Multiple Incentives For Parental Optimism and Brood Reduction in Blackbirds

Multiple Incentives for Parental Optimism and Brood Reduction in Blackbirds
Author(s): Scott Forbes, Richard Grosshans and Barb Glassey

Source: Ecology, Vol. 83, No. 9 (Sep., 2002), pp. 2529-2541
Published by: Ecological Society of America
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Ecology, 83(9), 2002, pp. 2529-2541

C 2002 by the Ecological Society of America
MULTIPLE INCENTIVES FOR PARENTAL OPTIMISM AND BROOD

REDUCTION IN BLACKBIRDS
SCOTT FORBES,13 RICHARD GROSSHANS,14 AND BARB GLASSEY2
'Department of Biology, 515 Portage Avenue, University of Winnipeg, Winnipeg, Manitoba, Canada R3B 2E9
2Department of Zoology, University of Manitoba, Winnipeg, Manitoba, Canada R3T 2N2
Abstract. Why do parent birds hatch their young asynchronously? This phenotypic
handicap exacts a cost of reduced growth and elevated mortality from the last-hatched or
"marginal" offspring, while conferring advantages to the more senior "core" brood. David
Lack long ago proposed that marginal offspring allow parents to track resources that are
uncertain at the time of clutch initiation. If food is insufficient for the entire brood, hatching
asynchrony allows surplus marginal offspring to be culled from the brood efficiently. This
in effect represents a secondary adjustment of clutch size. Today Lack's hypothesis remains
a central, although controversial, component, of discussions of hatching asynchrony and
avian brood reduction. Most field workers report results inconsistent with Lack's hypothesis
(e.g., a failure of asynchronously hatching broods to produce more fledglings than do
experimentally synchronized broods) and have offered a variety of alternative explanations.
Previous workers have focused, with limited success, on single explanations
for the complex
phenomena of hatching asynchrony. Here we use parental optimism theory to study the
secondary adjustment of clutch size in an integrated framework emphasizing ecological
(tracking uncertain resources) and developmental (unpredictable survival, quality and/or
sex of progeny) uncertainty. We present the results of a 6-yr study of Yellow-headed
Blackbirds
(Xanthocephalus
xanthocephalus),
a marsh-nesting
icterid. Marginal offspring
that are the products of hatching asynchrony serve multiple functions simultaneously: in
tracking resources that vary temporally; as a hedge against hatching failure; and in progeny
choice via sex-biased brood reduction. The effect of the phenotypic handicap was diminished
both by improving ecological conditions and by hatching failure/experimental clutch reduction. Marginal offspring represent high variance progeny, and synchrony represents a
high variance strategy. Brood reduction occurred at low cost and was rendered more efficient
by hatching asynchrony, particularly under conditions of stringency.
Key words: brood reduction; core offspring; hatching asynchrony; marginal offspring; parental
optimism; phenotypic handicap; resource tracking; Xanthocephalus xanthocephalus; Yellow-headed
Blackbird.
INTRODUCTION
Fatal sibling rivalry is ubiquitous among organisms

as diverse as tropical trees with embryos that produce
sib-killing chemicals, mammals and sharks that impale
or cannibalize siblings in utero, and birds whose younger nestlings routinely fall victim to siblicide or socially enforced starvation (Mock and Parker 1997). A
common thread for this grim practice is an initial parental overproduction of progeny, by which parents create more offspring than can or will be reared to independence. Such "parental optimism" (sensu Mock
and Forbes 1995) inevitably results in subsequent competition for postzygotic parental care and, hence, sibling rivalry. A puzzling feature of this behavior is that
parents often play favorites within their brood. The
phenomenon of hatching asynchrony in birds, by which
first-hatched nestlings enjoy substantial growth and
Manuscriptreceived 10 October 2000; revised 13 June 2001;
accepted 16 July 2001; final version received 10 January2002.
3 E-mail: s.forbes@uwinnipeg.ca
4 Present address: Department of Botany, University of
Manitoba, Winnipeg, Manitoba, Canada R3T 2N2.
survival advantages over last-hatched nestmates (Mock

1984, Magrath 1990), provides a striking example. The
obvious question is why?
Explanations for avian hatching asynchrony fall into
three categories: (1) asynchrony is directly adaptive;
e.g., by allowing tracking of uncertain resources (reviewed in Pijanowski 1992); (2) asynchrony arises as
a sequela to other adaptive trade-offs; e.g., avoidance
of whole-brood loss (Hussell 1972, Clark and Wilson
1981); or (3) asynchrony is an unselected by-product
of an underlying constraint; e.g., the hormonal transition from egg-laying to incubation (Mead and Morton
1985). The adaptive significance, if any, of avian hatching asynchrony remains highly controversial, and much
discussion surrounds Lack's original suggestion that
parent birds lay more eggs than they normally expect
to rear as a hedge against uncertain future food. If food
proves to be plentiful, parents might rear extra chicks
and derive a reproductive bonus. Conversely, if food
proves short, hatching asynchrony facilitates the selective removal of the surplus progeny (Lack 1947,
1954, 1968). This represents a secondary adjustment
2529

