Studies

The Auk 112(4):964-972, 1995
INFLUENCE
DECISION
OF BODY
AND
CONDITION
REPRODUCTIVE
BLUE
ON
REPRODUCTIVE
SUCCESS
IN THE
PETREL
OLIVIER CHASTEL,HENRI WEIMERSKIRCH,AND PIERREJOUVENTIN
Centred'EtudesBiologiques
de Chizg,CentreNationalde la Recherche
Scientifique,
79360 Beauvoir sur Niort, France
ABSTRACT.--Following
the prediction that body condition affectswhether eggsare laid in
birds, we examinedvariationsin prebreedingbody condition (i.e. body massone month
beforelaying,adjustedfor structuralsize,breedingexperience,and dateof return),incidence
of breeding,and reproductivesuccess
of male and femaleBluePetrels(Halobaena
caerulea).
Condition was inversely correlatedwith date of return and positively related to breeding
experiencein males,but not in females.In both sexes,condition displayedyear-to-year
variations; the 1991-1992 seasonwas distinguishedby poor condition, low incidence of
breeding,and low reproductivesuccess.
Conditionin both malesand females,and breeding
experiencein males had significanteffectson the "decisionto breed." Breedingexperience
also influenced the decisionto breed, especiallyin males. Condition, but not breeding experience,had a significantinfluenceon male incubationsuccess.
Femalebreeding performanceswere not related to condition but to breeding experience.Annual averagebreeding
success
was highly variable and significantlycorrelatedwith male condition.Prebreeding
visitsappearedto be criticalfor this pelagicseabirdand mayallow individuals,throughtheir
condition,to managetheir reproductiveinvestment.We suggestthat conditionmainly affects
egg formationin femalesand, togetherwith breedingexperience,influencesthe duration
and the quality of prebreeding colony attendancein males. The potential relationshipsof
body condition,breedingexperience,reproductiveinvestment,and future survivalare discussed.Received11 November1993, accepted25 February1994.
THE INFLUENCE
of body massor condition on
fitnesscomponents,suchas fecundity and sur-
trel populationstypically include high proportions of nonbreeders (for review, see Warham
1990).Nonbreeding yearsare of regular occurspecies.Studieshave demonstratedthat heavier rence among establishedbreeders(Mougin et
femalesproducelarger clutchesand that heavi- al. 1984, Woollet et al. 1989). In the order Proer males achieve greater reproductive success cellariiformes, intermittent breeding has been
and higher survival rates(Aldrich and Ravel- linked to ageor experience(Weimerskirch1990,
ling 1983,Newton et al. 1983,Petrie 1983,Gib- 1992), lossor changeof mate (Fisher 1976, O1bons 1989). Body condition also may influence lason and Dunnet 1988), food availability, enfood provisioning to the chick (Weimerskirch vironmental changes, or temporary inaccessiet al. 1993, Chaurand and Weimerskirch 1994a), bility of breedingsites(Prince 1985,Ainley and
pair-bond status(Robb et al. 1992),and second- Boekelheide 1990, Chastel et al. 1993). Only a
broodinitiation (De Laetand Dhont 1989).Long- few studies have examined the influence of interm studies have pointed out slight but sig- dividual traits, such as body condition, on renificant contributions of individual body mass productive decisionsand recruitment in longto lifetime reproductivesuccess(Gelbach1989, lived seabirds(Fisher 1967, Weimerskirch 1992).
Mills 1989).Additionally, broodmanipulations In fact, the influence of body condition on rehave establishedrelationshipsbetween female productive successhas not even been clearly
mass loss and experimentally enlarged brood establishedin pelagic seabirds.This is in consize (Nut 1984). However, body masshas been trast to the well documented studies of relashown to have no influence on fecundity or tionships between food supply, adult condisurvival in some species(Naylor and Bendell tion, and reproductive effort that have been
1989,Marjakangasand Aspegren 1991,Robbet conducted on inshore feeders such as terns
al. 1992, Meathrel et al. 1993).
