Sugawa et al.: Breeding site fidelity of Streaked Shearwater

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LONG-TERM TRENDS IN BREEDING SITE FIDELITY
OF STREAKED SHEARWATER CALONECTRIS LEUCOMELASHISASHI SUGAWA1, KIYOTAKA KARINO1, AKIO OHSHIRO1 & MASASHI HIRAI1,2
1Bird Banding Association, c/o Bird Migration Research Center, Yamashina Institute for Ornithology,
2115, Konoyama, Abiko, Chiba, 270–1145, Japan
Kyoto Prefectural University, 74 Oji, Kitainayazuma, Seika, Kyoto 619–0244, Japan (mhirai@kpu.ac.jp)
Submitted 24 March 2013, accepted 14 October 2013
SUMMARYSUGAWA, H., KARINO, K., OHSHIRO, A. & HIRAI, M. 2014. Long-term trends in breeding site fidelity of Streaked Shearwater
Calonectris leucomelas. Marine Ornithology 42: 11–15.
Site fidelity of Streaked Shearwater Calonectris leucomelas on Kanmurijima Island, Sea of Japan (35°40vN, 135°26vE) was analyzed basedon a 27-year banding record from 1984 to 2010. Two study sites were set in this colony, and return fidelity to the initial release site wasassessed. Most returns occurred at sites where they had been released. Only small percentages (0.7% and 0.2%) of the returns were recordedin the other study sites. Site fidelity was further analyzed using a 10 m × 10 m grid at one study site. Returning birds showed high site
fidelity. The rate of returns within 10 m of the release place among the total returns was defined as the site fidelity index. Based on a total of
4 154 returns, the index was calculated to be 0.73. The index was almost unchanged with increasing duration between the first release andthe return. Even after 20 years, birds returned to the vicinity of the initial release site. The annual change in the index fluctuated from 0.54
to 0.81, with a slight increase during the most recent study period.
Key words: banding and recapture, Kanmurijima Island, Streaked Shearwater, colony, long-term study, breeding site fidelity

INTRODUCTION
Streaked Shearwater Calonectris leucomelas is a large shearwaterof the Pacific Ocean (Warham 1990). Breeding colonies occur on
offshore islands in the northwestern Pacific Ocean, the Sea of Japanand the East China Sea (Kondratyev et al. 2000, Oka 2004). Outsideof the breeding season, the birds move to warmer areas in theequatorial Pacific, where they winter (Blaber & Milton 1994, Onley& Scofield 2007, Takahashi et al. 2008). The global population
is estimated to be ca. 3 000 000 individuals, and is thought to be
decreasing (Birdlife International 2012). However, the breeding
biology of this species is not well known (Yoshida 1962). Because
eastern Asia is highly populated by humans and has substantial
industrial fishing activity, human disturbance of breeding could
be significant. Disturbances may include the introduction of rats
to colonies, marine pollution, fishing gear bycatch and changes inthe environment of the colony site due to direct human activities(Everett & Pitman 1993, Tasker et al. 2000). The oil spill from a
Russian tanker, Nakhodka, in the Sea of Japan in 1997 resulted insevere damage to seabirds (Helm et al. 2006). Some of the smallsized colonies of this species on offshore islands in Japan havealready been lost (Oka 2004). A colony of this species in Korea wasseverely damaged by the introduction of rats (Lee & Yoo 2002).
Although Streaked Shearwater is classified as “Least Concern”(Birdlife International 2012), breeding status should be monitored
and knowledge of breeding biology should be increased toward
developing an effective conservation program. Although fidelity
to the nesting site is observed in a wide range of Procellariiform
species (Harris 1972, Croxall 1979, Ristow et al. 1991, Thibault
1994, Rabouam et al. 1998, MacDonald 2009, Gauthier et al.
2010), to the best of our knowledge it has not been investigated in
Streaked Shearwater.