2530
SCOTT FORBES ET AL.
of clutch size as more information about ecological

conditions becomes available (Forbes and Mock 1996,
Forbes et al. 2001).
Lack's hypothesis, however, fails as a general explanation for avian hatching asynchrony. It cannot explain, for example obligate brood reduction, in which
brood reduction is virtually certain, in a variety of
large, predatory birds (Mock 1984). As well, numerous
field tests in facultative brood reducers have produced
uneven results, stimulating field workers to propose a
variety of alternative hypotheses (reviews in Magrath
1990, Stoleson and Beissinger 1995, Stenning 1996).
As Magrath and Stoleson and Beissinger have emphasized, hatching asynchrony lies amid a series of coadapted traits, so no simple explanation is likely to
suffice. H. L. Mencken once noted: "For every complex problem there is a solution which is straightforward, simple and wrong." This pithy remark seems
particularly apt for the study of avian hatching asynchrony, which historically has emphasized single-factor explanations without obvious success.
Here we use the stochastic framework afforded by
the theory of parental optimism (Mock and Forbes
1995, Forbes and Mock 1996, 1998, 2000) to examine
the problem of avian family size and within-brood favoritism. Under parental optimism, surplus zygotes
serve a trio of adaptive functions: (1) tracking uncertain
resources such as food or gametes (Lack 1947, Temme
and Charnov 1987); (2) replacing failed or low-quality
offspring (an idea originally developed, with plants in
mind, by Buccholz [1922]; see Schtiz 1943, Dorward
1962, Lundberg 1985, Kozlowski and Stearns 1989,
Anderson 1990, Forbes 1990, Forbes and Lamey 1996,
Forbes and Mock 1998); and (3) facilitating the development of other family members (Mock and Forbes
1995).
Closely related to the problem of parental optimism

is why parent birds play favorites. Lack (1947) originally suggested that parent birds might reduce the cost
of surplus offspring by hatching nestlings asynchronously. This, in fact, is a special case of within-brood
favoritism, a taxonomically widespread trait (Mock and
Parker 1997) that we now know can include other forms
of phenotypic handicap, e.g., birth asynchrony as occurs in mammals; within-clutch variation in propagule
size or hormonal titer (Williams 1994, Schwabl et al.
1997). The phenotypic handicap in effect creates separate castes of "core" and "marginal" (those offspring
that are normally the first to succumb to partial brood
loss) progeny. Moreover, the phenotypic handicap creates a fundamental competitive asymmetry between
core and marginal progeny (Forbes et al. 1997, 2001,
Forbes and Glassey 2000). The fate of core progeny
has strong effects on that of marginal progeny; the
reverse is not true. The result is that core offspring are
largely buffered from the effects of overcrowding, and
parents can add extra marginal progeny to the brood
cheaply.
Ecology, Vol. 83, No. 9
We address the twin problems of parental optimism

and within-brood favoritism in the Yellow-headed
Blackbird (Xanthocephalus xanthocephalus), a marshnesting icterid with pronounced hatching asynchrony
(Richter 1984). Our work is framed around two key
questions. First, why do parent blackbirds play favorites? They do so by conferring the phenotypic handicap
of hatching asynchrony upon some of their progeny.
Does hatching asynchrony render brood reduction more
efficient, as Lack originally suggested, or is hatching
asynchrony unnecessary for a secondary adjustment of
clutch size as Clark and Wilson (1981) suggest? Second, why do parent blackbirds produce surplus marginal offspring? Here we focus on three potential functions of surplus progeny: resource tracking, insurance,
and progeny choice. Under resource tracking, the phenotypic handicap buffers core progeny from the inimical effects of food competition when resources are
scarce relative to demands (which can also vary; Forbes
and Mock 1996). Thus we expect greater variability in
growth and survival of marginal than of core nestlings.
Moreover, the phenotypic handicap must be reversible
when resources are plentiful, and the gap in growth and
survival between core and marginal nestlings should
narrow as feeding conditions improve. These are the
predictions of growth and mortality compression
(Forbes et al. 2001). Under the insurance hypothesis,
the phenotypic handicap buffers core progeny from the
effects of overcrowding for food or space, and again
must be reversible, but this time when preceded by the
early loss of other offspring, particularly the core brood
(Forbes 1990, Forbes et al. 1997, 2001). Thus we expect the survival of marginal nestlings to rise more
quickly than that of core nestlings when broods are
reduced either experimentally (by egg removal) or by
hatching failure. Under the progeny choice hypothesis,
parental optimism allow parents to upgrade mean offspring quality. In sexually dimorphic species such as
Yellow-headed Blackbirds, the lines between progeny
choice and resource tracking begin to blur. When resources are scarce, removal of male nestlings will yield
a greater benefit to the surviving nestlings than removal
of a female because of the greater nutritional demands
of males. Thus, we expect sex-biased brood reduction
under conditions of shortfall, with larger males suffering greater mortality.
METHODS
We studied Yellow-headed Blackbirds from midMay to July at sites near Winnipeg, Manitoba, Canada
in 1993, 1994, 1995, 1996, 1997, and 2000. The average clutch size ranged from 3.46 to 3.98 eggs over
the six years (Table 1), and the modal clutch size in
all years was four eggs. Nests were censused daily from
egg-laying until the eldest nestling reached 12 d of age.
Nestlings were not handled after the eldest chicks
reached 10 d of age to prevent premature fledging.
From 1994 on, eggs were marked as they were laid to