(Monaghan et al. 1989). However, general imMostseabirdsare long-livedspeciesand have provement in breeding performancewith age
low fecundity (Lack 1968). Albatrossand pe- or experience has been well established(see
vival,
has been established for several avian
964
October1995]
BodyCondition
ofBluePetrels
965
review in Newton 1989). Pelagic seabirds(e.g.
albatrossesand petrels) are excellent subjects
for studying the simultaneouseffectsof body
condition and breeding experience on reproductive investment and breeding performance
becausethesebirds are more likely not to jeop-
weighed,and the presenceor absenceof an eggwas
recorded.Femaleswere weighedat the beginningof
the secondincubationshift in mid-November(range
12-15 November).Hatchingsuccess
was determined
by inspectiona few daysafter hatchingwas to occur
(range28-30 December).Chickswere bandedin late
January.Annual breeding successwas calculatedas
ardize their future survival (Goodman 1974, the proportionof eggsresultingin fledgedyoung.
Drent and Daan 1980).
Two reproductivetraits (breeding decisionsand
Blue Petrels (Halobaena caeruelea) are small
breeding success)were studied and defined as folseabirds(200 g) that forage over subantarctic lows.Breedingdecisionincluded:(1) not seenduring
pelagic waters (Prince 1980, Steeleand Klages current breeding season,but seen subsequently;(2)
1986). These colonial burrowing petrels visit occupyinga burrow,but not breeding;and (3) breedincluded:(1) failure to hatchan
their breeding groundsmore than a month be- ing. Breedingsuccess
rearinga chickto fledging.
fore a period of highly synchronouslaying egg;and (2) successfully
Variablesassociatedwith thesereproductivetraits
(Paulian 1953, Jouventin et al. 1985, Fugler et
were defined as follows. (1) Prebreedingbody conal. 1987). This prebreeding period provides a dition (hereafter, "condition") was representedby
good opportunity to determine factorsinflu- body massscaledby linear, structuralmeasurements.
encingfuture reproductiveinvestmentand particularlyto testthe predictionof Drent and Daan
(1980)that the decisionto breedor not is clearly
relatedto femalebody condition,or that of both
sexes,where the two membersof the pair share
breeding duties equally (Lack 1968).
In this paper we examine the variation in
body condition (body masscorrectedfor linear
dimensions,breeding experience,and date of
This resulted in a condition index (Johnson et al.
1985).We combinedmeanbodymass,culmenlength,
wing length, and tarsuslength in a principal componentsanalysisfor 73 individuals weighed on at
leastthree occasions
(up to 13) during prebreeding
visits, incubationroutine, and postbreedingvisits.
Componentsscoresindicated that mean body mass
explainedmost of the variation. Loadingson principal componentI were: mean body mass,0.66; culmenlength,0.51;wing length,0.49;andtarsuslength,
return) of adult Blue Petrels to determine the
0.26.Additionally,culmenlengthwaspositivelycorextentto which prebreedingbodyconditionin- relatedwith mean body mass(r = 0.24, n = 73, P <
fluencesthe reproductivedecisionsand repro- 0.05). Condition,therefore,was estimatedusing the
ductive success of both males and females.
