全 文 ：第 26卷第 3期
2006年 3月
生 态 学 报
ACTA ECOL0GICA SINICA
Vo1．26，No．3
Mar．．2006
红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部IW)L山
地区的迁徙中途停歇生态
王 勇 一，常家传。，弗兰克·摩尔 ，苏立英 ，崔黎明。，杨显峰
(1．Center for Forestry and Ecology，Alabama A&M University，P．0．Box 1208，Normal，Al~ama 35762 USA；
2．北京师范大学生命科学学院，北京 100875；3．东北林业大学野生动物资源学院，哈尔滨 150040；
4．Department of Biological Science，University of Southern Mississippi，Hattiesburg，Mississippi，39406，USA；
5．International Crane Foundation．P．0．Box 447，Baraboo，Wisconsin，53913 USA)
摘要：中国迁徙鸣禽类的保护面对着与世界其他地区如欧洲和北美洲鸟类保护相似的挑战。迁徙鸣禽类具有复杂生活周期和
很大的空间关联。迁徙过程中发生的事件对迁徙鸣禽类种群动态具有决定作用。对于鸣禽类迁徙中途停歇期的生态，比如停
歇期的长短，能量的积 累，生境 的利用等 ，了解还非常有限。在中国东北部的一个鸟类迁徙停歇地对红胁蓝尾鸲(Tarsiger
cyanun~)的中途停歇生态包括迁徙时间、停歇时间、能量状态和性比进行了研究。2002年秋和 2003年春分别捕获了 l751只和
684只红胁蓝尾鸲。红胁蓝尾鸲的体重在秋季迁徙时要比在春季迁徙时重。春季雌性红胁蓝尾鸲停歇时的能量状态指数最
低 ；而秋季的红胁蓝尾鸲比春季的红胁蓝尾鸲停歇时间更长。无论季节和性别 ，红胁蓝尾鸲的能量状态指数和第 1次捕获的
时间早晚成正相关，间接证明红胁蓝尾鸲在停歇期间能够比较快地积累能量。秋季雄性红胁蓝尾鸲13体重净增率最大。估测
秋季停歇期的每13能量净增能维持红胁蓝尾鸲雌性 0．6h和雄性 3．1h的飞行。红胁蓝尾鸲的中途停歇生态与北美和欧洲一些
迁徙鸣禽类很相似。比如，春季迁徙过境的时间和脂肪积累的变化与 自然选择对雄性的要求 ：当食物和气候适宜时尽快到达繁
殖地的假设是一致的。对迁徙中途的停歇生态研究有利于更好地了解鸟类的迁徙行为和更有效地保护迁徙鸣禽类。
关键词：鸟类保护 ；鸟类迁徙；中途停歇生态；中途停歇地；红胁蓝尾鸲 Red—flanked Bush Kobin(Tarslger cyanurus)
文章编号：1000-0933(20o6)o3—0638—09 中豳分类号：Q958，Q959 文献标识码：A
Stopover ecology of Tarsiger cyanurus at M aoershan of northeast China
WANG Yong ，CHANG Jia—Chuan ，MOORE Frank R． ，SU Li—Ying5，CUI Li—Ming3，YANG Xian—Feng3 (1． Center
for Forestry and Ecology，Alabama A&M University，P．0．Box 1208，Normal，Alabama 35762 USA；2．Beijing Normal University，13e#ing 100875，China；
3．Colege of Wildl澹Resources，Northeast Forestry University，Harbin 150040，China；4．Department of Biological Science，University of Southern Mississ f，
Hatiesburg，Missisippi，39406，USA；5．International Crane Foundation，P．0．Box 447，Baraboo，Wisconsin，53913 USA)．ActaEeologiea Sinica，2006，
26(3)：638—646．
Abstract：The conservation of songbird migrants in China faces challenges similar to those in other parts of the world such as
Europe and North America．Songbird migrants have complex life history and are associated with large spatial scale．The events
occurred during migration play a critical role in determining population status．Little is known about stopover behavior of migratory
songbirds in China during passage，including stopover duration，energetic condition(e．g．，the amount of fat stores)，and habitat
use of these birds．We investigated migration timing，stopover duration，change of energetic condition，and sex—related variations
among Red flanked Bush Robin(Tarsiger cyanuru~)at a stopover site in northeast China．A total of I，75 1 and 684 Red-flanked
基金项目：中国国家林业局环志经费资助项目；美国林业部、美国国家 自然科学基金和阿拉巴马农工大学资助项目
收稿日期：2005—08—30；修订日期：2006—01—10
作者简介：王勇(1956一)，上海市人，博士，副教授，主要从事动物生态和鸟类迁徙研究．E．mal：yong．wang@email．∞mu．edu
Foundation item：The pioject wgs supported by the Bird Banding Grant of the National Forestry Bureau of China，and the Forest Service of United State Department
of Agriculture，the National Science Foundation of the United State and the Alabama A&M University
Reedved date：2005·O8·30；Accepted date：2006·01·10
Biography：WANG Yong，Ph．D．，Associate professor，mainly engaged in animal ecology and bird migration．E-mall：yong．wang@ema il
． aRlnu．edu
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3期 王勇 等：红胁蓝尾鸲(Tarsiger cyanunu)在中国东北部帽儿山地区的迁徙中途停歇生态 639
Bush Robins were captured in fall 2002 and in spring 2003，respectively．Body mass of fal birds was higher than that of spring
birds．Con~tion index(i．e．，body mass adjusted for body size)was lowest among spring females．Birds were more likely to stay
longer in fall than in spring．We detected a positive relationship between time of initial capture and condition index regardless of
season or sex，suggesting birds were able to replenish energy stores．The net daily mass gain was the highest among males in fall
(3％ bedy mass)．The net daily mass gain would sustain a flight of 0．6 h for females and 3．1 h for males in fal1．The stopover
biology of Red—flanked Bush Robins is similar to some songbird migrants of Europe and North America．For example，spring
passage time and fat store variation between sexes agree with the hypothesis that males are selected to arrive at their breeding