September 2002
PARENTAL OPTIMISM IN BLACKBIRDS
2531
monitor hatching sequence. Nestlings were colored nest, a conservative measure. Hatching spread was
with nontoxic felt markers to facilitate individual rec- based upon the number of days over which the nestlings
ognition, and colors were assigned at random to nest- hatched: e.g., a hatching spread of zero indicated that
lings. Nestlings were weighed daily to the nearest 0.1 all nestlings hatched between visits on consecutive
g with an Ohaus electronic balance (Ohaus, Pine Brook, days. We sexed nestlings using the method of Richter
New Jersey, USA). The frequency of parental food de- (1983, 1984) based upon the bimodal mass distribution
liveries was estimated from videotaped observations of chicks as they approach fledging. Yellowheads are
using 16X-zoom video cameras placed 3-8 m from strongly sexually dimorphic, with the average fledging
nests. Broods were videotaped for 4 h and nestlings mass of males being roughly 150% that of females; by
were weighed immediately before and after each taping the eighth day of the nestling period, the largest males
session to allow us to determine mass changes over the are more than twice the mass of the smallest females.
observation period. Nestlings were handled lightly be- The mass of 8-d-old chicks was used as an index of
fore a videotaping session to stimulate them to release growth performance. This age was chosen to allow for
fecal sacs. We used the relative mass changes of core comparisons between core and marginal chicks of the
and marginal nestlings over the observation period to same age, as marginal nestlings hatching two days beestimate the relative proportion of food received by hind core nestlings would be only 8 d old when core
each. For example, if in a two-nestling brood, the mass nestlings were last measured at 10 d old.
of a core nestling increased 2 g and that of a marginal
Statistical analysis
nestling increased 1 g over the same period, the relative
The data on nestling survival and hatching failure
food shares received by the core and marginal nestling
would be two-thirds and one-third, respectively. We were analyzed on a per brood basis to avoid pseudoused mass changes instead of food items received chief- replication (Hurlbert 1984). These data sets, as well as
ly because we could not always see which nestling those for hatching asynchrony, yielded polytochomous
received a particular food item. Parents appeared obliv- statistical distributions (limited numbers of discrete alious to the presence of cameras and resumed brooding ternatives) not amenable to conventional parametric
and feeding activities within minutes of our departure. statistics. In such cases, we used approximate randomMales assisted females in feeding nestlings at most ization for hypothesis testing (Edgington 1986, Noreen
nests when nestlings were older, but this contribution 1989, Manly 1991), where the null distribution is genwas relatively minor, as males made 6-31 % of the feed- erated by random permutation of the sample data on a
ing trips (x = 15.3%, n = 19 broods; excepting one computer and computed confidence intervals are obnest, where the female but not the male appeared to be tained by bootstrapping (Efron 1982). The methods
camera shy). Thus females delivered roughly five- used included pair-wise comparisons of means (test
sixths of all food items on average. The number of statistic D, the difference in sample means), one-way
feeding trips to the nest by both parents was used as ANOVA by randomization (test statistic T, equivalent
an index of parental input. This correlated strongly with to the within sum of squares), and factorial ANOVA
the rate of mass gain of individual chicks and the brood, by randomization (test statistic = SSb, the between sum
an estimate of the mass of food delivered to the nest of squares). All P values were based upon 25 000 ranper unit time. Videotaping was generally conducted domly generated permutations of the data set. Because
between 0700 and 1300 CST on days with fair weather data on nestling growth were drawn from continuous
for broods of 1-5 nestlings 2-10 d old. Broods were normal distributions, analyses were based on conventional parametric methods. All significance tests were
videotaped only once to avoid pseudoreplication.
We defined core and marginal offspring operation- two-tailed.
ally by the date upon which nestlings hatched. The core
Experimental methods
brood consists of those nestlings hatching on the first
=
In
1993 (n
22) and 1994 (n = 20), we artificially
day of the nestling period. Chicks hatching on subsequent days are defined as the marginal brood. When synchronized broods of Yellow-headed Blackbirds by
newly hatched nestlings were first found, we inspected exchanging nestlings across nests. Nestlings were usuthe clutch for pipped eggs, and we either returned later ally moved on the day of hatching (in a few cases on
in the day or used the mass distribution of nestlings the following day) to compress the normal hatching
the following day to assign such chicks to the core span among individuals within the brood. Most exbrood. Typically, in the modal clutch of four, the first- changes took <90 s to complete, and foster nestlings
and second-laid eggs hatched on day one, and the third- were quickly accepted into their new families, i.e., were
and fourth-laid eggs hatched one and two days later, fed by their foster parents. In most cases, the two lastrespectively (Richter 1984). For the purpose of as- hatched chicks in one nest were exchanged for the two
sessing hatching failure, we defined the first- and sec- first-hatched chicks in another, compressing the usual
ond-laid eggs as "core" eggs, and any subsequently 2-d hatching span to 1 d in one nest and 0 d in the
laid eggs as "marginal" eggs. We assessed hatching other. Because some of these broods are not completely
failure based on the presence of unhatched eggs in the synchronous, we shall hereafter refer to them as syn-

2532
SCOTT FORBES ET AL.
1. Interannual variation in components of reproductive performance in Yellow-headed Blackbirds with normal

hatching asynchrony in Winnipeg, Canada.
TABLE
1993
x
Reproductive component
1994
n
3.81
15.0
1.41
0.49
9.6-20.9
0.80
78
54
56
3.82
12.1
1.77
0.64
8.7-16.0
0.87
119
93
94
Total brood size at day 1

Core brood size at day 1
Marginal brood size at day 1
3.09
1.64
1.50
0.94
0.68
0.99
44
45
42
3.36
1.59
1.75
0.84
0.63
0.95
76
81
75
Core brood size at day 10

Marginal brood size at day 10
Total brood size at day 10
1.05
0.52
1.57
0.50
0.60
0.68
21
21
21
1.54
1.07
2.61
0.67
0.96
0.94
61
61
61
Clutch size
Hatching failure (%)t
Hatch span, all clutches (d)
Percentage of broods where day 10 brood
clutch size
SD
4.8
21
SD
20
60
Note: The sample size, n, is the number of clutches or broods.

t Bootstrap 95% confidence intervals are shown instead of standard deviation.
chronized, rather than synchronous, broods. Other than

the transfer of the chicks between nests, nestlings in
naturally asynchronous and experimentally synchronized nests were treated in an identical manner. Mean
clutch sizes for successful synchronized broods in 1993
(x = 3.70 ? 0.48 eggs, mean + 1 SD; n = 10 nests)
and 1994 (x = 3.80 + 0.56 eggs; n = 15) were virtually
identical to those in unmanipulated clutches (Table 1).
In June and July 2000, we performed reciprocal
swaps of nestlings on the day of hatching at 26 broods
as a sham control for the earlier synchronization experiment to determine if the effect of the manipulation
per se had any influence on the growth and survival of
the foster nestlings. Nestlings were handled in the identical manner as the earlier synchronization experiment,
except that individuals of the same age were exchanged
for each other. Fourteen core nestlings and six marginal
nestlings were exchanged in this manner at 20 broods
(one foster nestling per brood). We compared their
growth and survival to unmanipulated core and marginal nestlings in the same and other broods. Where
more than one unmanipulated core (or marginal) offspring was found in a brood, the brood mean was used
to avoid pseudoreplication. We found no detectable effect of this manipulation on either growth or survival;
the differences between the control and sham control
nestlings in factorial ANOVA by randomization were
small and nonsignificant (for growth, SSb = 11.50, P
=
0.58; for survival,
SSb = 0.01, P = 0.85).
We experimentally altered clutch sizes in 1995 to