following ratio: prebreedingbody mass(g) divided
by culmenlength (ram).A high value for this index
signifiesgoodbody condition.The date of return was
METHODS
the first record in the study colony (1 September=
The study was conductedon Mayes Island in the day 1, 6 October= day 36). Breedingexperiencewas
easternpart of the Kerguelen Archipelago (48ø38'S, thenumberof previousyearsduringwhichbreeding
68ø38'E).The island is free of alien predators.Every was attempted. In females, the degreesof freedom
year from 1986 to 1991, 150 burrows of Blue Petrels for breeding experiencewere smaller than for males
were monitored. Study burrows were fitted with a due to a smallersamplesize.In the 1988-1989season,
closeable observation window above the nest chamno data were availablefor the prebreedingperiod,
ber and individualswere bandedusingmonelbands. althoughburrowswerecheckedduringthebreeding
During the prebreeding period (8 September-6 Oc- period. Thus, breeding experienceof the birds was
tober),petrelsoccupiedburrowsby day and, except known. For several other calculations, birds were simasinexperienced(notrecordedbreeding
for the 1988-1989season,diurnal inspectionsof the ply classified
studyburrowswere conductedeverytwo daysto de- before)and experienced(at leastone previousbreedtermine datesof return and to measureprebreeding ing attempt). Year refers to breeding seasons1986bodymassfor eachindividual.Birdsweresexedusing 1987, 1987-1988, 1989-1990, 1990-1991, and 1991playbackrecords(Bretagnolle1990)and weighed to 1992. Return rate was the proportion of birds seen in
the nearest2 g with a 300-gPesolascale.The follow- the prebreedingperiod and subsequentlyrecovered
ing measurementswere taken: culmen length and the following year.
Relationships of condition and date of return,
tarsuslength to nearest0.1 mm; and flattenedwing
chord to nearest 1 mm. Burrows were checked for
breeding experience,previousbreeding status,and
eggseachyearin early November(range30 October- year were examinedusing one-way analysisof vari3 November);laying of large,singleeggswashighly ance (ANOVA). Comparisonsbetween means were
synchronous(Jouventinet al. 1985) and, at this time, made using two-tailed t-tests.To evaluatethe influtraits
98% of the birds are malesstarting their first incu- ence of these variables and noncontinuous
we used
bation shift (T. Chaurand unpubl. data). Males were (breedingdecisionand breedingsuccess),
966
CHASTEL,
WEIMERSKIRCH,
ANDJOUVENTIN
TABLE
1. Annual variation in mean (+SD) prebreeding body condition and date of return of Blue Petrels (malesand femalespooled),1986-1991.
Breeding
season
n
1986-1987
1987-1988
1989-1990
1990-1991
1991-1992
13
16
34
38
56
Prebreeding Date of return
body condition (September)
8.35
7.69
7.95
7.73
7.11
+
+
+
+
+
0.72
0.70
1.06
0.99
0.53
12.69
13.62
13.14
13.50
22.17
+
+
+
+
+
2.21
4.14
4.32
6.24
6.37
[Auk,Vol. 112
0 1991 - 1992
• ß
ß1986-198
to
•'- 10
1990- 1991
.
.
o
o
e. 6-
8 1'0 1• 1•41• 1• 20 22 24 26 28 30 32 34 •6
nonparametricKruskal-Wallis tests. Statisticalanalyseswere performedusing the STATITCF and LOGITHEQ packages(ITCF 1988).
Date of return (days after beginning of September)
Fig. 2. ?rebreedingbody conditionof individual
male Blue ?errels in relation to date of return (r =
-0.517,, = 50, P < 0.001), 1986-1991.
RESULTS
Prebreeding
period.--During the prebreeding pooled,F4,• = 9.36,P < 0.001)and wasparticperiod, 75.1%of the birds occupyinga burrow ularly poor in 1991-1992.
by day were males(n = 173). The averageduInexperiencedmaleshad a poorercondition
ration of a visit was 7.83 _ SD of 0.83 days (n index than did experiencedbirds 7.22 _ 0.71,
= 12), during which birds lost 4.45 g per day n = 23, and 7.84 _ 1.01, n = 46, respectively;t
(n = 7). Prebreedingbody massranged from = 2.63, P = 0.01), but the difference was not
165 to 234 g (CV = 11%)in males,and 168 to significantin females(7.30 + 0.65,n = 11, and
234 g (CV = 9%) in females.No female was 7.68 + 0.74, n = 17, respectively;t = 1.39, P =
found twice during diurnal surveys,indicating 0.17). In males,but not females,conditionsigthat they spent lesstime ashore.Condition did nificantly improved with breeding experience
not differ between males and females (FLu5=
(males, F3,49= 5.69, P < 0.01; females, Fz•0 =
0.76,P > 0.05).Conditiondifferedsignificantly 0.96,P > 0.05;year 1991-1992excluded;Fig. 1).