grounds as early as food resources or climatic conditions are adequate in s ng ．Further research on stopover ecology is urgently
needed in China for a better understanding of the migratory behavior and for the conservation of these songbird migrants．
Key words：bird conservation；bird migration；stopover ecology；stopover site； ‘ r cyanurus
The conservation of migratory birds is challenging because of their complex life history and the spatial scale over which
these migrants trave1．Migratory birds spend about one—third of their lifetime in migration．The events associated with
mi gration play a critical role in determining the population status of migratory songbirds⋯ ．Migration is an energetically
expensive event．Anticipating the energetic demand of migration and uncertainties en route， most songbird migrants
accumulate fat deposits for use as energy source during migration．However，these fat deposits usually are not suficient for
the entire migration joumey，birds have to periodicaly replenish depleted fat stores during migration at the stopover sites
along their migration route．During stopover birds must contend with fluctuations in food availability，the likelihood of
inter-and intra—specifc competition，and limited resources at stopover sites 引．The habitaf quality，behavioral and
physiological responses of the stopover birds，and the relationship between them are the main focus of avian stopover
ecology[。一
．
Songbirds are one of the most abundant avi~una groups in China，an d about one—third of these species are
migratory ．The effort to conserve migratory songbirds in China is facing challenges similar to those in other parts of the
wodd such as Europe and North America．Suitable habitats for many mi gratory bird species are declining on breeding
grounds，wintering grounds，and at stopover sites，and songbird migrants in China probably are affected more severely by
human—induced environmental and habitat changes than their counterparts in the North America．China’s natural resources
are more lim ted．For example，currently。China’s forest land is only about 10％ that of North America’s；and China is
one of the nations that had the highest rate of forest depletion in the world over the past 50 year ．China’s human
population has increased about 2．5 times．and timber production has more than tripled since later 1940’s 引． Forest
exploitation and monoculture in China has led to adverse consequence including degradation of forests and landscapes，loss
of biodiversity，unacceptable levels of soil erosion，and catastrophic flooding ．Over 6 1％ of wildlife species in China
have suffered from habitat loss ．China is among the countries with he most threatened birds and mammals in the
world ．The conservation of migratory songbirds in China is further hindered by the lack of available data．Ornithological
research has been limited in both quantity and quality in China．For example，between 1924 and 1989，China had only
715 research papers dealing with songbirds．10 of them addressed songbird migration and，to our knowledge，there is no
published research on stopover ecology of mi gratory songbirds．
In the early 1980’s．the Chinese government initiated a national bird banding program，and since then several
mi gratory bird protection agreements have been signed between China and other countries including Japan，United States，
and Australia。引
． Whereas the main objective of this banding program is to identify migratory routes。the program provides
an opportunity to collect other vital information such as site fidelity，life span，and habitat use of many bird species
including both migratory and nonmigratory bird species．
In this study，we used mist—netting data collected at one of the banding sites operated by the Chinese Bird Banding
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生 态 学 报 26卷
Ofice in northeast China and examined several stopover parameters of Red—flanked Bush Robin(Tarsiger cyanurus)．
Specifcally，we examined(1)timng of spring and fal migration，(2)stopover duration，(3)change in energetic
condition during stopover，and(4)sex related variations including body mass and rate of mass change．This study is also
intended to provide an indirect assessment of habitat suitab ility of the study site for the Red—flanked Bush Robins and those