examine the effect on core and marginal offspring survival. We followed the same protocol as Forbes et al.
(1997), adding or removing a single egg on the day of
laying. These manipulations generally took <90 s to
complete. Forbes et al. (1997), working on ecologically
similar Red-winged Blackbirds (Agelaius phoeniceus),
found no effect in an identical experiment performed
at the same time and location as the work reported here.
Brood reduction efficiency
We estimated the efficiency of brood reduction in
asynchronous and synchronized broods in the two years
of the experiment as the input into failed offspring. Our

currency for comparison was the total parental input:
estimated mass of food intake by an offspring just prior
to fledging (10 d of age). We used video observations
of parental feeding behavior to estimate age-specific
per capita provisioning rates, and determined the relative allocation of food to nestlings from mass changes
of individual nestlings during the 4-h observation sessions using the multiple regression equation: rate of
mass gain (grams per hour) = -0.071 + 0.42 nestling
age (days) + 0.155 yr + 0.165 nestling rank. Year and
rank were dummy variables (1993, 0, 1994, 1; marginal
nestling, 0, core nestling, 1). All individual regression
coefficients were significant (P < 0.008), as was the
overall regression model (F3 127 = 11 -l0, P < 0.0001)
From this equation, we were able to estimate the agespecific intake of core and marginal nestlings. We used
these behavioral data, in conjunction with observations
of the age at death of core and marginal nestlings in
synchronized and asynchronous broods, to determine
the per capita input prior to death. We also computed
the cost of rearing a core nestling to 10 d old (nestlings
fledge at 11-13 d old), and used this as a benchmark
for comparison. Brood reduction efficiency was estimated from VA/VS where vi is the mean per capita input
in victims of brood reduction, and the subscripts A and
S denote asynchronous and synchronized broods, respectively. We calculated vi according to
( jC E
f(t, M)
t=I f(t, C)) + jME
t=l
nC +
nM
(1)
Here, j denotes chick, C and M denote core and

marginal nestlings, and f(tC) and f(tM) represents the
food consumed by nestling j on day t of the nestling
period by core and marginal chicks, respectively; td is
the last day a chick was observed alive, and nc and nM
are the numbers of core and marginal victims of brood
reduction. In words, we estimated the total food con-

TABLE
1.
2533
September 2002
Extended.
2000
1997
1996
1995
1SD
1 SD
1 SD
3.98
12.1
1.60
0.45
6.3-18.3
0.90
48
26
45
3.67
9.0
1.83
0.87
0-19
1.33
9
5
6
3.80
21.8
1.33
0.36
9.0-37.2
0.65
14
13
12
3.46
18.2
1.11
0.69
10.8-26.2
0.90
54
30
35
3.50
1.77
1.73
0.65
0.71
0.78
28
28
28
3.33
1.50
2.20
0.82
0.55
0.84
6
6
5
3.14
1.71
1.43
1.03
0.73
0.76
14
14
14
2.74
1.61
1.14
0.98
0.60
0.91
35
38
35
1.71
1.17
2.88
0.69
0.82
0.85
26
26
26
1.25
1.25
2.50
0.50
0.96
0.58
4
4
4
1.78
0.67
2.44
0.67
0.50
0.73
9
9
9
1.43
0.62
2.05
0.92
0.81
0.92
21
21
21
24
Xc
20.8
sued by core victims of brood reduction prior to their

demise (the first term in the numerator on the right
hand side (rhs) of Eq. 1) and similarly for marginal
victims of brood reduction (the second term in the numerator on the right-hand side of Eq. 1. We divided
these sums by the total number of core and marginal
victims of brood reduction (denominator on the rhs of
Eq. 1) to arrive at an overall mean per capita investment
in victims of brood reduction. We computed confidence
intervals for the costs of rearing brood reduction victims in a Monte Carlo simulation. We generated bootstrap samples for age at death for each nestling category, and used the point estimates (means of the
coefficient and standard error of the estimate) to generate random Gaussian distributions for the per capita
provisioning and mass change regressions. The simulated coefficients were used to estimate age-specific per
capita provisioning for core and marginal nestlings in
synchronized and asynchronous broods in each of 5000
simulation runs for both 1993 and 1994. In each simulation run, we calculated the cost of a brood reduction
victim in asynchronous and synchronized broods, and
computed the ratio of the two estimates according to
Eq. 1; unity indicates equal brood reduction efficiency
in synchronized and asynchronous broods. A value
< 1.0 indicates greater efficiency of brood reduction in
asynchronous nests; a value >1.0 indicates the opposite. We also computed two further measures in our
simulations: (1) the proportion of cases in which the
efficiency ratio was less than unity; and the proportionate cost of a brood reduction victim in relation to
rearing a core nestling to 10 days of age (a rough estimate of the pre-fledging costs of parental input).
RESULTS
We start by establishing a prima facie case for parental optimism. Was there evidence of an initial overproduction of offspring? Indeed there was. Over the
study, only 23 of 143 (16.1%) unmanipulated broods
fledging at least one nestling succeeded in rearing
broods that matched the initial clutch size (Table 1).
Partial brood loss or hatching failure occurred in the
1 SD
25
20
other 120 broods. Thus, parents generally began with