between years (Table 1; males and females
There was no effect of sex (Fu•s = 0.23, P >
0.05) or breedingexperience(males,F•,so
= 0.29,
P > 0.05; females, F4a• = 2.28, P > 0.05; year
1991-1992 excluded) on date of return. However, the mean date of return differed notably
11
5
amongyears(Table1;malesandfemalespooled,
F,,•5•= 23.4,P < 0.001)due to 1991-1992,when
the mean date of return was delayed by nearly
10 days.Condition and date of return were in-
8.5
4 4
8.0
11
0
1991-1992
ß
1986-1987
14
to
1990-1991
•= lO
es
7.5
T
¸ Females
16
7.0
0
*So
I
Breeding
*
o
8
3
experience
Fig. 1. Relationshipbetweenbreedingexperience
and prebreedingbodyconditionin maleand female
Blue Petrels.Samplessizes given adjacentto each
point and datapresentedas œ+ SD. Significantdifference in malesonly (P < 0.01).
ß
o
8 8
Oate of ret-;n (days after beginning of September)
Fig. 3. ?rebreedingbody condition of individual
female Blue Petrels in relation to date of return (r =
-0.062, n = 22, P > 0.05), 1986-1991.
October1995]
TABLE 2.
BodyCondition
ofBluePetrels
Results of Kruskal-Wallis
tests for overall
Sinceconditionand experienceinfluencedthe
decision to breed, especially in males, we consideredthese two variablessimultaneously(Ta-
effectsof severalvariableson breedingdecision(82
males and 34 females) and breeding success(51
males and 23 females) in Blue Petrels, 1986-1991.
Males
Source of variation
df
ble 3). Experiencedmalesthat did not breed in
the current seasonhad poorer condition than
birds that bred. For inexperienced males, the
same trend was not significant. Nonbreeding
malesshowedno significantdifferenceswheth-
Females
H
967
df
H
10.20'*
7.62*
2
2
9.53**
0.44
Breedingexperience 4
15.40'*
2
7.75*
0.61, n = 17; t = 1.53, P > 0.05). In females,
Year
15.54'*
4
0.24
experiencedbirds that bred appearedto be in
better condition than nonbreeding ones, but
the difference was not statistically significant
2.57
0.57
(Table 3).
Breeding decision
Prebreeding body
condition
Date of return
2
2
4
er or not they were experienced(experienced,
7.46 + 0.66, n = 18; not experienced, 7.13 +
Breedingsuccess
Prebreeding body
condition
Date of return
2
2
8.92**
7.02*
2
2
Breedingexperience 3
4.27
2
15.78'*
4
Year
4
The decisionto breed alsowas influencedby
year in males,but not in females(Table 2). The
strong effect of year on breeding decision in
malesis due mainly to the poor 1991-1992 season, during which nearly 70% of the males
present did not breed. If the 1991-1992 data are
removed,breedingbirdsshowedconditionand
were more experiencedthan nonbreedingones
13.54'**
3.04
0.05; **, P <• 0.01; ***, P '• 0.00J..
versely correlatedin males (r = -0.517, n = 50,
P < 0.001;Fig. 2), but not in females(r = -0.062,
n = 22, P > 0.05; Fig. 3).
Breedingdecision.--For both males and females,the "decisionto breed" was significantly
influencedby condition (Table 2). Condition in
the two categoriesof nonbreedingmales(birds
not presentand birds presentbut not breeding)
(Table 4). However, date of return was similar,
indicating that condition and experiencehad
the mostimportantinfluenceon malebreeding
decision during most years.