songbird mi gratory species with similar habitat requirements during stopover．We hope that this research wil promote the
study of stopover ecology of migratory songbirds in China．
1 M匝THoDS
1．1 Study species Red flan ked Bush Robin is a small migratory passerine species of the Old World Turdidae．Th e
species breeds in boreal and mountain thick mossy conifer forest in Russia，north Asia to the Himalayas and western
China，and is most abundant in taiga of moist soil with undergrowth．It is an insectivore，but also feeds on fruits and seeds
during the nonbreeding seasons，foraging in low trees，shrubs，an d on ground．It nests on the ground in hoHows among
tree roots，in bankholes，or slishtly above the ground in stumps or fallen logs．111e species winters in Southeast Asia
including the area south of the Yangtze River in China，Taiwan，and the Hainan Islands．There are two subspecies in
China：T．c．cyanurus and T．c．rufilatus ．The former is the northern long—distance migratory subspecies breeding
in northern Eurasia；and the latter is the southern subspecies，breeding in the Himalayan region，exhibiting mainly short—
distance altitudinal movements，although some birds reach Assam，Burm a，and northern Th ailand．Th e individuals in this
study belong to the first subspecies．Although the study site is within the breeding region of this subspecies ，individuals
captured are migrating birds because(1)there is no breeding population at the study site or adjacent areas，and the
nearest breeding populations are over 100 km north on the northern slope of Xiaoxinganling Mts(Chang pets．obs．)；and
(2)few individuals were recaptured in subsequent season or years at the study site(Chang pers．obs．)．
1．2 Study site The study was conducted at Laoyeling Banding Station(45。22 N，127。32 E)located within the
Maoershan Experiment Forest of Northeast Forestry University in Heilongjiagg Province of northeast China㈣ ．Northeast
China is one of the few areas in China that still supports extensive natural forests．The Experiment Forest covers an area of
30km×20 km， about 26，507 hm ， and is located in the northwest remnant hills of Zhangguangchai of the Changbai
Mountains．Th e average elevation of the study site is 300 m above sea level with a 10—15。slope．Th e area experiences
cold winters an d warm summers with an annual average precipitation of 723 mm．Natural vegetation is deciduous broadleaf
forest dominated by oak or a mixture of hardwood species．Canopy species are comprised of Japanese white birch(Betula
platyphyla)，aspen (Populus davidiana)，Mongolian oak (Quercus mongolica)，Manchurian ash (Fraxinus
mandshurica)，Manchurian walnut(Juglans mandshurica)，Amur cork tree(PheUodendron amurense)，color maple
(Acer mono)，Tilia amurensis，and elm(Ulmus propinqua)．Stands of Scotch pine(Pinus sylvestris)and Korean pine
(Pinus koraiensis)occur in some places．Common shrub species include wilow bushes(Salix spp．)，Amur maple(Acer
ginnala)，Amur lilac(Syringa amurensis)， bushclover(Lespedeza bicolor)，honeysuckle(Lonicera spp．)，red
elderbery(Sambucus racemosa)，hazel(Corylus heterophyla)，Dahurian buckthorn(Rhamnus dahuricus)，hawthorn
(Crataegus pinnatifda)，and Dahurian rose(Rosa dahurica)．
1-3 Capture and recapture We collected data in fall from 5 August to 27 October 2002 and in spring from 10 March to
22 May 2003 We used 173 mist nets(10 m long，2．5 m hi【gh，5 tiers，30 mm mesh)，which were permanently
established as three clusters within the banding station．Nets were placed opportunistically among forest，shrub，grasslan d，
and edges of agriculture fields and fish ponds to maximize the capture rate．111e maximum distance among the three netting
clusters is about 3 000 m．Mist nets remained open through the banding season．Nets were checked every 1～ 1
．5 h
between 6 am and 8 pm daily，travel among the nets were assisted with bicycles．We took all birds captured to a central
location for data colection and banding．We measured body mas (4-O．5 g)using a spring scale and unflatened wing
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3期 王勇 等：红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽儿山地区的迁徙中途停歇生态 641
chord(±O．5 mm)with a wing ruler．Each bird wag banded with a uniquely numbered metal leg band issued by the
