senses, however, would be required in an infrasound-
based avian navigational system for orientation during
the return flight home. The source location of infrasonic
signals cannot be detected during direct flight, but birds
could circle again, change direction, or fly in zigzag pat-
terns to check or reestablish the bearing of their home-
ward signal.
We suggest that older birds do not abandon their innate
magnetic compass, which is always available, but use
it, and not the visual landscape, to register directions of
interest; birds likely dedicate their sense of vision during
flight to detection and avoidance of predators. The sun
compass is calibrated during development by the mag-
netic compass (Wiltschko and Wiltschko 1981), but even-
tually becomes the dominant one (e.g., Kramer 1959).
Pigeons might use their sun compass, as migratory birds
do, to adjust their magnetic compass for inaccuracies
(e.g., Muheim et al. 2006) resulting from, for example,
local magnetic declination. Pigeons shifting their mag-
netic reference frame (including the homeward direc-
tion determined by circling) to agree with an artificially
altered circadian rhythm and sun compass, would also
depart release sites well away from the homeward bear-
ing, as expected, depending on the sense and amount of
clock shift. Departures of older experienced birds show-
ing less than the predicted shift (Wiltschko et al. 1994)
could result from partial resolution of the conflict between
sun compass and acoustic bearings.
Sound intensity decreases with distance from the source
according to the inverse square law for point sources, and
birds far from the loft could be listening for stronger sig-
nals generated by collections of appropriately oriented
terrain features within much larger geographic areas. As
birds approach their loft the source area would dimin-
ish in size, and the 1–2 km accuracy of pigeons homing
without sight (Schmidt-Koenig and Walcott 1978) might
reflect the minimum dimension of a proximate source
area, which theoretically would be equivalent to the infra-
sonic signal’s wavelength (~1.7 km). It can be calculated
how much area would be needed to generate sound levels
at various distances, and as given in Hagstrum (2013) it
would take the combined effect of an area totaling ~1 km2,
vibrating in unison at 0.2 Hz with an amplitude of 1 μm
(typical ground motion due to microseisms), to create a
sound intensity of 120 dB SPL at a range of ~1 km. From
the pigeon audiogram presented in Kreithen and Quine’s
(1979) paper the threshold of sound detection steadily
increases as the frequency drops, reaching 120 dB SPL at
0.2 Hz.
Like any other natural system, difficulties with an acous-
tic navigational system can arise from intermittent com-
plications like shadow zones (e.g., Jersey Hill, Hagstrum
2013), deflection of signals by topographic features (e.g.,
release-site biases, Hagstrum 2013), changes in atmos-
pheric conditions (e.g., temperature inversions, Wagner
1978), and jamming by local or distant sources (e.g., sur-
face waves of lakes, Wagner 1972; sonic booms, Hagstrum
2000). In addition, microbaromic waves generated by local
terrain features could mask distant signals, but one means
of distinguishing signals from distant sources is their angle
of arrival: they would tend to arrive at steeper angles from
reflections within ephemeral or stratospheric ducts (Fig. 4).
Signals from proximal sources, on the other hand, arrive
more or less horizontally as they have not yet sustained
the cumulative effects of upward refraction by the tropo-
sphere’s temperature profile (Figs. 4, 6). The transverse
speed of an infrasonic wave across a stationary array indi-
cates its angle of incidence: a speed equal to the ambient
sound speed indicates the wave traveled horizontally. As the
angle of incidence increases the transverse speed increases,
and instantaneous arrivals across an array indicate the wave
arrived vertically (Christie and Campus 2010). Birds could
possibly establish an airborne synthetic-aperture ‘array’ by
circling, but knowledge of airspeed would be an important
factor; a future paper will consider how they might accom-
plish this.
Microbaroms are confined to a relatively narrow fre-
quency band making it easier to filter them from other
atmospheric noise mechanically and/or during signal pro-
cessing, and animals can perform remarkable feats of sig-
nal recognition. For example, bats can identify their own
complex echolocation signals from those generated by a
large number of conspecifics, and although the solution to
this problem of interference is not yet fully understood, big
brown bats (Eptesicus fuscus) apparently improve the situ-
ation by reducing their rate of vocalizations during group
flight (Chiu 2008). It will probably take many years to
completely understand the sophisticated methods by which
homing pigeons possibly detect and exploit infrasonic sig-
nals for navigational purposes, because, by analogy, even
though we have known for ~75 years that bats use ultra-
sonic signals to echolocate (Griffin 1958) we still do not
fully comprehend their highly evolved techniques (Denny
and McFadzean 2011).