Returnratein relationtobreeding
decision.--Birds
was similar (t = 1.06, df = 37, P > 0.05) and, that occupieda burrow during prebreeding,but
therefore,datawere pooled.Nonbreeding males did not breed, were mainly the inexperienced
averaged lower condition (7.34 + 0.68, n = 37) (malesand femalespooled,57.1%,n = 91). Conthan breeding birds (8.04 + 1.05, n = 45; t = sideringexperienced
birds,nonbreeders
showed
3.49, P < 0.001). Similar results were found in
a similar return rate to breeding birds (Table 5;
females;breeding birds were in better condi- X 2 = 2.29, n = 113,P = 0.15). Nonbreeding intion (7.90 + 0.75, n = 18) than those that did experiencedbirds had a significantlylower rate
of return (Table 5; X 2 = 12.98, n = 91, P < 0.001)
not breed (7.19 + 0.44, n = 16; t = 3.31, P <
0.01). Breeding decision also was related sig- than nonbreeding experienced birds. Many
nificantly to date of return in males(breeding, birds that were first seen in the prebreeding
14.22 + 5.89 days, n = 45; nonbreeding, 19.27 period,but did not breed that season,were nev+ 7.95, n = 37; t = 3.30, P < 0.01), but not in
er seenagain; however, all inexperienced birds
females(P > 0.05).More experiencedmaleswere successfulyrecruited into the breeding popumore likely to breed;experiencein femaleshad lation were seen in later years.
less effect than did condition (Table 2).
Breedingsuccess.--Inmales, condition, year,
TABLE
3. Comparisonsof means(+SD, with n in parentheses)of prebreedingbody conditionof male and
female Blue Petrelsin relation to breeding decisionand previousexperience,1986-1991.
Experienced
Inexperienced
Sex
Breeding
Nonbreeding
t
Breeding
Nonbreeding
t
Males
Females
8.08 + 1.10 (29)
7.87 + 0.72 (13)
7.46 + 0.66 (18)
7.08 + 0.45 (4)
2.17'
2.05a
7.50 + 0.96 (9)
7.68 + 1.45 (2)
7.13 + 0.61 (17)
7.21 + 0.47 (9)
1.12
0.917
*, P < 0.05.
a p = 0.06.
968
CHASTœL,
WEIMERSKIRCH,
ANDJOUVENTIN
[Auk,Vol. 112
TABLœ
4. Mean (_+SD,with n in parentheses
in boxhead)valuesof prebreedingbody condition,date of
return (daysof September)and breedingexperience(yearsof previousbreeding)relativeto breeding
decisionand breeding successin male Blue Petrels, 1986-1990.
Variable
Breeding
(38)
Nonbreeding
(13)
t
Prebreeding body condition
8.19 _+1.03
7.54 _+0.91
2.02*
Date of return
13.02 _+ 5.14
Breedingexperience
2.29 _+0.95
13.38 + 4.11
0.23
1.54 _+0.77
2.56*
Successful Unsuccessful
(20)
(15)
8.50 _+0.89
12.25 _+ 3.97
2.54 _+0.94
7.67 _+ 1.09
14.06 + 6.83
t
2.48*
0.98
2.20 _+1.01 0.75
*, P < 0.05.
and to a lesserextent date of return had a significant effect on breeding success(Table 2),
whereasbreedingexperiencedid not. Malesthat
successfullyreared a chick were in better condition (8.36 +__
0.90, n = 23) than males that were
unsuccessfulat the egg stage(7.59 + 1.03, n =
17; t = 2.43, P = 0.02). In females, breeding
successwas only significantly influenced by
breeding experience(Table 2). In our set of data
(n = 23), femalesrecordedbreeding for the first
time were morelikely to fail (10 of 11 breeding
attempts) than females recorded breeding for
the secondtime (2 of 7 breeding attempts;Z =
wasrelatedto the poor 1991-1992seasonduring
which 72% of eggs (n = 71) failed to hatch.
Becauseegg mortality occurredmainly during
the first male incubationshift, many females
were not identified (seeMethods), causingthe
observeddiscrepancyin year effect. With the
1991-1992dataon males(Table4) removed,only
conditiondiffered significantlybetween failed
and successful
males.