Center of Bird Banding of China and released immediately after measurements were taken．Recaptured birds were re．
weighed．Identification of the species an d sex is based on several resources【 6]
． No attempt was made to identify the age
0f the birds because the information or technique for reliably identifying age(e．g．，skul osifcation)of this species has
not been documented yet．Handling time was usualy less than 30 min．
1．4 Data analysis Netting efort was constant among dates during the study period．We estima ted stopover duration by
subtracting the date of fimt capture from the date of last capture of recaptured birds．This method yielded a conservative
estimate because we assumed that birds arrived on the day of initial banding and departed on the day of last recapture[17]
．
Individuals that were not recaptured were assumed to have departed from the study site on the same day they were banded，
and were assigned a stopover duration of zero．We refer to birds captured after the day of initial capture as“recaptures’
an d all other birds as“non—recaptures．’Rate of body nlass chan ge during stopover of recaptured birds was estimated
Mass change(g／d)= (Ml—Mi)×Stopover duration。 (1)
where Mi is body nlaBS(g)at initial capture，and M is body mas at last capture after adjusted to body mas at 12 pm
using a regression approach between body mass and the time of capture．Folowing Winker’s method『18]
， we calculated a
condition index for every individual：
Condition index=100×mass×wing (2)
where wing is the wing chord (mm)，and mass is the initial body mass．Condition index provides an estimate of body InaBs
adjusted for body size ．We then regressed condition index on the time of initial capture with a linear model to estimate
the daily body mass chan ge．Th e capture time was transformed to decimal unit and time since sunrise before the regression
analysis．The slopes of the regresion models provide an estimation of the hourly condition change( (100 mm·h))．We
used analysis of covariance to test the equality of the slopes of these regression models among sex—season groups．We
estimated the mass change of an average individual during a 24一h stopover by first calculating the gross daily gains of al
individual with average body mass and wing chord and an assumed 8 h each day available for gaining body mass during the
netting period ．Th e average day length was about 13 h during spring and 12 h during fal at the study site．Our estimate of
mass chan ge is a more conservative index than that of Winker’s method ，which calculated average daily gain based Ol
the actual avemge day length at his study site in Veracruz，Mexico．A nocturnal loss of 4．5％ of body mass[18_l9j is then
subtracted from gross gain to obtain the net body mass gain，and the net ITIaSS gain is then divided by body mass at initial
capture to estimate the percentage of mass gain．We estima ted flight cost for each sex group in each season by using a
statistical model developed by Castro and Myers[加_：
Metabolism(kJ／h)：2．23×M 咖 ×W (3)
where M =average body mass in g．and W average wing chord in cm．Th e values for metabolic rate are converted using
the equivalent of 39．5 kJ，g of fat．We assume that water gain or loss during stopover is negligible，and mass change
reflects the variati0n of the amount of fat deposit[18l ，
． Th e likelihood rati0 chi-square test was used to examine if the
recapture rate difered between spring and fall and between males and females．To test the efect of season and sex and
their interaction on body mass，wing chord，and condition index，we used a two—way ANOVA(General Linear Mode1)．
Means and standard deviations(SD)are reported throughout the manuscript， and a statistical signifcance level of o／=
0．05(two．tailed) is used unles indicated otherwise． Al analyses were conducted with SAS V8．2 圳 or SPSS／PC
V10．2I
．
2 RESULTS
We captured a total 1，751
．
and 684 Red-flanked Bush Robins during fall 2002 an d spring 2003，respectively．We
randomly measured body mass and wing chord for a subsample of 769 individuals(390 females，367 males，and 12
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642 生 态 学 报 26卷