13

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References Keeton WT (1971) Magnets interfere with pigeon homing. Proc Nat
Acad Sci 68:102–106
Keeton WT (1973) Release-site bias as a possible guide to the “map”
Baldaccini NE, Benvenuti S, Fiaschi V, Papi F (1975) Pigeon naviga-
component in pigeon homing. J Comp Physiol 86:1–16
tion: effects of wind deflection at home cage on homing behav-
Keeton WT (1974) The orientation and navigational basis of homing
iour. J Comp Physiol 99:177–186
in birds. Advan Study Behav 5:47–132
Batschelet E (1981) Circular statistics in biology. Academic Press,
Kiepenheuer J (1978) Pigeon homing: a repetition of the deflector loft
London
experiment. Behav Ecol Sciobiol 3:393–395
Bedard AJ Jr, Georges TM (2000) Atmospheric infrasound. Phys
Kiepenheuer J (1979) Pigeon homing: deprivation of olfactory infor-
Today 53:32–37
mation does not affect the deflector effect. Behav Ecol Sociobiol
Brown F Jr (1971) Unconventional theories of orientation, panel dis-
6:11–22
cussion. In: Adler HE (ed) Orientation: sensory basis, Part V.
Kiepenheuer J (1993) The ambiguity of initial orientation of homing
Ann NY Acad Sci 188:331–358
pigeons. Birds, Humans and Other Animals. Royal Institute of
Brown I, Larkin T, Emlen ST (1984) Fourteen years of pigeon hom-
Navigation, Oxford, UK, In Orientation and Navigation
ing data. Auk 101:622–623
Kramer G (1959) Recent experiments on bird orientation. Ibis
Chiu C, Wei X, Moss CF (2008) Flying in silence: echolocating bats
101:399–416
cease vocalizing to avoid sonar jamming. Proc Nat Acad Sci
Kramer G, von St. Paul U (1954) Das heimkehrvermögen gekäfigter
105:13116–13121. doi:10.1073/pnas.0804408105
Brieftauben. Ornithol Beob 51:4–12
Christie DR, Campus P (2010) The IMS infrasound network: design
Kreithen ML, Quine DB (1979) Infrasound detection by the homing
and establishment of infrasound stations. In: Le Pichon A, Blanc
pigeon: a behavioral audiogram. J Comp Physiol 129:1–4
E, Hauchecorne A (eds) Infrasound monitoring for atmospheric
Lauwers M, Pichler P, Edelman NB, Resch GP, Ushakova L, Salzer MC,
studies. Springer, Berlin, pp 29–75
Heyers D, Saunders M, Shaw J, Keays DA (2013) An iron-rich
Denny M, McFadzean A (2011) Engineering animals. Harvard Uni-
organelle in the cuticular plate of avian hair cells. Curr Biol 23:1–6
versity Press, Cambridge
Manley GA, Haeseler C, Brix J (1991) Innervation patterns and spon-
Donn WL (1978) Exploring the atmosphere with sonic booms. Amer
taneous activity of afferent fibres to the lagenar macula and api-
Sci 66:724–733
cal basilar papilla of the chicken’s cochlea. Hear Res 56:211–226
Drob DP, Emmert JT, Crowley G, Picone JM, Shepherd GG, Skin-
Mitra SK (1952) The upper atmosphere. The Asiatic Society, Calcutta
ner W, Hays P, Niciejewski RJ, Larsen M, She CY, Meriwether
Muheim R, Moore FR, Phillips JB (2006) Calibration of magnetic
JW, Hernandez G, Jarvis MJ, Sipler DP, Tepley CA, O’Brien
and celestial compass cues in migratory birds—a review of cue-
MS, Bowman JR, Wu Q, Murayama Y, Kawamura S, Reid IM,
conflict experiments. J Exp Bio 209:2–17
Vincent RA (2008) An empirical model of the Earth’s hori-
Norris D, Gibson R, Bongiovanni K (2010) Numerical methods to
zontal wind fields: HWM-07. J Geophys Res. doi:10.1029/200
model infrasonic propagation through realistic specifications of
8JA0113668
the atmosphere. In: Le Pichon A, Blanc E, Hauchecorne A (eds)
Evers LG, Haak HW (2010) The characteristics of infrasound, its
Infrasound monitoring for atmospheric studies. Springer, Berlin,
propagation, and some early history. In: Le Pichon A, Blanc E,
pp 541–573
Hauchecorne A (eds) Infrasound monitoring for atmospheric
Papi F (1986) Pigeon navigation: solved problems and open ques-
studies. Springer, Berlin, pp 3–27
tions. Monitore Zool Ital 20:471–517
Green DN, Guilbert J, Le Pichon A, Sebe O, Bowers D (2009) Mod-
Papi F, Ioalé P, Fiaschi V, Benvenuti S, Baldaccini NE (1984) Pigeon
eling ground-to-air coupling for the shallow ML 4.3 Folkestone,
homing: the effect of outward-journey detours on orientation.
United Kingdom, earthquake of 28 April 2007. Bull Seis Soc Am
Monitore Zool Ital 18:53–87
99:2541–2551. doi:10.1785/0120080236
Picone JM, Hedin AE, Drob DP, Aiken AC (2002) NRLMSISE-00
Griffin DR (1958) Listening in the dark. Yale University Press, New
empirical model of the atmosphere: statistical comparisons and
Haven
scientific issues. J Geophys Res 107:1468
Griffin DR (1969) The physiology and geophysics of bird navigation.
Quine D (1982) Infrasounds: A potential navigational cue for hom-
Quart Rev Biol 44:255–276
ing pigeons. In: Papi F, Wallraff HG (eds) Avian navigation.
Hagstrum JT (2000) Infrasound and the avian navigational map. J
Springer, Berlin, pp 373–376
Exp Biol 203:1103–1111
Quine DB, Kreithen ML (1981) Frequency shift discrimination: can
Hagstrum JT (2013) Atmospheric propagation modeling indicates
homing pigeons locate infrasounds by Doppler shifts? J Comp
homing pigeons use loft-specific infrasonic ‘map’ cues. J Exp
Physiol 141:153–155
Biol 216:687–699
Rind D (1980) Microseisms at Palisades, 3. Microseisms and micro-
Harada Y (2002) Experimental analysis of behavior of homing
baroms. J Geophys Res 85:4854–4862
pigeons as a result of functional disorders of their lagena. Acta
Ritz T, Witschko R, Hore PJ, Rodgers CT, Stapput K, Thalau P, Tim-
Otolaryngol 122:132–137
mel CR, Wiltschko W (2009) Magnetic compass of birds is
Hedin AE, Fleming EL, Manson AH, Schmidlin FJ, Avery SK, Clark
based on a molecule with optimal directional sensitivity. Biophys
RR, Franke SJ, Fraser GJ, Tsuda T, Vial F, Vincent RA (1996)
J 96(8):3451–3457
Empirical wind model for the upper, middle, and lower atmos-
Schermuly L, Klinke R (1990) Origin of infrasound sensitive neu-
phere. J Atmos Terr Phys 58:1421–1447
rons in the papilla basilaris of the pigeon: a HRP study. Hear Res
Jones RM (1982) Algorithms for reflecting rays from general topo-
48:69–78
graphic surfaces in a ray-tracing program. NOAA Technical
Schiffner I, Fuhrmann P, Wiltschko R (2013) Homing flights of
Memorandum ERL WPL-98
pigeons in the Frankfurt region: the effect of distance and local
Jones RM, Riley JP, Georges TM (1986) HARPA: a versatile three-
experience. Animal Behav 86:291–307
dimensional hamiltonian ray-tracing program for acoustic waves
Schiffner I, Vo HD, Bhagavatula PS, Srinivasan MV (2014) Minding
in the atmosphere above Irregular Terrain. NOAA, Boulder
the gap: in-flight body awareness in birds. Front Zool 11:64
Kedar S, Longuet-Higgens M, Webb F, Graham N, Clayton R, Jones
Schmidt-Koeing K (1987) Bird navigation: has olfactory orientation
C (2008) The origin of deep ocean microseisms in the North
solved the problem? Quart Rev Biol 62:31–47
Atlantic Ocean. Proc R Soc A 464:777–793

13
J Comp Physiol A (2015) 201:983–1001 1001
Schmidt-Koenig K (1966) Über die entfernung als parameter bei der
anfangsorientierung der brieftaube. Z Vergl Physiol 52:33–55
Schmidt-Koenig K (1979) Avian orientation and navigation. Aca-
demic Press, London
Schmidt-Koenig K, Walcott C (1978) Tracks of pigeons homing with
frosted lenses. Anim Behav 26:480–486
Schwarzkopff J (1949) Über Sitz und Leistung von Gehör und Vibra-
tionssinn bei Vögeln. Zeitschrift für vergleichende Physiologie
31:527–608
Sullivan JD (1998) The comprehensive test ban treaty. Phys Today
51:24–29
Wagner G (1972) Topography and pigeon homing. In: Galler SR,
Schmidt-Koenig K, Jacobs GJ, Belleville RE (eds) Animal ori-
entation and navigation SP-262. NASA, Washington, DC, pp
259–273
Wagner G (1978) Homing pigeons’ flight over and under low stratus.
In: Schmidt-Koenig K, Keeton WT (eds) Animal migration, nav-
igation, and homing. Springer, Berlin, pp 162–170
Walcott C (1992) Pigeons at magnetic anomalies: the effects of loft
location. J Exp Biol 170:127–141
Walcott C (1996) Pigeon Homing: observations, experiments, and
confusions. J Exp Biol 199:21–27
Walcott C (2005) Multi-modal orientation cues in homing pigeons.
Integr Comp Biol 45:574–581
Walcott C, Brown AI (1989) The disorientation of pigeons at Jersey
Hill. In: Orientation and Navigation: birds, humans and other
animals. Royal Institute of Navigation, Cardiff, Wales
Waldvogel JA, Benvenuti S, Keeton WT, Papi F (1978) Homing
pigeon orientation influenced by deflected winds at home loft. J
Comp Physiol 128:297–301
Wallraff HG (1966) Über die Heimfindeleistungen von Brieftauben
nach Haltung in verschiedenartig abgeschirmten Volieren. Z
Vergl Physiol 52:215–259
Wallraff HG (1970) Über die flugrichtungen verfrachteter brieftauben
in abhängigkeit vom heimatort und vom ort der freilassung. Z
Tierpsychol 27:303–351
Wallraff HG (1972) Homing of pigeons after extirpation of their
cochleae and lagenae. Nature New Biology 236:223–224
Wallraff HG (1979) Goal-oriented and compass-oriented movements
of displaced homing pigeons after confinement in differently
shielded aviaries. Behav Ecol Sociobiol 5:201–225
Wallraff HG (2001) Navigation by homing pigeon: updated perspec-
tive. Ethol Ecol Evol 13:1–48
Wallraff HG (2005) Avian navigation: pigeon homing as a paradigm.
Springer-Verlag, Berlin
Wallraff HG (2013) Ratios among atmospheric trace gases together
with winds imply exploitable information for bird navigation: a
model elucidating experimental results. Biogeosci 10:6929–6943
Wiltschko R, Wiltschko W (1978) Evidence for the use of outward-
journey information in homing pigeons. Naturwissenschaften
65:112–113
Wiltschko W, Wiltschko R (1981) Disorientation of inexperienced
young pigeons after transportation in total darkness. Nature
291:433–434
Wiltschko W, Wiltschko R (1982) The role of outward journey infor-
mation in the orientation of homing pigeons. In: Papi F, Wallraff
HG (eds) Animal navigation. Springer, Berlin, pp 239–252
Wiltschko R, Wiltschko W (1985) Pigeon homing: change in naviga-
tional strategy during ontogeny. Anim Behav 33:583–590
Wiltschko R, Wiltschko W (1995) Magnetic orientation in animals.
Springer, Berlin
Wiltschko R, Wiltschko W (1998) Pigeon homing: effect of various
wavelengths of light during displacement. Naturwissenschaften
85:164–167
Wiltschko R, Wiltschko W (2009) Avian navigation. Auk
126:717–743
Wiltschko R, Kumpfüller R, Muth R, Wiltschko W (1994) Pigeon
homing: the effect of Clock-shift is often smaller than predicted.
Behav Ecol Sociobiol 35:63–73
Wiltschko R, Schiffner I, Siegmund B (2007) Homing flights of
pigeons over familiar terrain. Anim Behav 74:1229–1240
Wu L-Q, Dickman JD (2011) Magnetoreception in an avian brain in
part mediated by inner ear lagena. Curr Biol 21:418–423
Wu L-Q, Dickman JD (2012) Neural correlates of a magnetic sense.
Science 336:1054–1057
Yodlowski ML, Kreithen ML, Keeton WT (1977) Detection of atmos-
pheric infrasound by homing pigeons. Nature 265:725–726
Young JM, Greene GE (1982) Anomalous infrasound generated by
the Alaskan earthquake of 28 March 1964. J Acoust Soc Am
71:334–339
Zapka M, Heyers D, Hein CM, Engels S, Schneider N-L, Hans J,
Weiler S, Dreyer D, Kishkinev D, Wild JM et al (2009) Visual
but not trigeminal mediation of magnetic compass information in
a migratory bird. Nature 461:1274–1277
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