A significant relationship existed between
mean condition of males and breeding success
of the study colony (Fig. 4), while in females
the relationshipwas not significant(r = 0.773,
3.09, P < 0.01), and for the third or more times
n = 5, P > 0.05). During 1987-1988 and partic(2 of 5 breeding attempts, Z = 2.51, P < 0.05). ularly 1991-1992,overall breeding success
was
Since condition influenced male reproduc- low (28 and 25%;respectively)and correspondtive performance, we examined relationships ed to the period of lowest male prebreeding
betweenprebreedingbody massand body mass body condition. However, during 1989-1990,
at the onset of the first incubation shift (egg breeding success
(62%)and male condition were
loss during this period accountedfor 47% of particularly high.
total breeding failure; n = 125). These two categorieswere significantlycorrelated(r = 0.318,
DISCUSSION
n = 47, P = 0.03) in males, but not in females
(r = 0.098, n = 18, P > 0.05). Body condition of
males at the start of their
first incubation
shift
was better for birds that successfullyhatched
eggs than for those that failed; in females, no
differences were found (Table 6). Male condition at the start of incubation
was not different
for experienced and inexperienced birds (experienced, 7.64 + 0.65, n = 56; inexperienced,
During the course of this study, body condition varied significantlyamong Blue Petrels.
Differenceswere related to breeding experience, date of return, year, reproductive decision, and breeding performance.
Individualvariationin bodycondition.--Adults
exhibited great individual variation in body
conditionduring their prebreedingvisitsto their
7.79 + 0.77, n = 21; t = 0.79, P > 0.05). The
strongeffectof year on breeding success
in males
TABLE6. Mean (+SD, with n in parentheses)hatch-
TABLE
5. Return rate (with n in parentheses)during
year y + I for Blue Petrels (males and females
pooled)in relation to breedingdecisionand breeding experienceduring year y, 1986-1991.
Experienced
Inexperienced
Breeding
Nonbreeding
0.93 (74)
1.00 (10)
0.82 (39)
0.42 (52)
ing successin relation to body condition on the
first day of first incubationshift in male and female
Blue Petrels, 1986-1991.
Hatching
Males
Successful
7.88 + 0.72 (43)
Unsuccessful 7.55 + 0.63 (43)
t-value
*, P < 0.05.
2.30'
Females
7.64 + 0.78 (46)
7.66 + 0.78 (16)
0.09
October1995]
BodyCondition
ofBluePetrels
colony. Males in good condition tended to arrive early. This may reflect individual differencesin foragingsuccess.
Heavy malesacquired
reservesto visit early and stay longer. In our
study,femaleconditionwasnot relatedto date
of arrivalprobablybecausethey visitthe colony
during a morecontractedperiodthan their mate.
However, the samplesize for femaleswassmall.
After removingthe 1991-1992datafor males,
our resultsshowed a significant improvement
in condition with experience. Similar effects
havebeenestablishedin Manx Shearwaters(Puffinuspuffinus;Brooke 1978) and Wandering Albatrosses(Diomedeaexulans;Weimerskirch 1992).
The observedimprovementcouldbe due to improvedforagingefficiency,asestablishedin the
Herring Gull (Larusargentatus;
Greiget al. 1983),
thus supporting Lack's hypothesis(1968) that,
in seabirds,the lower skill of young birds is
responsiblefor the delayedonsetof breeding.
Among experiencedbreeders,male BluePetrels
continued to improve their condition with each
breeding attempt. In the very long-lived Wandering Albatross, body conditions increase
throughoutthe breedingcareerof males,while
in femalesit only improves during the first 20
years,perhapsdue to a progressiveselectionof
high-qualitybirdsat older age,rather than from
an improvement in individual skill (Weimer-
skirch 1992).Contrary to what was found for
male petrels, female condition was not related
to breeding experience.In September, females
visited the colony only briefly with the result
that insufficient sample sizes were obtained.