more incipient offspring than were reared to independence.
Parental optimism and resource tracking
We observed wide variation in reproductive success
across years (Table 1): 1993 showed the lowest fledging
success ever reported for Yellow-headed Blackbirds
(see Richter 1984, Beletsky and Orians 1994, Barber
and Evans [1995] and references therein). From this
literature, in conjunction with our own results, we obtained data on the average fledging success of Yellowheaded Blackbirds in five populations. From this, for
the purpose of comparison, we computed the mean
number of fledglings produced in a successful nest (i.e.,
that yielded one or more fledglings) as 2.44 + 0.38
fledglings (mean + 1 SD; n = 16 breeding seasons).
We used these data to transform observed reproductive
success in our population (unmanipulated nests only)
to standard normal deviates, which we interpret as a
crude index of environmental quality, with the following results: 1993, Z = -2.29; 1994, Z = +0.45; 1995,
Z
+ 1.16; 1996, Z = +0.16;
1997, Z = 0.00; 2000,
Z = - 1.03). These results suggest that 1993 was a

classic "bad" year and 1994 and 1995 were above
average or "good" years. Our data on fledging masses
are consistent with this conclusion: chicks fledged in
1993 were the smallest of the six years in the study by
a wide margin. Thus there was a strong contrast in
environmental quality in the two years (1993 and 1994)
of the synchronization experiments.
Yellow-headed Blackbirds showed significant interannual variation in some, but not all, components of
reproductive performance (Table 1). We note that data
for 1996 and 1997 are based on small samples. Variation across years in clutch size (6-yr mean = 3.76
eggs) and brood size at hatching (6-yr mean = 3.19
nestlings) was modest, with coefficients of variation of
4.6% and 8.4%, respectively. By contrast, the brood
size at day 10, just prior to fledging, ranged from 1.57
to 2.88 nestlings (6-yr mean = 2.34 nestlings), with a
19.8% coefficient of variation. Most variation in the

2534
SCOTT FORBES ET AL.
number of fledglings, therefore, stemmed from posthatch variation in nestling survival and not from differences in clutch size and hatching success. Table 2
shows the coefficients of variation in the mean numbers
of core and marginal offspring at hatching and at day
10; from this, two results are obvious: variability in
offspring number increases with age, and the numbers
of marginal offspring are more variable, particularly
near fledging age, than are the numbers of core offspring. These results mirror those obtained by Forbes
et al. (2001) for Red-winged Blackbirds.
Multiple regression analysis showed that parental delivery rates (number of food deliveries per chick per
hour) increased with brood age (t = 6.19, P < 0.001),
but when year and treatment (asynchronous vs. synchronized) were added as dummy variables to the bestfit regression model, neither explained additional var-
12 -
0.4
1994
1993
10
.0.3
8
45
1a
V~~~~~~~~~~~~
4~~~~7
(6
0.2(
~~~~~~~C,
Ca ~
60
0
iation in feeding rate (for year, t = -0.09, P = 0.93;

for treatment, t = -0.34, P = 0.73) and hence were
deleted. Evidently, parents did not step up deliveries

to synchronized broods, nor did they make more trips
in 1994 than in 1993. Multiple regression analysis revealed that the rate of mass change (in grams per hour)
for individual nestlings during videotape sessions in-
2and magia
was higher in core than in marginal chicks (t = 2.70,

2.76, P = 0.007). The year X chick interaction (core
vs. marginal) was weak and nonsignificant (t = 1.09,
P = 0.28),
as was treatment (synchronization)
(t =
-0.11, P = 0.92), and these two terms were deleted

from the best-fit model. These results suggest that parents brought less food per feeding trip in 1993, while
holding the number of trips steady.
We examined the age at which victims of brood reduction perished in asynchronous and synchronized
broods in 1993 and 1994 (Fig. 1). There were three
main results. First, marginal nestlings in asynchronous
broods died earlier than core nestlings (all comparisons
are by factorial ANOVA by randomization: SSb = 46.5,
P < 0.0001). Second, nestlings in asynchronous broods
died earlier in 1993 than in 1994 (SSb = 14.8, P =
0.02). The effect, however, was much weaker in synchronous broods in 1994 (SSb = 1.7, P = 0.39). Third,
the age at death of marginal nestlings, the most frequent
victims of brood reduction, was almost identical in synchronized and asynchronous broods in a given year
(Fig. 1; SSb = 0.43, P = 0.67). The data are too few
for core nestlings to reach any definitive conclusion
(Fig. 1;
SSb
= 7.8, P = 0.13).
The survival of core nestlings in asynchronous

broods varied modestly across years, from 80% to 96100% (Fig. 2; the 100% survival observed in both 1996
and 1997 is based on small samples and is probably
an overestimate). By contrast, the survival of marginal
nestlings in asynchronous broods varied widely over
the same time span, from 41% to 72% (Fig. 2). Across
all years, the survival of core nestlings in asynchronous
ga
bas0htaevcisoro
0.0
__ha
creased linearly with nestling age (t = 2.68, P = 0.008),

P = 0.008), and was higher in 1994 than in 1993 (t =
6etig
FIG. 1. Mean age at death of core nestlings (open bars)

and marginalnestlings (gray bars) that are victims of brood
reductionin asynchronous(A) and synchronized(S) broods
of Yellow-headedBlackbirdsin 1993 and 1994, with sample
sizes and95%bootstrapconfidenceintervals.The right-hand
y-axis shows the efficiency of brood reduction(medianand
95% ci estimatedin MonteCarlosimulation)in asynchronous
(circles) and synchronized(diamonds) broods of YellowheadedBlackbirdsin 1993 and 1994. Here, parentalinputto
victims of broodreductionis estimatedfrombehavioralstudies and weighted accordingto the fractionof offspringthat
perish and the age at death; it and is measuredagainst the
cost of rearinga core nestling to 10 d of age (n).
broods was significantly higher (Fig. 2; factorial ANOVA by randomization, SSb = 9.2, P < 0.0001), and
less variable than that of marginal nestlings (Table 2).
In synchronized broods, the survival of core and marginal offspring was sharply higher in 1994 than in 1993
(Fig. 2; SSb = 1.55, P = 0.0001), and core nestlings
in synchronized broods enjoyed higher survival than
marginal nestlings (Fig. 2; SSb = 0.82, P = 0.006). In
1993, the survival of both core and marginal offspring
was higher in asynchronous than in synchronized
broods
(SSb
= 0.65, P = 0.03). In 1994, however, the
survival of core and marginal offspring in synchronous

and asynchronous broods differed little, if at all (SSb
=
0.0005,
P = 0.96).
One further measure of reproductive performance is

simply the number of nestlings that reach fledging age,
in our case, 10 d old. Asynchronous broods produced
an average of 1.57 and 2.61 nestlings per successful
nest in 1993 and 1994, respectively (Table 1); synchronized broods produced, on average, 1.60 ? 0.70

September 2002
1.0
Asynchronous
42-48 -266947426
Synchronized
.
16
2535
2. Coefficients of variation (%) in the mean numbers

of core and marginal offspring at hatching and at day 10
from asynchronous broods.
TABLE
~~61
21
At hatching
15
39
Year
Marginal
brood
Core
brood
Marginal
brood
1993
1994
1995
1996
1997
2000
All years
41.5
39.6
40.1
36.7
42.7
37.0
39.6
66.0
54.3
45.1
38.2
53.1
79.8
56.1
33.3
43.5
40.4
40.0
37.6
41.8
39.4
115.4
89.7
70.1
76.8
74.6
130.0
92.8
21
0.88
10
57
> 0.6
At day 10
Core
brood
a)
Parental optimism and insurance
Z 0.4-
5
0.2
0.0
1993 1994
1993 1994 1995 1996 1997 2000
FIG. 2. Mean survival of core (open bars) and marginal

nestlings (gray bars) in synchronized and asynchronous
broods of Yellow-headed Blackbirds in 1993, 1994, 1995,
1996, 1997, and 2000. Sample sizes and 95% confidence
intervals are shown.
We compared the hatching spans of clutches of three

and four eggs in 1993 and 1994 by approximate randomization; there were too few data from clutches of
two and five eggs for a similar comparison. This analysis revealed that hatching spans were substantially
shorter in 1993 than in 1994 (factorial ANOVA by
randomization, SSb = 2.49, P = 0.02). We also examined interannual variability in the rate of hatching
failure (calculated on a per brood basis to avoid pseudoreplication): this ranged widely, from 9% to 22% per
egg, but not significantly across years (Table 1; AN471
44
nestlings (n = 10 nests) in 1993 and 2.81 ? 0.98 nestlings (n = 16 nests) in 1994. In short, asynchronous
and synchronized broods produced virtually identical
numbers of fledglings in 1993 (t = 0.11, df = 29, P
= 0.91), and synchronized broods produced slightly
more nestlings than asynchronous broods in 1994 (t =
0.78, df = 75, P = 0.44).
Core nestlings in asynchronous broods enjoyed superior growth, and marginal nestlings in asynchronous
broods exhibited roughly comparable growth to core
chicks in synchronized broods in 1993 (Fig. 3). Twoway parametric ANOVA indicated a strong effect of
experiment (F146 = 4.76, P = 0.034) and a weaker
effect of chick type (F,,46 = 3.91, P = 0.054) in 1993.
In 1994, there was a stronger effect of chick type (F. 118
= 6.43, P = 0.0 13), but a weaker effect of the experiment (F1 118 = 0.99, P = 0.21). Thus, in the good year
the gap between asynchronous and synchronized
broods narrowed (Fig. 3). The masses of both core and
marginal nestlings in asynchronous broods in 1993
were substantially below those observed in all other
years (Fig. 3; the mass of core nestlings in other years
was 32.2-35.6 g; the mass of marginal nestlings in
other years was 28.6-32.8 g) 1993 was indeed a very
bad year.
41
1996
1994
38 -
1M9
44
(number ofbod)aeson28
1 46
35,
Cn?3
E 32
CD
1997
9
2000
1
3j59
Cu
':
1995
44,
1993
20
20
1714
Mean mass of 8-d-old core nestlings (open symbols) andmarginalnestlings(solid symbols)in asynchronous
(circles) and synchronizedbroods (diamonds) of Yellowheaded Blackbirds in 1993, 1994, 1995, 1996, 1997, and
2000. Mean, 95% confidence intervals, and sample sizes
(numberof broods) are shown.
FIG. 3.

2536
SCOTT FORBES ET AL.
64
1.0
75
HF
ENHF

6
1.0
1
4211
Reduced
Control
Enlarged
9
49
0.8
_~
_79
.20.6 -
0.8
> 0.6-
Z 0.4/
0.2
0 .0
0.0
Core nestlings
..
. ..
..
Marginal nestlings
4. Mean survival of core and marginal nestling Yellow-headed Blackbirds to fledging age in broods with hatching failure (HF) and with no hatching failure (NHF). Vertical
lines show 95% confidence intervals. Sample sizes (number
of broods) are shown above bars.
FIG.
OVA by randomization, T = 0.19, P = 0.43). The

survival of marginal offspring rose sharply at nests in
which one or more eggs failed to hatch (Fig. 4; comparison of means by approximate randomization, D =
0.15, P = 0.05). In contrast, hatching failure had virtually no effect on the survival of core nestlings (Fig.
4; D = 0.01, P = 0.83). We examined the survival of
core and marginal offspring with experimentally enlarged and reduced clutches in 1995 and obtained results parallel to those for hatching failure. For both
core and marginal offspring, survival was highest in
reduced clutches and lowest in enlarged clutches, although the effect was greater for marginal offspring
(Fig. 5). The overall effect of experimental alteration
of clutch size was not quite significant, probably due
to modest sample sizes and substantial variability (Fig.
5; factorial ANOVA by randomization, SSb = 0.354, P
0.5
~ ~
37 ..
.......
0.2A
0.4 21
35..
27.......
~~ ~ ~
CD~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..
E
. . .
C,,
...
..
.
16~~~..
0.6broods)
above the bars.
0.2
0.1
~ ~
0.0
~ ~
.......
.......
1993
1995.200
1994.
6.Prportion .o maesi.snhrnze
.grybas
and asynchrnous
brood (oenbrs.f.elo.eae
Coe nestlIng
Margina~l
ntlIng
Blackbirs
in 193, 1994
.5
195.ad200.oosra
FIG
5.
Suvalo
cor
marina
an
netln
confienceintevalsarehown
wih.amlesze.(ume Yellowof broods)
above the.bars.
FIG.
heade Blcbid
Portion8
ofmlds innssyincbrooizds fgromexpr-)
imntallyncreduced conrols aopnd exprimentaYllyw-enarged
bersiabove batrvasarshownwt sample sizes (numberofbod)
= 0.065).
Parental optimism and progeny choice

The sex ratio of nestlings approaching fledging (810 d old) varied modestly across years (ANOVA by
randomization, T = 0.70, P = 0.13), but differed significantly between synchronized and asynchronous
broods (Fig. 6; factorial analysis by randomization, SSb
= 0.49, P = 0.04). Fewer males, evidently, survived
in synchronized broods. This effect was most clear in

September2002
PARENTALOPTIMISMIN BLACKBIRDS
1993, when only a single male out of 10 synchronized

broods reached fledging age.
Is brood reduction efficient?
In both 1993 and 1994, brood reduction victims died
earlier in asynchronous broods than in synchronized
broods (Fig. 1; factorial ANOVA by randomization for
1993,
SSb =
0.65, P = 0.03; for 1994,
SSb =
0.36, P
= 0.03). This result is consistent with nearly all other

brood synchronization studies (see review in Magrath
1990). Curiously, the age at death was greater in 1993
(the "bad" year) than in 1994 (the "good" year) for
both asynchronous and synchronized broods (Fig. 1;
factorial analysis by randomization; asynchronous
broods, SSb = 14.8, P = 0.02; synchronized broods,
SSb = 1.55, P = 0.0001). One might expect earlier
rather than later brood reduction when resources are
scarce. This mystery, however, is easily resolved. In
both years, brood reduction occurred substantially earlier in marginal than in core chicks, and in 1994, nearly
all core chicks survived. In 1993, however, core survival was lower than in any other year (Fig. 2) in both
synchronized and asynchronous broods, which died relatively late in the nestling period. The apparent delayed
onset of brood reduction in 1993 is a result of a higher
proportion of core offspring in the partial brood loss
sample that year. Evidently, 1993 was not just a "bad"
year: it was a very bad year.
Interestingly, there was no difference in the age at
death within an offspring class (core or marginal) in
asynchronous and synchronized broods. The earlier age
at death in asynchronous broods is due simply to the
fact that of those chicks that died, there were a higher
proportion of marginal offspring in asynchronous than
in synchronized broods. The key result is that core
offspring die later than marginal offspring, and in synchronous broods, a higher proportion of core offspring
perish. We suspect that this fact explains the oftenobserved pattern of apparently delayed brood reduction
in synchronized broods (see the review in Magrath
1990).
We compared the efficiency of brood reduction in

synchronized and asynchronous broods in 1993 and
1994 by estimating the input that parents made in
chicks before their death. Our results indicate that parents with asynchronous broods enjoyed greater brood
reduction efficiency in both years, markedly in 1993,
but only marginally so in 1994 (Fig. 1). In 1993, the
average cost of a brood reduction victim in an asynchronous brood was only four-fifths that in a synchronized brood. In 1994, that cost rose to 97%, a slim
advantage, if any. We also estimated the relative likelihood that brood reduction was cheaper from the distribution of efficiency ratios in our Monte Carlo simulation. In 1993, that likelihood was 9.97 to 1; in 1994,
it was 1.41 to 1. These are, in effect, Bayesian estimators where the prior probabilities for each outcome
were equal (0.5). Our results also show that brood re-
2537
duction in both synchronized and particularly asynchronous broods occurred at low cost, from 24% to
30% of the input required to sustain a core nestling to
10 d of age (Fig. 1). Perhaps most importantly, brood
reduction was cheaper in 1993, the year of greatest
stringency, than in 1994 for asynchronous broods, but
not synchronized broods.
DISCUSSION
The regulation of avian family size is a complex

question because it consists of numerous deterministic
and stochastic elements that are thoroughly intertwined. The now-familiar trade-off between offspring
number and quality, a central focus of life history theory (for a discussion specific to birds, see Amundsen
and Slagsvold 1996), belongs to the deterministic class.
With finite resources, parents must choose between a
small number of progeny with a large per capita investment and a larger number of progeny with a smaller
investment. Superimposed upon this simple trade-off,
however is uncertainty in ecology and development.
Ecological conditions vary from location to location,
microenvironment to microenvironment, minute to minute, week to week, and year to year. Uncertainty in
offspring development stems from an array of causes
ranging from meiosis to mishaps during birth or egglaying. Although some of this variety of life is predictable (e.g., seasonal variations in weather conditions), much is not.
Yellow-headed Blackbirds engage in a reproductive
lifestyle that, at times, appears to involve profligate
waste. In some years, more than half of all offspring
fall victim to partial brood loss, and almost no parents
succeed in rearing a full brood. The obvious question
is why? Our results suggest that parental optimism,
coupled with within-brood favoritism (by imposing
phenotypic handicaps), is an adaptation to the twin
challenges of ecological and developmental stochasticity: resource supplies and demands are uncertain, as
are the developmental fates of individual progeny. Our
results showed that asynchronous hatching yielded the
greatest benefits to parent blackbirds when needed
most: in 1993, the year of lowest nestling growth and
highest mortality (Fig. 1). Conversely, asynchrony
yielded little benefit (or cost) in a "good" year. The
greater efficiency of brood reduction in asynchronous
broods stemmed not from earlier brood reduction per
se, but rather from the fact that a greater proportion of
core nestlings perished in synchronized than in asynchronous broods. There were, in fact, no obvious differences in the age at death of marginal nestlings between asynchronous and synchronized broods in either
year. Because marginal nestlings perish before core
nestlings, our data give the appearance of an accelerated onset of brood reduction in asynchronous broods
(as a greater proportion of brood reduction victims are
marginal nestlings), but such data are misleading. The
question is more who is dying, rather than when core

2538
SCOTT FORBES ET AL.
and marginal nestlings are dying. Here the concept of

core and marginal offspring is essential to unraveling
the mechanism of brood reduction. Most previous studies of brood reduction that have examined age at death
report that brood reduction occurs earlier in asynchronous broods (reviewed in Magrath 1990).
Multiple incentives for parental optimism
Resource tracking.-To test the resource-tracking
hypothesis with a synchronization experiment, we
needed to verify two key assumptions: (1) that the requisite ecological variation had been encountered to provide a robust test of the resource-tracking function of
parental optimism; and (2) in the context of the brood
synchronization experiment, that the results were not
confounded by changes in parental effort and, in particular, that parents did not elevate provisioning rates
to synchronized broods.
It is clear that the first three years of the study provided the wide contrast in resource levels that one
would desire in a multi-year test of Lack's hypothesis:
1993 was a classic "bad" year and 1995 was a "very
good" year. Thus we are satisfied that we encountered
a sufficient range of ecological variability to provide
a fair test of Lack's hypothesis. Assessing provisioning
level is also crucial to interpreting the results of synchronization experiments, as parents may buffer nestlings from the effects of increased food competition by
stepping up workloads (Mock and Forbes 1994). Yellowhead parents, however, did not work harder to provision experimentally synchronized broods (see Results), thus removing a second impediment to providing
a strong test of Lack's hypothesis.
Our experimental results provide strong support for
the quality assurance hypothesis (Slagsvold 1986,
Slagsvold et al. 1995, Amundsen and Slagsvold 1996);
Amundsen and Slagsvold (1998) provide an example
using information on nestling mass in tandem with survival data. We view this hypothesis as a permutation
of the logic of Lack's resource-tracking argument. In
the most stringent year, 1993, the average mass of nestlings in synchronized broods fell below that of both
core and marginal nestlings in asynchronous broods.
In 1994, a relatively "good" year, the average mass of
nestlings in synchronized broods fell between that of
core and marginal nestlings in asynchronous broods
(Fig. 2). Thus, synchrony most certainly resulted in a
sacrifice in offspring quality under the conditions of
ecological stringency, evidently a result of delayed
mortality within synchronized broods (Fig. 1). In a year
of comparative ecological bounty, the picture is less
clear. Both the highest and lowest quality offspring
came from asynchronous broods. The advantage or disadvantage of synchrony under such conditions will depend upon the precise relationship between fitness and
fledging mass (Amundsen and Slagsvold 1996), for
which information is generally scant.
Survival of marginal chicks showed a nearly twofold
range of variation, whereas the survival of core chicks

varied within a high and narrow range. The survival
of core offspring in 1993, the "bad" year, plunged to
levels roughly in the midpoint of the range of survival
of marginal nestlings in asynchronous broods across
years (Fig. 2). By contrast, the survival of core and
marginal nestlings in asynchronous broods showed no
overlap over the same time span. Evidently, hatching
asynchrony does buffer core offspring from overcrowding for food or space under stringent conditions. Marginal offspring in synchronized broods fared no better
in 1993, when their survival crashed to levels well
below that in seen in asynchronous broods (Fig. 2).
Our work suggests that marginal offspring are a caste
of high-variance progeny, consistent with predictions
of the resource-tracking component of parental optimism. The survival of marginal offspring varied more
widely than that of core offspring both within and
across seasons, due chiefly to post-hatch variation in
marginal chick survival. For example, in all six years,
the coefficient of variation for the number of marginal
nestlings at day 10 was considerably greater than that
for core nestlings, although only modest differences in
the coefficient of variation existed for the size of the
core and marginal brood at hatching (Table 2). Growth
and survival both exhibited far greater variation across
years for marginal than for core chicks. Intriguingly,
variability in the growth of core nestlings rose sharply
in synchronized broods (Fig. 7). Because variance
alone can affect fitness, asynchrony may yield a benefit
over synchrony from bet-hedging (Amundsen and
Slagsvold 1998), although such an advantage is likely
to be modest in iteroparous species (Seger and Brockmann 1987).
Our results also show quite clearly that the phenotypic handicap of hatching asynchrony serves to buffer
the core brood from the mortality costs of resource
shortfall, thus verifying the prediction of mortality
compression from the resource-tracking hypothesis.
Even in the worst year, 1993, the average survival of
core progeny exceeded the average survival of marginal progeny in the best year. However, the phenotypic
handicap imposed upon marginal offspring was indeed
reversible. In the four years of the study when sample
sizes were relatively large (1993-1995 and 2000), we
can see that the average survival of core offspring rose
relatively modestly with improving ecological conditions, whereas the survival of marginal offspring rose
sharply (Fig. 2). Thus, as expected under resource
tracking, the fate of marginal offspring was conditional
upon prevailing ecological conditions; the fate of core
offspring was much less so. Still, improving conditions
for growth did not erase the entire cost of the phenotypic handicap for marginal offspring, as the best average survival for marginal offspring still fell below
the range of average survival rates for core offspring.
As we shall see, the extent of the cost of the phenotypic

September2002
Synchronized
Asynchronous
2539
ing failure had no obvious effect upon the survival of

core offspring (Fig. 4). