In 1899, Mortensen initiated the systematic marking of wild birds
with numbered leg bands (Wood 1945), but apart from the use of mod-
ern alloys and color bands there have been few major developments in
bird-band technology. Most types of bird bands used today still require
the recovery of a dead bird or the recapture of a living bird in order
to identify the band, even though these two sources of information have
obvious disadvantages for the study of non-sedentary free-living birds.
Many individual life histories are derived from banded birds being shot
or found dead and hence for each bird there are only two records, and
where an individual is retrapped at a particular colony during breeding
seasons there are no data on where the bird went during the non-
breeding season.
In this paper we report results from the use of bands with large
numerals which were developed to enable individual identification of
free-living Australian Silver Gulls (Larus novaehollandiae) at distances up
to 50 m. The effectiveness and durability of the bands are evaluated,
the nature and frequency of errors detected in field observations dis-
cussed, and the quality of the data returned is compared with that from
other studies which used standard bands.
MATERIALS AND METHODS
Description of the number bands and color bands.--Carrick and Murray
(1970) described the manufacture and specifications of the bands they
developed to enable positive identification of free-living Silver Gulls. In
brief, number bands were made from 1 mm thick aluminum alloy, and
when flattened were 16 mm high x 25 mm. A unique 5-digit number
was impressed into the band four times, the impressions were coated
with black enamel paint, and the bands then sprayed twice with "Scotch-
lite Finishing Clear." When a number band was fitted to the leg of a
gull, the numbers read vertically (Fig. 1), and the whole sequence could
be seen from any angle. A return address for the bands was impressed
onto the inside. Color bands were made from 0.9 mm thick commercial
aluminum sheet, cut to 13 mm high x 24 mm for 1-color bands or 19
mm high x 24 mm for 2-color or 3-color bands. "Scotchlite" reflective
colors and "Scotchcal" non-reflective black or white were heat-bonded
FIGURE 1. Examples of number bands recovered from Silver Gulls: 61942, taken from
chick which died on nest; 15567, standard CSIRO 080 series band; 63180, recovered
after 9 yr 7 mo; 61029, recovered after 12 yr 11 mo; 51264, from dead chick on
nest; 59638, 8 mo; 65966, 9 yr 10 mo; 66192, 9 yr 10 mo; 63583, 11 yr 10 mo. The
large 081 aluminum bands are 16 mm (high) x 25 mm, I mm thick, and were sprayed
with Scotchlite Finishing Clear (c) after coating the number impression with black paint.
These bands read from top to bottom. As can be seen, most wear occurs at the bottom,
and it is recommended that bands with vertical number sequences should be placed
on birds to read from bottom to top.
to the outside of the bands in various color and sequence combinations
(Fig. 2). (Scotchlite (c), Scotchcal (c) and Scotchlite Finishing Clear (c) are
registered trademarks of the 3M Company.) White Scotchcal was heat-
bonded to the inside of color bands to increase durability of the alu-
minum.
Gull banding.--Between June 1967 and November 1970, 17,410 Sil-
ver Gulls were banded in South Australia, using the bands described
above (Table 1). Some 15,950 gull chicks were banded on the nest at
15 island breeding colonies, 173 juveniles were clap-netted and banded
in the Beachport area, and 1287 adult gulls were banded after nest-
trapping or clap-netting: 142 on islands at Encounter Bay and 1145 in
the Beachport area (Fig. 3). Each banded gull was given a unique number
band on one leg and a color band on the other. Some 467 types of color
bands were used, being various combinations of blue, green, red, yellow,
black, and white, size of band, vertical, horizontal, or oblique color
sequencing. One-color and 2-color combinations were used for mass-
banding of groups (e.g., each of 3530 chicks hatched on Penguin Island
in Spring 1968 was given a 13 mm high yellow band on the right leg,
referred to in text as R-Y), while 432 unique 3-color bands were used
FICuRE 2. Examples of color bands used on Silver Gulls. RWR, new; YGBk, 1 yr 1 mo;
YG-(--YGR), 8 yr 11 mo; BkGBk, 9 yr; GBk-(-GBkB), 10 yr; Bare metal (--WRW),
11 yr 9 mo; vBkW, new; vBkW, 12 yr 10 mo; WG, new; WG, 12 yr 2 mo; Y, new; Y,
12 yr 9 mo; v, each color runs vertically; B, blue; Bk, black; G, green; R, red; Y,
yellow; W, white. The large bands are 19 mm (high) x 24 mm; the two small bands
are 13 mm (high) x 24 mm. The color materials are Scotchlite (c) or Scotchcal (c) heat-
bonded to aluminum.
on nest-trapped breeding adults (e.g., only one gull had the band black/
red/green on the left leg, and one gull had the same color combination
on the right leg. These are referred to as L-BkRG or R-BkRG).
Band resightings and field records.--During October 1968-August 1981,
teams of professional ornithologists made extensive searches of south-
eastern Australia to find and identify these banded gulls. During 1969,
1970, and 1971, the teams spent 848 days searching for banded gulls,
but only 56 days during 1972-1981. About 60 occasional observers,
scattered through South Australia and Victoria, also watched for banded
gulls, and members of the public returned bands, found on dead or
injured gulls, to the Australian Bird-Banding Scheme. The professional
ornithologists concentrated on shores and coastal areas between Ade-
laide, South Australia, and Cowes, Victoria (Fig. 3) with occasional forays
to more distant areas where banded gulls had been reported.
Banded gulls could be detected at distances up to about 100 m, al-
though it was necessary to be much closer to identify color bands and
read number bands. Usually, a flock of gulls was approached by 2-4
observers in a vehicle suited to the particular terrain. Gulls were not
disturbed unduly by the slow movement of the approaching vehicle, but
TABLE 1. Summary of Silver Gull bandings,June 1967-November 1970, and resightings,
October 1968-August 1981, in south-eastern Australia.