Annualvariationin bodycondition.--Condition
of adult BluePetrelsvaried annually. The 19911992 breeding season(and to a lesserextent
1987-1988)was characterizedby the lowest average prebreeding condition recorded during
the study.Suchyear-to-yearvariation in body
condition hasnot been describedpreviouslyin
pelagicseabirds.In coastalseabirds,for example
Arctic Terns (Sterna paradisea)and Common
Terns (S. hirundo),poor body condition under
conditionsof food shortagehas been detected
(Monaghan et al. 1989, Monaghan et al. 1992,
Uttley 1992).The low conditionrecordedin our
studyin 1987-1988and particularly in the 19911992 could be explainedby a decreasein food
availability.
Bodyconditionand reproductive
decision.--Our
resultsshowedthat bodyconditionof maleand
female
Blue Petrels influenced
whether
an in-
dividual bred, consistentwith the prediction of
969
70
1989-1990
ß
60
1986-1987
50
1990-1991
ß
ß
40 •
• 30
1987-1988
1991-1992
ß
ß
20
Prebreeding
body condition
Relationshipbetween mean prebreeding
body conditionin maleBlue Petrelsand meanannual
breedingsuccess
of studycolony(r = 0.903,n = 5, P
Fig. 4.
< 0.05).
Drent and Daan (1980). It is possiblethat only
birds attaining a threshold condition decide to
breed,assuggestedby Weimerskirch(1992)for
femaleWanderingAlbatrosses.
BluePetrelpopulations are characterizedby a significantproportion of experiencednonbreeders,as is the
casefor other speciesof Procellariiformer(Cory's Shearwater,Calonectris
diomedea
[Mougin et
al. 1984];Short-tailed Shearwater,Puffinustenuirostris[Wooller et al. 1989]; Manx Shearwater
[Brooke 1990]). In our study, nonbreeding but
experiencedbirds showeda rate of return similar to their breeding counterparts. Former
breedersthat skip a seasoncould have experiencedlow foragingsuccess
beforethe breeding
seasonand may not have acquired sufficient
reservesto invest in reproduction.As for the
inexperiencednonbreeders,many were never
recordedlater in the study colony. These presumablyareyoungerbirdsthat may either have
sufferedhigher mortality(asobservedin Shorttailed Shearwaters;Bradley et al. 1989) or have
moved to another part of the colony.
Extensiveoccurrenceof nonbreedingduring
a particular year by establishedbreedershas
been related to food shortageand to large-scale
environmental perturbations (Coulson 1984,
Prince 1985, Ainley and Boekelheide 1990,
Chastel et al. 1993). During the 1991-1992
breedingseason,up to 70%of malesthat visited
the colonyrefrainedfrom breedingand showed
poor condition. Breeding Blue Petrels experience severe declinesin body massduring incubationand chickrearing (Chaurandand Wei-
970
CHASTEL,
WEIMERSICIRCH,
ANDJOUVENTIN
[Auk, Vol. 112
merskirch 1994), as is the case in many other depend on the female'scondition because,even
procellariids (Weimerskirch 1990); under ad- when in goodcondition, a male will not breed
verse conditions, life-history theory (Stearns if his mate fails to lay an egg. Moreover, in
1976) suggestsa trade-off between future reproductive potential or survival. A threshold
value of prebreedingbody condition may allow
individual Blue Petrels to "predict" the likely
outcomeof a reproductive decision or to aban-
procellariiforms,
materetentionis high and pair
formation is a long processthat is known to
affectbreeding frequency(Fisher 1967, Oilasson and Dunnet 1988).
Bodyconditionandreproductive
success.--Inour
don a breedingattemptif the "perceived"risks study, condition did not play the samerole in
to their survival are too great (Drent and Daan both sexes. In males, condition and not breed1980, Reznick 1985, Pugesek 1987).