unknown)during the fal of 2002 and 572 individuals(1 87 females，383 males，and 2 unknown)during the spring of
2003(Table 1)．The median capture date was Julian day 276(October 4)±9 in fal，and Julian day 106(April 17)±
6 in spring(Fig．1)．We were not able to assess sex and age related variation because gender was not identifed for all
birds，and age identification was not performed．Two high capture periods occurred in fall：the first on 24 and 25
September with about 200 captures each day；the second on 14 October with 358 captures．During spring，daily capture
rate was relatively constant between 10 and 23 April，with the maximum capture of 65 individuals on 22 April．Males
(Julian day 106±6)moved through the site earlier than females(Julian day 108±7)。
Table 1 Sample size。唧IdIH仰 Index。body m ss．and wing chord(mean±SD)of Red．flanked Bush Robin(， l r cyanurus)captured during fal
(2002)and spring(2003)migration in Heilongllang Provin∞of northeast China
a Percent of total captures of the season；b Condition index=mas×wing ×100(Winker 1995)；c Percent of capture for each 8ex group in each season；d
Stopover duration for birds recaptured after the initial capture date(recaptures)；e Mass change per day of recaptures；f F statistics from two-way ANOVA ( by
髓as0n)with unknown individuals excluded from analysis：ns=not signifcant， P(O．05。* P《0．O1， ** P(O．001
The average body mas of captured birds was higher(tl瑚 =4．1，P((13．3±1．1)g)．Two-way ANOVA suggested that body mass had a signifcant interaction between season and sex(Table
1)．This interaction is because body mass of females was lower in spring than that in fal while males had similar body
mltss between the two seasons(Table 1)．Body mass of males was higher than that of females in spring(￡56g=4．7，
P<0．001)；body mass between males and females was not different in fall(Table 1)．Males had longer wing chords
((77。9±2．0)mm)than females((76．3±2．0)mm)；F1
．
1323=168．0，Padjusted the body mas by body size(wing chord)，showed the same patern of body mas variation：a signifcant
interaction between season and sex(Table 1)．Females and males had similar condition upon capture in fall，while the
males had a hilgher condition index than females in spring(Table 1)．
The recapture rate was not different between fal(39 birds，2．2％ of total capture)and spring(17 birds，2．5％ of
total capture)，and the recapture probability was not affected by sex(Likelihood—ratio test)in either seasons．Recaptured
birds tended to stay longer(Fl ：4．0，P<0．05)in fal((5．7±4．3)d)than in spring((3．4±3．4)d)；and males
tended to stay shorter than females in both seasons although it was not statisticaly signifcant(Table 1)．Among
recaptures，females(4．7％ ±12．3％)gained mass while males lost mass(一1．5％ ±5．9％)during their stopover
(Table 1)．
Th e linear regression an alysis suggested that there is a positive relationship between condition index and initial time of
capture for each sex group in both seasons(Fig．2)．The slopes棚 similar among sex-season groups except females in
spring which had a lower rate of mass change(F¨ Ⅲ =3．4，P维普资讯 http://www.cqvip.com
3期 王勇 等：红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽儿山地区的迁徙中途停歇生态 643
O9．1O O9．16 O9．22 O9-28 1O O4 lO-1O 1O．16 1O-22 1O．28 O3-l5 03-21 03-27 04-02 04-08 04-14 04-20 04-26 05-02
09．13 O9．19 O9．25 1O．01 IO．O7 1O-13 1O-19 1O-25 IO-31 O3-l8 03-24 o3-3O O4-∞ 04-I1 04-17 04-23 04．29 05·05
Month-day(2002)
Fig．1 Migration timing of the Red—flanked Bush Robin during舢 (A)and spring(B)al Lanyeling Banding Station of Maoershan Experiment Forest of
Northeast Forestry Univemity in Heilongjiang Province of China
females both in spring and fall(Table 2)．The nocturnal metabolic loss of body mass was about 0．6 g on average based on
the estima tion of4．5％ ofbody mass．It is apparent that on average the birds had a positive net body m聃s gain for a 24h
diel cycle，except females in spring(Table 2)．The net gain was the highest among the fal males．Assuming the net gain
was the fat deposit．the net daily Inass gain of average individuals could sustain a flight of 0．6 h for females and 3．1h for
males in fal and a flight of 0．5 h for males in spring(Table 2)．Average females in spring did not gain enough fat for
migration at this site based our calculation．
3 DISCUSSION