Gulls Gulls Total
Banding site a banded resighted (%) resightings
1. ICI, St. Kilda, Adelaide 2178 40 1.84 77 1.93
2. Outer Harbour, Adelaide 1780 59 3.31 108 1.83
3. West Island 1331 113 8.48 188 1.66
4. Wright Island 731 70 9.58 121 1.73
5. Lake Mulgundawa 933 40 4.29 52 1.30
6. Seagull Is., Coorong 94 14 -- 46 --
7. Lake Hawdon 2025 385 19.01 1373 3.57
8. Nora Creina 1084 240 22.4 1144 4.77
9. Lake George Islands 1154 388 33.62 1927 4.97
10. Cowrie Rock 536 203 37.87 3812 18.78
11. Pleasant Cove 63 58 92.06 1282 22.10
12. Penguin Island 4446 1629 36.64 12,363 7.60
13. Rivoli Bay 679 609 89.69 13,663 22.44
14. Southend Beach 121 100 82.64 1394 13.94
15. Harrington Rock 7 1 -- 20 --
16. Cullens Rock 79 9 -- 54 --
17. Mounce and Battye Rocks 151 17 11.26 78 4.59
18. Carpenters Rock 18 2 -- 72 --
Totals 17,410 3977 22.84 37,774 9.50
a Colonies and banding sites 9-14 were within the intensive study area of the resighting
program.
b g = mean number of resightings per resighted gull.
were always wary of observers on foot. With favorable conditions, num-
ber bands could be read with the unaided eye at distances of less than
2.5 m, with binoculars from 2.5-20 m, and with telescopes from 18-
50 m distance. Color bands could be identified at up to 60 m. These
approximate maximum distances for legibility depended on quality and
clarity of the band, weather and light conditions, behavior of the banded
gull at the time of observation, the individual observer's visual acuity
and experience with reading gull bands, and the quality of the optical
equipment. If possible, bands were read and checked by several ob-
servers. Where a digit or color remained in doubt after a gull had left
the observation site, an appropriate coded comment, which remained
with the record, was made in the field notebook (e.g., 6(5)047 signified
that the digit 5 was doubtful). A resighting record was considered valid
if the number band, color band, and leg sequence of bands recorded at
the time of sighting corresponded with the banding records. Where
discrepancies were found between observation records and banding rec-
ords, a subjective decision was made whether to reject the observation,
whether to accept it as possibly valid but requiring further field confir-
mation, or to accept it as valid despite the discrepancy. This decision
was based on consideration of the combination of number band and
o 100km
I J
3
5
6
9
10
11
12
,- ...................
13
14 FAIRY
15
16 18
17
1. ICl, St. Kilda, Adelaide
2. Outer Harbour, Adelaide
3. West Island
4. Wright Island
5. Lake Mulgundawa
6. Seagull Island, Coorong
7. Lake Hawdon
8. Nora Creina
9. Lake George Islands
10. Cowrie Rock
I 1. Pleasant Cove
12. Penguin Island
13. Rivoli Bay
14. Southend Beach
15. Harrington Rock
16. Cullens Rock
17. Mounce and Battye Rocks
18. Carpenters Rock
FICURE 3. Banding locations of Silver Gulls in southern Australia, 1967-1970.
color band observed, the place of sighting, and the nature of the dis-
crepancy (Table 2).
By 1978, banded gulls were much scarcer, and most number bands
and all color bands were showing signs of wear. To provide further
quantitative data on the effectiveness of these bands and the band re-
sighting program, a sample of the surviving banded gulls (N = 237) was
shot in 1978 and 1981. Although selection of the gulls to be shot was
not based on formal randomization (for reasons of both public safety
and local government regulations), the gulls were shot during all daylight
T^mE 2. Examples of appraisal of incorrect sighting records of Silver Gulls.
Num-
ber
of Correct record
Incorrect record similar
color possibilities* Decision
Left Right bands Left Right Left Right
leg leg used leg leg Comment leg leg
53724 GBG 1 GBkG 53724 This bird was re- accept as:
57358 GBG ported as found dead GBkG 53724
GBG 57433 at a village 40 km
57364 GBKG from banding place.
The leg order was
incorrectly report-
ed by the finder, and
one of the profes-
sional team incor-
rectly recorded
GBkG as GBG.
53757 GYW 1 BkYW 53757 At least two errors reject record
53744 GYW involved.
GYW 57411
vYG (53970) 66 vYG 55970 At time of record, accept as:
observer noted un- vYG 55970
certainty about num-
ber, possibly because
the bird flew away
before confirmation
was possible. Confu-
sion of 5 and 3 was a
common error. (In
this instance band
53970 was not a pos-
sibility since it had
not been used.)
55927 66 55927 vYG The bird was stand- accept as:
ingon oneleg, which 55927 vYG
carried the number
band. Left and right
legs were often con-
fused when only one
leg could be seen.
L-55927 had been
seen on 9 occasions
at the place of the in-
correct sight record.
GWR
56787 1 56787 vGW At least two errors reject record
GWR 57420 involved.
57351 GWR
57304
Bk-W 6 57304 BkRW Middle color strip accept as:
(red) had faded or 57304 BkRW
worn away.
TABLE 2. Continued.
Num-
ber
of Correct record
Incorrect record similar
color possibilities* Decision
Left Right bands Left Right Left Right
leg leg used leg leg Comment leg leg
(573(7)6) BkGW 1 57316 BkGW This band number, accept as:
57376 BRBk and especially the 57316 BkGW
BkGW 53751 fourth digit, was
noted to be doubtful
at the time of obser-
vation. R-BkGW was
a unique band/leg
combination.
Y 60296 3530 60296 Y Leg reversal. accept as:
6O296 Y
63745 W 750 63745 RW This incorrect re- accept as:
53780 WWW cord came 5 yr 5 mo 63745 RW
WWW 65748 after banding. Pre-
vious observations
indicated that R fad-
ed or wore off within
2-4 yr.
66681 Y 3530 GWG 66681 Unlikely that R-Y reject record
was confused for L-
GWG. More proba-
bly 1 or 2 digits of
the number were in-
correct (for in-
stance, could have
been L-56581).
Some 3530 chicks
were banded R-Y,
hence the bird
sighted could be one
of many possibili-
ties.
69300 Nil 463 69300 WG This bird lost the accept as:
color band within 4 69300 (WG)
months. It had 41
subsequent confir-
matory resightings.
* B -- blue; Bk = black; G --- green; R -- red; W = white; Y = yellow; v --- an individual
color runs vertically on the band, otherwise assume individual colors run horizontally
around band (see Fig. 2). Hence, L-GYW indicates green over yellow over white on the
left leg.
110 -
100-
90
"5 80
Q. 70-
ß
60-
o
z,O-
3o-
ß
ß
,.
'o .,../. ., . . .
ß ..-.:'.:. ...". .
0 '
Time between banding and last record(years)
FlouRs 4. Correlation between the time Silver Gulls were known to be alive and the
number of records for individual gulls, for a sample of 188 breeding adults banded
at Beachport, South Australia, 1968 and 1969 (rxy = .385).
hours and at many different sites in order to achieve a representative
sample of survivors. Recovered number bands were washed, dried, and
weighed to the nearest 0.001 g on an analytical balance. Where the
serial number of a band could not be identified, bands were opened,
flattened, washed in water, then 100% ethanol and finally xylene. After
drying, the bands were swabbed with a solution of 30% hydrofluoric
acid, 15% nitric acid, and 55% glycerine (Villella's solution). The alu-
minum affected by stamping of the serial number at the time of band
manufacture etched away faster than the surrounding metal, and the
digits were revealed, even on number bands which had been worn smooth.
Recovered color bands were carefully examined for remaining traces
of Scotchlite or Scotchcal, but were not treated further.
RESULTS
There was a significant (r = .385; P < .01) positive correlation be-
tween number of records per individual gull and length of time that
individual gulls were known to be alive; however, there was great vari-
ation in the frequency with which individual gulls were resighted (Fig.
4). Of the 17,410 Silver Gulls banded, 3977 (22.8%) were seen again,
T^BLv. 3. Band recoveries from 237 Silver Gulls banded in 1967-1970 and shot in 1978
and 1981.
Condition of bands No. %
Both bands, legible
Both bands, number only legible
Both bands, color only legible a
Both bands, neither legible a
Number band only, legible
Number band only, illegible a
Color band only, legible b
Color band only, illegible b
116 48.9
53 22.4
4 1.7
1 0.4
21 8.9
1 0.4
36 15.2
5 2.1
Totals 237 100
a All illegible number bands were readable after acid etching.
b By 1978 most Scotchlite and Scotchcal had worn off the aluminum bands. Where all
traces of color were gone, the color band was considered illegible.
and of 6999 gulls banded in the intensive study area, Lake George to
Southend, 2987 (42.7%) were seen again (Table 1). A total of 37,774
valid sightings was accepted to August 1981 (R = 9.5 resighting records
per resighted gull): 35,189 resightings from the professional teams and
2585 from the occasional observers. Some 26 gulls had > 100 records
and 61 gulls had histories spanning > 12.75 yr from banding.
Loss and wear of bands.--Loss and wear data were derived: (1) from
general observations on the condition of bands while resighting banded
gulls, (2) from specific observations on 246 adult gulls in the Beachport
intensive study area, which were given unique 3-color bands in addition
to their unique number bands, and (3) from a sample of 411 bands taken
from 237 gulls shot in 1978 and 1981.
Some band losses were discovered within 4 months of banding, al-
though until the end of 1971 the total incidence of gulls known to have
lost either band was considerably less than 0.5%. By 1978, 5.4% of known
surviving gulls had lost color bands, and 10.0% had lost number bands.
Of the 246 Beachport adults given unique color bands, 3 (1.2%) were
known to have lost color bands and 5 (2.0%) were known to have lost
number bands by August 1981, after 12-13 yr; however, many of these
adults are thought to have died within a few years of banding. Of 237
banded gulls shot in 1978 and 1981, 41 (17.3%) had lost number bands
and 22 (9.3%) had lost color bands (Table 3). Of the 196 number bands
recovered, 160 (81.6%) were considered to have been still legible on
the free-living gulls, 30 (15.3%) could be read only in the hand, and 6
(3.1%) required treatment with Villella's solution to be rendered legible.
Of 215 color bands taken from the shot gulls, none was clearly identi-
fiable on the free-living gulls, but 157 (73.0%) were identified in the
hand. Some 58 (27.0%) had been worn to bare aluminum, with no traces
of color.
T^BLE 4. Comparison of resighting records from Silver Gulls, shot in 1978 and 1981,
found to have fast-wearing, average-wearing, and slow-wearing bands.
Number Total
of indi- number
vidual of re-
Wear rate gulls Location of resighting records a sighting
of band in records
group group Beach Swamp Town Dump Colony (Z n)
Fast 30 186 (37.0) 61 (12.1) 145 (28.8) 44 (8.7) 67 (13.3) 503
Average 30 172 (32.2) 59 (11.0) 160 (30.0) 70 (13.1) 73 (13.6) 534
Slow 30 191 (34.1) 39 (7.0) 158 (28.2) 44 (7.8) 128 (22.8) 560
Numbers in parentheses are percentages of total number of observations (Z n) in wear
rate group.
All recovered number bands (n = 196 from shot birds + 13 others =
209) were used in the regression of band weight on time (Fig. 5). This
calculation gives a biased estimate of band wear, because the sample
does not include the fast-wearing bands which were lost from gulls (41
of the gulls shot in 1978 and 1981 had lost the number band). However,
it is clear that the mean band wear is at least .0411 g/yr. Inspection of
recovered number bands indicated that properly closed bands were
unlikely to drop off as a result of wear until the bands were lighter than
about .45 g. Since unused number bands weighed .9556 g (___.0124 g
S D), significant band losses (> 5 %) would have occurred from these Silver
Gulls by 9 yr. By 1978, there was a marked variation in the weights of
number bands, with some having lost almost twice as much aluminum
as others (Fig. 5). These differences seem unlikely to be due to differ-
ences in the quality of the metal, but may be due to differences in
behavior of the individual gulls. For instance, on gulls habitually fre-
quenting sandy beaches and shores, bands may wear faster than on gulls
which frequent villages or dumps. To test this, 3 band groups were
selected from the plotted regression of band weights on time: one group
contained the 30 bands showing the fastest rates of band wear, one the
30 slowest rates of band wear, and one the 30 bands showing rates of
wear most similar to the calculated overall mean rate. The age distri-
butions of gulls in each group were similar. The location records for
the gulls carrying these bands were compared. There is significant het-
erogeneity among the 3 groups in the locations where the gulls were
resighted (x22 = 37.7; P < .001). Inspection of the data (Table 4), how-
ever, reveals no clear trends. In part this may be because most resighting
records were made during 1968-1972, whereas much, perhaps even
most, band wear occurred during 1973-1981. If the behavior of indi-
vidual gulls is not consistent over years, the paucity of observations
during 1973-1981 would confuse elucidation of any relationship be-
tween gull behavior and rate of band wear. Even so, this comparison
1'0
0'9'
0.8
0-6-
.o.5
, o.4,
o.3-
0'2-
0-1
FIGURE 5.
Time from banding to band recovery(years)
Regression of weight of number band (n = 209) on time (years). Regression
equation is y -- .9556 - .041 lx (years).
does indicate considerable potential for field study of band wear and
the reasons for differential band wear.
With color bands, the problem was not so much one of when bands
were lost, but rather when the color of the Scotchlite or Scotchcal was
not identifiable. Red Scotchlite showed some fading in 9 months, marked
fading by 2 yr, and appeared white by 3 yr. All other color materials
showed deterioration and evidence of peeling from the aluminum base
by 4 yr. By 1978, no color bands had sufficient color left to enable
positive identification on the free-living gulls. In the hand, from the
1978 and 1981 shootings, the proportions of color bands which could
be identified positively ranged from ( 10% for bands originally bearing
some red (e.g., WR), 16% for unique 3-color bands (e.g., BBkG), 54%
for Y, and 73% for WG. Red Scotchlite appeared to wear away faster
than the other materials, in addition to fading much faster, but with
the exception of that material, the durability of colors on the bands
appeared to be a function of the actual sizes of material bonded to the
aluminum base. The larger the individual pieces of color material, the
longer color could be identified on the bands.
TABLE 5. Problems detected in 37,774 resightings of number bands on Silver Gulls (68
unresolved errors not included).
Problems a Correct digit
detected 0 1 2 3 4 5 6 7 8 9
Digit not read 5 3 16 43 9 29 27 12 14 19
Digit read as:
0 -- -- 1 ..... 4 1
1 1 .... 1 1 39 -- 3
2 2 -- -- 5 .... 3 5
3 -- -- 2 -- 1 14 3 2 2 --
4 -- 1 2 2 -- -- 1 1 1 --
5 -- -- -- 9 -- -- 16 -- 2 --
6 3 -- -- 5 -- 37 -- -- -- 6
7 -- 15 3 8 2 .....
8 7 -- -- 4 -- 2 4 2 -- 3
9 2 3 4 1 -- 5 8 1 3 --
Recorded as
dubious, but
later checked
as correct 18 26 17 30 28 75 67 24 25 44
Totals 38 48 45 107 40 163 127 81 54 81
Some 329 number bands were misread leading to an incorrect field record. In addition,
68 other resighting records contained possible digit errors in combination with possible
color errors. These multiple errors could not be resolved, and the 68 records were rejected.
Errors in resighting bands, recording and processing data.--Regular checks
during field work and data processing indicated that almost all errors
were made at the time of the original field observation, sometimes by
the recorder, but usually by the observer. The accuracy and efficiency
of most observers improved markedly with an experience of only a few
days, but some observers never improved to the level of accuracy where
their observations were accepted. Common early errors were reading
digits incorrectly (e.g., reading 7 as 1), and reversing the sequence of
two adjacent digits. Such errors sometimes went undetected when a bird
flew away before the bands could be checked by a second observer, or
when the bands were seen at such a distance that only one observer had
sufficiently acute vision to read the bands. For the purpose of this section,
all errors will be considered together, regardless of the source.
The most common error was leg reversal. Some 328 observations
(.87% of all resightings) had the incorrect leg sequence for bands. Leg
reversal errors occurred mainly when a gull, standing sideways to the
observer, was seen near the limit of an observer's ability to read a band,
or when a gull was standing on one leg with the second leg hidden. Leg
reversal errors also occurred when a gull was in the hand: e.g., L-GBkG,
R-53724 was sighted and recorded correctly on 21 occasions over 2 yr,
but on the last occasion it was found dead by a local villager who reported
TABLE 6.
Leg Bands for Visual Identification
Number of problems detected per 10,000 readings of each digit.
[299
Problems Correct digit
detected 0 1 2 3 4 5 6 7 8 9
Digit not read 4.4 2.2 12.4 24.2 7.9 9.1 6.7 7.9 11.7 8.4
Digit read as:
0 -- 0.7 3.3 0.4
1 0.9 -- 0.3 0.2 25.6
2 1.8 -- 2.8 2.5 1.3
3 1.5 -- 0.9 4.4 0.7 1.3 1.7 2.2
4 0.7 1.5 1.1 -- 0.2 0.7 0.8
5 5.1 -- 4.0 1.7
6 2.6 2.8 11.7 -- 2.6
7 11.2 2.3 4.5 1.8 --
8 6.1 2.3 0.6 1.0 1.3 -- 1.3
9 1.8 2.2 3.1 0.6 1.6 2.0 0.7 2.5 --
Recorded as
dubious, but
later checked
as correct 15.8 19.5 13.2 16.9 24.6 23.6 16.6 16.4 20.8 19.4
Totals 33.4 35.8 34.7 60.3 35.2 51.3 31.4 53.9 45.0 35.6
the bands as L-53724, R-GBkG. The team member noting the recovery
added a second error, by recording the bands as L-53724, R-GBG.
Table 5 shows the frequency of digit errors, omissions, and uncer-
tainties in field records. Of 37,774 records, 329 (.87%) were found to
contain digit errors. A further 68 (. 18%) contained errors which could
not be explained, so those records were rejected totally and are not
included in the total of 37,774 records analyzed. Table 6 shows relative
occurrences, corrected for the different frequencies with which digits
occurred on the bands actually used, and then standardised per 10,000
digit resightings. The most common errors were sighting or recording
7 as 1, 5 as 6, and 1 as 7 (Tables 5 and 6). Considering the relative
frequencies of digits on the resighted bands, the digit 3 caused the most
problems, followed by 7 and 5 (Table 6).
With color bands, the most common errors were to read green or
blue bands as black, when the bands were in shadow, and dirtied white
bands as yellow. The most common error made while recording color
bands was to write blue (B) instead of black (Bk). Color bands were
recorded incorrectly 109 times when the bands were known to be in
good condition (.3% error). Once the color bands showed signs of wear
or fading, color band errors and color uncertainties increased markedly.
From 1973 onwards, 841 records had color band discrepancies which
could be attributed to wear or fading (53.7% error). Even so, most of
the worn or faded color bands retained enough identifying color traces
TABLE 7. Comparison of data derived from Silver Gulls banded in 1967-1970 and
recovered in 1978.
Recovery method of band
Non-recaptured
Shooting resighting
Number of birds
Mean number of resighting records
Range of resighting records
Percentage of gulls with only one
record after banding (including
recovery record)
Mean number of days between penulti-
mate record and last record
171 125
17.5 19.0
0-104 0-134
10.5 11.2
1815 1501
to be useful for additional confirmation of the accuracy of number bands
recorded (see Table 2).
Comparison of records from a shot sample with records from a non-shot
sample.--Shooting is a common, effective method for recovering bands
from free-living birds; however, it has the obvious disadvantage of ter-
minating the return of data. With Silver Gulls, it was particularly quick
and easy to shoot a selected individual within a range of 6-60 m. In
order to assess the effectiveness of the visual resighting program, the
histories of the 171 gulls shot during 1978 were compared with the
histories, prior to the start of the shooting program, of 125 gulls which
were seen-only during 1978 (Table 7). The shot sample was taken con-
currently with routine field searches, during otherwise standard obser-
vation procedures. Excluding all records made on the seen-only gulls
after the 1978 shooting program, the shot birds still had slightly fewer
resighting records (2 = 17.5) than the seen-only gulls (2 = 19.0), but in
all of the comparisons shown (Table 7) there were no significant differ-
ences between the data sets (Fig. 6). We conclude therefore that, with
respect to the number of individual gulls "recovered" by the two meth-
ods over the particular trial period, resighting of these bands by trained
observers was as effective as the shooting program. We note, however,
that some of the gulls shot in 1978 had bands which were no longer
readable on the free-living gulls. These were the fast-wearing bands,
and had the intensity of sightings been greater in 1976-1977, it is highly
probable that there would have been significant differences.
By the end of the field program, the range of resighting frequency,
for the 3977 resighted gulls, was 1-204. There were major differences
in distances of dispersal and foraging movements between individual
gulls, even considering gulls from a particular colony and the same age
cohort, which in part accounts for the great range in resighting fre-
quency. However, the proportion of gulls recorded in 1978 for the first
time since banding (18/171 of shot gulls, 14/125 of seen-only gulls:
30-
25.
20
10.
15-
u>,1o
,t-5
B
0 20 0 60 80 100 120 10
Number of records after banding
FIGURE 6. A--Frequency of resighting records for 171 Silver Gulls shot in 1978; B--
Frequency of resighting records for 125 Silver Gulls seen but not shot in 1978. The
gulls were banded in 1967-1970.
10.8% of total sample) suggests that the behavior of these individuals
had changed with time.
Thus, of the two methods, resighting of readable band numbers on
uncaptured birds has overwhelming advantages for Silver Gulls and
similar species: the method leaves the birds unharmed for further re-
sightings and, with trained observers, allows for convenient searches in
areas of urban development. Shooting is quicker, and will always be
effective at greater distances than a band can be read, but it destroys
the potential for future information and is unsafe or prohibited in many
areas of human activity.