Experiencealso played a significant role in
the decisionto breed,especiallyfor males.Low-
er breeding incidenceamongyoung individu-
ing experienceclearly influenced breeding success and particularly hatching success.Although an improvment in breeding performance with age or experience has been well
als is characteristic of all seabirds studied, in-
established (for review, see Newton 1989), no
cluding Black-leggedKittiwakes (Rissatridac- study on pelagic seabirdshas related breeding
to body condition.IncubatingBluePettyla; Wooller and Coulson1977),Short-tailed success
Shearwaters(Woollet et aL 1989), Ad•lie Pen- rels have a threshold massat which they sponguins(Pygoscelis
adeliae;Ainley et al. 1983),Ant- taneouslydesert the egg (Chaurand and Weiarctic Fulmars (Fulmarus
glacialoi•ies;
Weimer- merskirch 1994), and males with lower body
skirch 1990), South Polar Skuas (Catharactamac- conditionwould attain this value more rapidly
cormicki;
Ainley et al. 1990),LaysanAlbatrosses and particularlybeforethe return of their mate.
(Diomedeaimmutabilis;Fisher 1976), and Wan- Consistentwith this idea, nearly one-half of the
dering Albatrosses(Weimerskirch 1992). This total hatching failure occurredduring the first
generaltrend for lessexperiencedbirds to post- incubation shift and concerned males in poor
pone their breeding to the next year could be condition. The importance of male body mass
explained in terms of a higher cost of repro- on incubation success has also been demonduction(Williams 1966),especiallyif they show stratedin the Moorhen (Gallinulachloropus;
Giba lower body condition than more experienced bons 1989).
breeders.
In contrastto males,breeding experienceand
In our study,factorsmediatingreproductive not prebreeding body condition had a signifidecision differed between the sexes. In female
cant effecton reproductivesuccessfor females.
Blue Petrels, condition was the most important After laying, female Blue Petrels forage at sea
factor and may influence the ability of the in- while their mate incubatesthe egg for 12 days.
dividual to form the egg during the prelaying Breedingexperiencemay affectthe foraging efexodus.However, Meathrel et al. (1993) did not ficiencyof femalesand, consequently,the time
find a relationshipbetweeneggsize and female neededto restorebody condition.
body condition in the Short-tailedShearwater.
Our study of Blue Petrelsshowsthat condiIn male Blue Petrels, the constraints on future
tion in both males and females
reproductionare different.During prebreeding
visits, males undergo a long period without
feeding and they perform energy-consuming
activitiessuchascourting,digging and burrow
defense.Poor condition may render them less
able to perform theseactivitiesefficiently.Furthermore, condition probably determines the
duration of their prebreedingattendance.Birds
in poor conditionmay spendlesstime ashore,
productive decision and reproductive performance (males).The obvious linkages between
breeding experience, body condition, reproductivedecision,and reproductivesuccess
raise
the questionof the consequencesfor future survival or future fecundityof reproductiveeffort
madeby individualsfacingpoorenvironmental
with the consequent penalties on mate reten-
tion or re-mating ability and, therefore, breeding probability (Fisher 1976,Ollassonand Dunnet 1988).
Considering both membersof the pair, the
ability of the male to reproduce will mainly
affects the re-
conditions.
ACKNOWLEDGMENTS
The administration
of "Terres
Australes
and An-
tarctiquesFrangaises"
providedlogisticand financial
support.We thank: all the successivefield workers
involved in banding and recoveryon Mayes Island;
October1995]
BodyCondition
ofBluePetrels
971
F. Genevoisfor greatassistance
in the field; D. Besson,
ent:Energeticadjustments
in avianbreeding.Ardea 68:225-252.
N. Huin, T. Micol, and D. Capdeville for help in data
processing;S. Hall, D. Ainley, R. D. Woollet, and G. FISHER,
H.I. 1967. Bodyweightsin LaysanAlbatross
D. Schnell for critically commentingon the manuscriptand for help with the English;and L. Ruchon
for drawing the figures.
Diomedea immutabilis. Ibis 109:373-382.
FISHER,H. I. 1976. Some dynamics of a breeding
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FUGLER,S. R., S. HUNTER, I. P. NEWTON, AND W. K.
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