Kanmurijima Island, also written Kammuri-jima Island (35°40vE,
135°26vN), an uninhabited island in the Sea of Japan, about 23 km
north of Maizuru City, Kyoto, is home to one of the largest colonies
of Streaked Shearwater (Yoshida 1962, 1981, Oka 2004). The colony
has long been known and protected by local fishermen because flocksof this bird indicate the presence of schooling fish. The island was
designated a Natural Monument by the Japanese government in 1924,
and landing there is prohibited by law except for scientific and religious
purposes. Another uninhabited island, Kutsujima Island, which lies
about 4 km north, has a small colony of this species (Sato et al. 2010).
Yoshida (1962) clarified the breeding cycle of Streaked Shearwateron Kanmurijima Island and banded a small number of the birds.
Continuous banding began in 1971, but the results have not yet beenreported. Here we report site fidelity of Streaked Shearwater to theirbreeding island and any changes in fidelity over a period of 25 years.
STUDY AREA AND METHOD
Study area
The top and low flat areas of Kanmurijima Island are covered bynatural evergreen forests, dominated by Persea thunbergii, mixed
with some Castanopsis cuspidata and Ilex integra (Maesako 1999).
Elsewhere are steep rocky cliffs, which are almost bare. Island area isabout 22.3 ha; the highest point is 169 m (Fig. 1). Soil under the forest
is deep and is suitable for burrow-digging seabirds. Two study sites,
Site A (ca. 1.0 ha) and Site B (ca. 0.1 ha) were set under a tree canopyon the southeastern part of the island. Site A extended from near sea
level to 50 m; Site B was set on flat areas near sea level (Fig. 1). A
grid system was established within Site A by dividing the site into
10 × 10 m quadrats. Site B was used for banding without setting
quadrats. From center to center, these two sites are about 200 m apart.

Marine Ornithology 42: 11–15 (2014)

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Sugawa et al.: Breeding site fidelity of Streaked Shearwater

Bird banding

Calculation of distance between release and recapture points

Birds were banded and recaptured by hand, mainly on the groundsurface at night when the birds returned or were moving to thelaunching point. Some birds in burrows were also captured andbanded. The location information, time and bird age (chick or adult)
were recorded. A “return” was defined as a recapture in any later
breeding season. Only the first recaptures of the given breeding
season were analyzed for this study. Hence, “number of returns”
does not mean the number of returning birds, because some birds
returned more than one time. The recaptures within the same
breeding season were treated as “repeat recaptures,” and were notused in the present analysis. Data on the banding of adult birds from1984 to 2010 were used.

Our location data were based on quadrats, not on burrows.
The distance between the banding and returning points wascalculated on the assumption that the birds were caught in thecenters of quadrats in question. When the bird was recaptured in
the same quadrat, the distance was recorded recorded as 0 m. In
the case of directly or diagonally adjacent quadrats, the distanceswere 10 m or 14.1 m, respectively.

A total of 19 172 birds were newly banded from 1984 to 2010.
However, 2 062 of the banded birds lacked banding site information,and these data were discarded. Among the remaining 17 110
birds, 16 446 were banded as adults and were included in thepresent analysis. A substantial portion of these banded birds
returned to this island in later breeding seasons. Details of the return
rate will be reported elsewhere.
Data selection
Analysis of return fidelity between the two study sites. Banding
continued from 1984 to 2010 at Site A, but at Site B it started in1988 and finished in 1997. Therefore, banding and recapturing dataof both sites from 1988 to 1997 were used in this analysis. During
this period, a total of 4 103 returns were recorded at both sites.
Return site fidelity within Site A. Return fidelity of breeding birds
was further analyzed using the quadrat data at Site A. Birds banded
as adults and released from 1984 were used in the analysis. The
banding data in 1986 lacked quadrat information and were notincluded. A preliminary analysis showed an obvious effect of
movement to the launching point, and such movement started after
midnight. Since the purpose of the present analysis is to describe
the fidelity to the nesting site, we excluded the capture data from24h00 to 05h00. Capture data for the quadrat at the launching point
were also excluded.