DISCUSSION
Following Lack's classic field study of Robins (Lack 1939, 1940), re-
sighting records have been used very successfully to establish the pres-
ence of uncaptured, uniquely color-banded birds in many ecological and
behavioral studies, for example, on corvids (Rowley 1971, 1973), a crac-
ticid (Garrick 1960, 1963, 1972), larids (Garrick and Murray 1970;
Garrick 1972; Mills 1972, 1973, 1979) and a procellariid (Garrick and
Dunnet 1954). Attempts have been made to read standard number
bands on uncaptured, free-living birds (Garrick et al. 1957; Kadlec and
Drury 1968, 1969; Kadlec 1975; Butler et al. 1980), but most number
bands in use today require the banded bird to be in hand for the unique
serial number to be read. To our knowledge, the number bands de-
scribed in this paper are the first developed to enable resighting of the
bands to be used as the primary method of bird identification.
Of 17,410 Silver Gulls given these aluminum number bands, 22.8%
were seen again. These gulls had a mean of 9.5 resightings. Considering
only those 1287 gulls banded as adults, 85.6% were seen again, with a
mean of >20 resighting records per bird. These figures may be com-
pared with returns from other seabird programs which used the con-
ventional types of bands. Recovery rates range from about .1-20%
(Hitchcock and Carrick 1958; Dunnet et al. 1977), with, typically, a
single record after banding for any bird. Even with gulls, which are
amongst the most accessible seabirds, returns from standard bands are
usually about 5%, rarely approach 20% (Table 8), and, again, typically
consist of just a single record after banding. Such limited data can
represent an extremely poor return for the expense and time invested
in the initial banding programs. Using the new bands on Silver Gulls,
we now have many detailed life histories which show the movements of
individuals after they left the banding locations. Some individuals were
identified at their breeding or natal colony and then up to 450 km away
in the non-breeding months, in 3 different years. Over the 14 years of
resightings, 1.1% of number band records were found to contain digit
errors or digit omissions, even with observers in the field checking each
other. This emphasizes that only those actively involved in the details
of such a study can make the final decision of validity of a recovery.
Some 2.5% of all color-band resightings were also in error, but this
includes errors due to wear and fading of colors after about 4 yr. Some
.9% of records showed leg reversal errors. Investigation of the sources
of errors revealed that almost all were made in the field by the observers,
rather than at some later stage; hence, where possible all observations
must be checked and rechecked by 2 or more people. This greatly
minimizes the level of errors, and also identifies any observers who are
prone to make errors. When errors were found in the data files, field
records were compared with original banding records, and only 68 of
37,842 sightings (.2%) had to be rejected because the errors could not
be resolved with a reasonable degree of certainty. Of concern was the
thought that errors could be made which might not be detected. For in-
stance, if bands L-62817 R-Y were incorrectly read or recorded as
L-62811 R-Y, which was another valid band combination, the latter
Vo. 55, o. Leg Bands for Visual Identification [303
record would be accepted unless L-62811 was known to be elsewhere
at the same time, or dead. We can only speculate that, in our study,
undetected errors should be few, and at least an order of magnitude
less than the number of errors detected, because of the large number
of different types of color bands used. Clearly, undetected errors would
be reduced to negligible levels by avoiding multiple use of color bands
to denote cohorts and by using discontinuous number band sequences.
With a banding program of 20,000 birds, it would be best to avoid a
sequence such as 50000-69999 and instead use, for example, 20000-
24499, 45000-49999, 60000-64499, and 75000-79999. These number
bands could be paired with one of the 216 three-color combinations of
black, blue, green, red, yellow, and white, used randomly. The proba-
bility of an erroneous sighting then resulting in another valid number
band and color band combination is minute. Since most band wear
occurs near the foot, we suggest that bands with vertical number se-
quences should be placed on the bird's leg to read from bottom to top.
A most urgent need is to continue the experiments of Kadlec (1975),
and others, to find an alloy that will increase the durability and reada-
bility of bands. For long-lived species of seabirds, exceptionally durable
alloys are needed to last 30 yr or more with intermittent immersion in
the sea, brackish water and mud, and abrasion by sand. Poulding's (1954)
data suggested that Larus argentatus actively removed leg bands, result-
ing in a 50% loss of bands in the first year. Paynter (1966) thought that
band loss from Herring Gulls was probably about 5% per year. Kadlec
and Drury (1968) reported that by the sixth year about 20% of standard
aluminum bands were being lost per year from Herring Gulls. Later
this estimate was amended to 56% loss by the fourth year (Kadlec and
Drury 1969 ), although Kadlec's (1975) most recent figure was 3% loss
to the end of 7 yr. Mills (1972) used various types of bands on the Red-
billed Gull of New Zealand, Larus novaehollandiae scopulinus, and found
significantly fewer lock bands were lost than butt-to-butt bands (24%
versus 89% loss) over 10 yr, but lock bands wore more quickly because
of their imbalanced weight. Spear (1980) found that 23% of number
bands on Larus occidentalis were lost by 3.7 yr, and the loss rate thereafter
was 16% per year. Clearly, the loss rate from gulls is very variable.
Paynter (1966) considered that losses could be due to bands weakening
with wear, bands being removed by the birds, and "an inconsistent or
fluctuating loss" due to such factors as time immersed in saline water
"or because the new bands are not of uniform strength." With the size
and weight of our bands, it seems unlikely that even adult Silver Gulls
(weighing ca. 400 g) could remove bands unless the metal had inherent
defects or the banding technique was faulty. Removal of defective bands
by the gulls would explain the low early losses noted. By 9-10 yr, 10%
of surviving Silver Gulls were known to have lost number bands and
5% were known to have lost color bands. It is possible to recapture
banded birds to replace worn or lost bands, but this sometimes results
in bird mortality, which we wished to avoid. Number bands of the type
we describe would have increased durability, and readability, on larger
birds. Relatively increased in size for use on Herring Gulls, for instance,
duration and distance of readability should be doubled on the figures
we give for Silver Gulls.
The lower incidence of loss of color bands, compared with number
bands, was attributed to the Scotchlite and Scotchcal bonded to the
outside and inside surfaces of the aluminum, protecting the metal from
wear and corrosion. Even so, there were obvious differences in wear
rates of similar color bands on different gulls, as for the number bands.
We have not been able to explain this adequately, although we have
suggested that it may be a consequence of individual gull behavior.
Durability of number bands might be improved by cementing a plastic
layer to the inside, as we did for the color bands, finding a durable clear
coating that could be sprayed or painted on the outside of the bands
before they are placed on the gulls, and increasing the band size and
thickness when used on larger birds.
We consider that the results of this program give an unequivocal
indication of the effectiveness and usefulness of number bands which
are readable on free-living birds, and also of the value of combining
these number bands with clearly visible color bands. Our band return
rates were far better than those which have been achieved by most other
large-scale gull-banding programs (Table 8), approaching the returns
that could have been achieved by actively seeking out and then shooting
the banded gulls. A major and obvious advantage of "visual recapture"
is that it is a non-destructive method of sampling: once banded, the gulls
need not be handled or disturbed again. Another equally important
advantage is that part-time observers can be trained and encouraged to
read the bands, needing only binoculars and notebooks instead of trap-
ping equipment or guns. Many valuable records came from part-time
observers who were scattered throughout southern Australia, and who
were prepared to search their home ranges on a regular basis looking
for the banded gulls. Having color bands and number bands on the
birds was especially essential here, because most of the part-time ob-
servers were alone when they were reading bands, so the match of color
and number with the banding record was the only check of validity. As
colors faded or wore away, the advantage of having single bands with
3 colors, rather than 3 individual bands each with a single color, was
demonstrated. Where one color of a sequence was lost from a 3ocolor
band (e.g., L-BkRG becoming L-Bk/metal/G), there were far fewer
original combinations possible (6) than if 1 of 3 single-color bands had
been lost (e.g., if L-Bk/R/G became L-Bk/R, there are 18 possibilities
for the original combination. The problem of loss of colors may be
resolved by bonding a few layers of the color base to the band so that
a fresh layer is revealed as each outer layer becomes abraded. Anderson
(1980) has indicated that polyvinyl chloride (Darvic) color bands could
be more durable, although at present these are not available commer-
cially.
SUMMARY
Carrick and Murray (1970) described a new type of aluminum band that was designed to enable identification of large numbers of uncaptured, free-living birds. During 1967-1970, these number bands, used together with color bands on the other legs, were placed on 17,410 Silver Gulls in South Australia. The number bands had a maximum readable life of about 10 yr on these free-living gulls, although after this period the numbers could be revealed on smooth bands by etching the aluminum with Villella's solution. The color bands used had a maximum readable life of about 3-4 yr. Our results from these bands indicate enormous potential for studies using readable number bands, with returns far exceeding those from conventional bands. The nature and frequency of the errors detected in data files are discussed. Errors were made mostly during field observations, rather than subsequently, and the necessity is emphasized to continuously check and recheck all sighting records while the banded bird is still present at a location. Because all gulls had 2 bands, a number band and a color band, both of which were readable on the uncaptured birds, less than .2% of records were rejected as a consequence of unresolved errors. Undetected errors were probably few, but these could be reduced to negligible levels by the appropriate use of color bands to confirm the accuracy of reading of number bands. To our knowledge, this is the first study that has used resightings of number bands on free-living, unrecaptured birds as the primary method of data acquisition.
ACKNOWLEDGMENTS
We gratefully acknowledge the assistance of Mawson Institute staff
with banding, field observations, data processing and logistical support:
D. Creighton, D. Grey, D. Ferrett, S. R. Harris, S. E. Ingham, G. Jacobs,
D. MacKenzie, and P. D. Shaughnessy. National Parks Rangers G. Har-
rington and I. May also assisted with field work. Our unpaid, part-time
field observers included G. Bowker, C. Carrick, P. Fox, W. R. Garrett,
I. Savage, V. Yeoman, and over 50 more. We thank them all. The help
of D. Purchase, Secretary, Australian Bird-Banding Scheme, is acknowl-
edged. Permission to conduct field work was given by the South Aus-
tralian National Parks and Wildlife Service. Professor G. M. Dunnet,
University of Aberdeen, gave very helpful criticisms of the manuscript.
We especially thank Dr. F. Jacka, Director of the Mawson Institute
for Antarctic Research, for support during the entire project. Grants
and support came also from the Australian Research Grants Committee,
the University of Adelaide, the Australian Commonwealth Scientific
and Industrial Research Organization (CSIRO), Flinders University of
South Australia, and the Utah Foundation.
LITERATURE CITED
ArmRso, A. 1980. The effects of age and wear on color bands. J. Field Ornithol. 51:
213-219.
BUTLER, R. W., N. A.M. VERBEEK, AND R. G. FOOTTITß 1980. Mortality and dispersal
of the Glaucous-winged Gulls of southern British Columbia, Canadaß Can. Field Nat.
94:315-320ß
CARRICK, R. 1960. The Australian bird-banding scheme, 1953-1960. Australian Mus.
Mag. 13:222-227ß
ß 1963. Ecological significance of territory in the Australian Magpie, Gymnorhina
tibicen. Proc. Int. Ornithol. Congr. 13:740-753.
--. 1972. Population ecology of the Australian Black-backed Magpie, Royal Penguin,
and Silver Gull. In Population Ecology of Migratory Birds: a Symposium. U.S. Dep.
Int., Wildl. Res. Rep. 2:41-99.
--, AND G. M. DUNNET. 1954. Breeding of the Fulmar Fulmaris glacialis. Ibis 96:
356-370ß
, AND M.D. MURRAYß 1970. Readable band numbers and "Scotchlite" colour bands
for the Silver Gull. Australian Bird Bander 8:51-56.
--, W. R. WHEELER, AND M.D. MURRAYß 1957. Seasonal dispersal and mortality in
the Silver Gull, Larus novaehollandiae Stephens, and Crested Tern Sterna bergii Li-
chenstein, in Australia. CSIRO Wildl. Res. 2:116-144.
DUNNET, G. M., W. R. P. BOURNE, J. C. COULSON, M.P. HARRIS, C.J. MEAD, H. MILNE,
J. L. F. PARSLOW, C. M. PERRINS, R. C. STEELE, AND M. A. BOOTH. 1977. The report
of a working group on ecological research on seabirds. Natural Environment Res.
Council (U.K.) Publ. Series C, No. 18, 48 pp.
HITCHCOCK, W. B., AND R. CARRICK. 1958. Fourth annual report of the Australian Bird-
Banding Scheme, July 1957-July 1958. CSIRO Wildl. Res. 3:115-141.
HOFSLUND, P. B. 1959. Fall migration of Herring Gulls from Knife Island, Minnesota.
Bird-Banding 30:104-114.
KADLEC, J. A. 1975. Recovery rates and loss of aluminum, titanium, and incoloy bands
on Herring Gulls. Bird-Banding 46:230-235.
--, AND W. H. DRURY. 1968ß Structure of the New England Herring Gull populationß
Ecology 49:644-676.
--, AND --. 1969. Loss of bands from adult Herring Gullsß Bird-Banding 40:
216-221.
LACK, D. 1939. The behaviour of the Robinß I. The life history, with special reference
to aggressive behaviour, sexual behaviour and territory. II. A partial analysis of
aggressive and recognitional behaviourß Proc. Zool. Soc. 1091:169-200ß
ß 1940. The behaviour of the Robinß Population changes over four years. Ibis 4:
299-324.
MiLLS,J. A. 1972. A difference in band loss from male and female Red-billed Gulls, Larus
novaehollandiae scopulinus. Ibis 114:252-255.
1973. The influence of age and pair-bond on the breeding biology of the Red-
biiled Gull, Larus novaehollandiae scopulinus. J. Anim. Ecol. 42:147-162.
. 1979. Factors affecting the egg size of Red-billed Gulls, Larus novaehollandiae
scopulinus. Ibis 121:53-67ß
MURRAY, M.D., AND R. CARRICK. 1964. Seasonal movements and habitats of the Silver
Gull, Larus novaehollandiae Stephens in south-eastern Australiaß CSIRO Wildl. Res.
9:160-188.
OLSSON, V. 1958. Dispersal, migration, longevity and death causes of Strix aluco, Buteo
buteo, Ardea cinerea and Larus argentatus. A study based on recoveries of birds ringed
in Fenno-Scandia. Acta Vertebratica 1:91-189.
PARSONS, J., AND N. DUNCANß 1978. Recoveries and dispersal of Herring Gulls from the
Isle of May. J. Anim. Ecol. 47:993-1005.
PAYNTER, R. A. 1966. A new attempt to construct life tables for Kent Island Herring
Gulls. Bull. Mus. Comp. Zool. Harvard Univ. 133:489-528.
POULDING, R. U. 1954. Loss of rings by marked Herring Gulls. Bird Study 1:37-40.
ROWLEY, I. 1971. Movements and longevity of Ravens in south-eastern Australiaß CSIRO
Wildl. Res. 16:49-72.
ß 1973. The comparative ecology of Australian corvids. I. Introduction, study areas,
and methods of study. CSIRO Wildl. Res. 18:1-23.
SMITH, W.J. 1959. Movements of Michigan Herring Gulls. Bird-Banding 30:69-104.
SORENSEN, L. H. 1972. An analysis of Common Gull, Larus canus, recoveries recorded
from 1931 to 1972 by the Zoological Museum in Copenhagen. Gerfaut 67:133-160.
S'E^R, L. 1980. Band loss from the Western Gull on Southeast Farallon Island. J. Field
Ornithol. 51: 319-328.
S'rNCrR, R. 1971. Report on bird-ringing for 1969. Br. Birds 64:137-186.
THRrLF^LL, W. 1978. Dispersal of Herring Gulls from the Witless Bay Sea Bird Sanctuary,
Newfoundland. Bird-Banding 49:116-124.
Woon, H. B. 1945. The history of bird-banding. Auk 62:256-265.
(JRO) Mawson Institute for Antarctic Research, University of Adelaide, North
Terrace, Adelaide, S.A. 5000, Australia (Current address: Department of Con-
servation and Environment, 1 Mount St., Perth, W.A. 6000, Australia),' (RC)
Mawson Institute for Antarctic Research, University of Adelaide, North Terrace,
Adelaide, S.A. 5000, Australia (Current address: "Achandean," Meadows
Road, Dornoch, Sutherland, Scotland); (MDM) Division of Tropical Animal
Science, CSIRO, McMaster Laboratory, Private Bag No. 1, P.O., Glebe, N.S.W.
2037, Australia. Received 7 Oct. 1984; accepted 28 Mar. 1984.