Red．flanked Bush Robins in this study"winter mostly south of the Yangtze River and breed in northeastern China．Our
study site is within the species’reported breeding range although no local breeding population was detected at the site
(Chang pers．obs．)．Many birds captured in this study have almost arived at their breeding grounds in spring and just
initiated their migration journey to their wintering grounds in fal1．Two high capture periods were detected in fal1．Such
multiple high capture periods are also common among songbird migrants in North Americafl71
． We suspect that the two
peaks of capture in spring and fall may reflect populations that have diferent breeding and／or wintering distributions．
Although we did not identify gender of all individual captured，we observed that in general males tended to move through
our site eadier than females both in spring and fal(Chang pers．obs+)．This patern is consistent with some species in
Europe and North Americal261
．
Table 2 Estimated dally body m日ss clmnges using the regression models In Figure 3 for average individuals in each sex group and~．ason of Red-flanked
Bush Robin(Tarsiger cyanurus)captured during fal(2002)and spring(2003)migration In HelongJlang Province of northeast China
a，Mass g8in during one day stopover for individuals of the average wing length and an 8 h day estimated from regression models in Figure 2；b，Body fat used
in nocturnal metabolism=4．5％ of晌 n bodymass after one day stopover(mean arrival bodymass+diurnalmass change)；c，Net dailymass gainin 24一hperiod
曲er suht咖ting nocturnal metab c loss；d。Net daily mass gain as a percentage of arrival body mass；e，Night COSt estimated based on M=2．23×Mas’。柏7×
Wing一 ×39．5一 ；f，Potentialflight durationfor nocturnalmigration gained from one day stopover assuming net gainisallfat
The gross daily mass gain of Red—flanked Bush Robin about 1 g／day was similar to that of Bluethroat(Luscinia s·
枷 瑚 姗 挪 抛 m m 如 0
旨量坤矗 晴I|nP 眚 暑．尹0 d马nN
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644 生 态 学 报 26卷
svee／~a)in Europe ．We expected that the body lnass would be higher in fall than in spring because our study site is
near the breeding grounds，and the birds captured at our site just started migration in fal while they wlere a1most 0n their
breeding grounds in spring． Interestingly，we found seasonal body IIlass variation was mainly due t0 l0wer b0dv rnass 0f
females in spring，while females in fal an d males in both season had simi lar body mass
． Wilson’s Warblers f 0
pusila)in New Mexico of USA showed simlar paterns between seasons and between sexes[29]． The higher fat stores and
faster rate of mass gain of males in spring may indicate that the selective pressures of stopover are diflferent fbm that 0f
females．Like many songbird species，male Red—flan ked Bush Robins usually arrive at breeding grounds earlier than
females(Chang pens．0bs．)．These paterns are consistent with the hypothesis that males are selected t0 arrive at their
breeding grounds as early as food resources or clima tic conditions a】 adequate，whereas fle脚ale吕 ve later，closer t0 the
time when they Can successfully begin nesting ．Elegren[28]found that the stopover patterns including rate 0f fat
accumulation were simi lar between sexes in Bluethroats during fall mi gration in Europe
．
Springfcanal~ Springfcnnalc
6 8 10 12 14 16 18 20
Time
Fig·2 Seasonal and sex rdated daily body c。ndition change based on linear regression8 betweon conditi。n index and capture time
The condition index was estimated as body ITla8s×wing chord一 x 100[ ]
The estimates of body mass gain and rate of mass gain based on recaptured birds were l0wer than the estimates 0f th
e
same variables based on the regression analysis between condition index and time of initial capture of all the birds
．
Capturing birds mi ght interrupt birds’normal behavior，cause stress，and lead to body mas8 decline．which was 0bserved
in other studies ．Recaptured male birds lost body mass while female gained maBs while the pattem was reversed based
。n the regression analyses of all birds captured
． Th ese diferences are probably because recaptured birds may reflect a
biased sample of all the birds that used 0ur stopover site
．
Carlisle et a1．[ 3。found general agreement betwleen regressi0n
data and l鹅s change among recapture birds．Average females in spring did not gain enough fat for migration 0n dailv base
at th te based on our estima tion．However no local breeding population Was detected
． Th is suggests that female8 might
h ve to tay longer to gain energy stores for further migration；our recapture data did suggest that females tended t0 have
l0nge t0pover duration both in spring and fal1．Th e other possibility is that females already had SUmcient ene y st0res f0
r
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" ：宝：2 H ¨
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3期 王勇 等：红胁蓝尾鸲(Tarsiger cyanurus)在中国东北部帽&1．LI地区的迁徙中途停歇生态 645
migrating to their breeding grounds during sprint stopover，our capture data did support this possibility：females had higher
bodyⅡlass than that of males in spring．Balrlein found that many tran s-Saharan songbird mi grants used an intermitent
migratory strategy，with regular stopovers during the day and flight at night．The female Red-flanked Bush Robins in spring
might stop at our stopover site for resting and continued migration at night．
Migrants require suitable stopover sites for resting and replenishing depleted fat stores㈨
． e suitability of stopover
habitats in relation to migran ts’fitness is dificult to measure directly；indirect measurements include the fat load at
departure and stopover duration of stopover birds【笛-
．
e optimal time mi nimization model of migration【 ’]predicts that
migran ts should deposit fat stores faster and depa~ sooner at suitable stopover habitat than at a lower quality site．Th e
quick mass gain，short stopover duration，and low recapture rate of stopover Red-Flanked Bush Robins might suggest that
the habitat at our study sites is suitab le for this species．To our knowledge，this is the first study of stopover ecology of
songbird migrants in China．Because financial and logistic limitations，we did not coHect similar data for other species or
from other locations for the same species。which excludes the possibility for comparisons to our results．Collect and analysis
of data for other migrant species，and for the same species in muhiple years should give us a beter understanding of the
spatial an d temporal dynamics of mass gain during migration of these birds and their associated habitat【蚓
．
Northeast China is one of the last remaining areas that preserves natural habitat including grasslands and forests in
China． However， these habitats are under threat because of demands for economic development such as urban
encroachment，agricultural expansion，an d timber production．In 1998，the Chinese government established the Natural
Forest Conservation Program(NFCP)．One of the main objectives of this program is to protect existing natural forests．
Most forest lands in the Northeast China are listed as the first priority for protection and conservation by this program．Th e
fact that Red—flanked Bush Robins are able to gain body mass，and to do so quickly，at our study site is encouraging．We
know little about mi gration routes，paterns of habitat use，and stopover behavior of songbird migrants in China．Avian
ecologists such Baidein【 ]caled for migration research aiming at much more comparative research and a more integrative
approach at various spatial and temporal scales and using currently advance research techniques．To determine patterns
observed in Red—flanked Bush Robin hold up among other migratory songbird species at our study site，and at other
stopover habitats along migration routes，further study of stopover behavior is urgently needed for understanding the
evolution of migration systems in songbirds，and for the conservation of these species in China．
ACKNOWLEDGEMENTS：We are very grateful to Hong Enrong，Wu Wei，Tang Jingwen，and Zhu Kunjie for assisting field
data colection．Th e research would be impossible without the support from the Northe：ast Forestry University， Maoershan
Experiment Forest，and Bird Banding Center of China．We thank Zachary Felix，Dr．Frances C．James，and two anonymous
reviewers for their ideas and constructive comments on earlier drafts of the manuscript．
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