Fig. 1. Map of Kanmurijima Island and the study sites, based on a
map by Geospatial Information Authority of Japan.

RESULTS
Return analysis
As a result of data selection, a total of 4 103 returns were analyzed.
The birds released at Site A produced 1 586 returns. Most of the
returns were those to the same site, but 11 returns were recorded in
Site B (0.7%). A total of 2 517 returns were recorded among birds
released at Site B. However, four were recovered in Site A (0.2%)(Table 1).
In order to examine fidelity within Site A, the selected 4 154
returns were further used to calculate the distance between thequadrat where the bird was newly banded and the quadrat to which
it first returned. Among the 4 154 analyzed returns, about 44%
were recorded in the same quadrat, and 73% (3 017 returns) wererecorded within 10 m of the release point (Fig. 2).
TABLE 1
Summary of returns to the two study sites, 1988–1997(see text for definition of “return”)
Banding site toreturn site

No. of returns% of returns

A to A

1 575

99.3

A to B11

0.7

B to A4

0.2

B to B

2 513

99.8

Total

4 103

Fig. 2. Breeding site fidelity of Streaked Shearwater. A total of
4 154 returns at Site A were classified based on the distance
between banding and recapture points.

Marine Ornithology 42: 11–15 (2014)

Sugawa et al.: Breeding site fidelity of Streaked Shearwater
The rate of return within 10 m of the release point among thetotal number of returns is defined as the site fidelity index. In its
estimation, the return data were grouped based on duration between
the first release and recapture (Table 2). Even after 15 years, we
obtained 38 returns showing a high site fidelity index. As ourpresent longest record, five birds were recaptured 20 years after the
first release: two in the same quadrat, two in the next quadrat, andone in a diagonally adjacent quadrat (Table 2). The fidelity index
was mostly constant for 20 years. The regression slope was not
significant (P = 0.30, data not shown).
To determine the annual variation in site fidelity index, the index
for returns in each breeding season was calculated (Table 3). The
fluctuation of the index was rather large during the 1980s, butlater became smaller. The index as a whole gradually increased
during the study period. The regression slope was 0.0063(r2 = 0.53, P < 0.05) (Fig. 3).
DISCUSSION
The present data show a high rate of site fidelity in Streaked
Shearwater, as reported in other seabird species (Harris 1972,
Ristow et al. 1991, Thibault 1994, Rabouam et al. 1998). Most of
TABLE 2
Effect of time elapsed between release andrecapture on the site fidelity index
Duration, Total no. Returns Returns Returns
years
of returns to the within 10 within
m (B)
20 m
(A)
same
quadratSitefidelity
index
(B/A)

1

960

466

738

855

0.77

2

697

318

520

595

0.75

3

557

240

385

452

0.69

4

393

162

272

320

0.69

5

294

122

205

250

0.70

6

262

113

190

211

0.73

7

192

96

141

168

0.73

8

140

52

102

115

0.73

9

132

58

96

108

0.73

10

101

42

69

79

0.68

11

86

35

62

75

0.72

12

83

30

56

66

0.67

13

76

30

49

60

0.64

14

56

21

41

49

0.73

15

38

19

27

32

0.71

16

30

11

18

25

0.60

17

18

7

12

15

0.67

18

24

15

21

23

0.88

19

10

3

9

9

0.90

20

5

2

4

5

0.80

4 154

1 842

3 017

3 512

0.73

Total

13

TABLE 3
Annual change in the site fidelity index
Year

Total no. of
returns

Returns within
10 m

Site fidelity
index

1985

82

57

0.70

1986

–

–

–

1987

112

60

0.54

1988

133

83

0.62

1989

135

101

0.75

1990

247

180

0.73

1991

172

121

0.70

1992

170

110

0.65

1993

204

144

0.71

1994

181

124

0.69

1995

80

52

0.65

1996

85

57

0.67

1997

190

125

0.66

1998

59

44

0.75

1999

194

141

0.73

2000

183

130

0.71

2001

78

54

0.69

2002

211

153

0.73

2003

187

143

0.76

2004

164

128

0.78

2005

196

159

0.81

2006

205

162

0.79

2007

179

140

0.78

2008

262

198

0.76

2009

232

178

0.77

2010

213

173

0.81

Total

4 154

3 017

0.73

Fig. 3. Annual change in site fidelity index.
Marine Ornithology 42: 11–15 (2014)

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Sugawa et al.: Breeding site fidelity of Streaked Shearwater

the birds returned to their exact previous study site. Although a few
switched sites, it should be noted that the distance between Sites
A and B is only 200 m (Fig. 1, Table 1). In addition, the quadrat
analysis of the returns within Site A showed that birds returnedvery close to the release point. Since birds were caught mainlyon the surface, the results do not necessarily indicate return to thesame burrow. However, results are comparable to those of Cory’s
Shearwater Calonectris diomedea, in which site fidelity has been
investigated in greater detail (Thibault 1994, Ristow et al. 1991,
Rabouam et al. 1998).

Headquarters Maizuru District, Japan Maritime Self-Defense
Force, for transportation to the island. We thank Mr. F. Sato and
Mr. S. Komeda of the Yamashina Institute of Ornithology, Abiko,
Japan, for their advice and support on archiving banding data. We
are also grateful to Dr. M. Lawson of Suzuka National College ofTechnology, who improved the English of the manuscript. Helpwith banding on the island by students of universities in Kyoto
was indispensable.

The site fidelity index was high and rather constant throughout
the period. Even after 20 years, the birds returned to the vicinityof the original release point (Table 2). The age of these birds was
not known, but the birds were banded as adults, and had alreadyreached or approached breeding age since they came to the colony.
The longevity record of this species is 36 years (Yamashina Institutefor Ornithology / Ministry of Environment 2012).

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Apparently, Streaked Shearwater maintains a strong tendency tobreed at the same site within the colony for almost all its life. The
site fidelity index varied little from year to year, except for a few
years during the 1980s when it was particularly low. These results
indicate that each bird, or each pair, uses only a small limitedarea in the colony, even if the colony itself is very large. The
implications are that, if a given area within the colony was lost orchanged for some reason, the pair nesting there would have to finda new site for breeding. This could lead to increased conspecificcompetition for a breeding site, depending on the amount of
habitat available.
On a longer time scale, breeding site fidelity increased slowly
during the study period, possibly due to one or more factors. The
colony site examined was flat and well covered by large, mature
evergreen trees. Although this island is sometimes subject totyphoons or heavy rains, it does not collect pooled water; the areawith burrows is rather stable over a long time. Since Streaked
Shearwater is the only burrowing species breeding on this island,no competition with other seabird species exists. A ground
predator, the Norway Rat Rattus norvegicus, was unintentionallyintroduced to this island when army troops were stationed here
before World War II. The rat can possibly attack the nestlings.
However, there seems to be no serious effect on the breeding of
this species.
Site fidelity is also related to various factors other than those of
the colony itself. Breeding failure is known to lower the mateand site fidelity in seabird species (Thibault 1994, MacDonald2009). Breeding success is, on the other hand, affected by foodsupply (Harris & Wanless 1990, Phillips et al. 1996, Furness& Camphuysen 1997, Furness & Tasker 2000). Changes in
the population dynamics may affect fidelity, but we have no
information on that subject. Therefore, the catalyst for the increasein the fidelity index observed in the study period is unknown.
Further work is needed on breeding success, mate fidelity anddemography of Streaked Shearwater in this colony.
ACKNOWLEDGEMENTS
We thank Mr. T. Matsumoto of the Department of Education
and Cultural Asset, Maizuru Municipal Government, for hiskind and continuous support. We also express our thanks to the

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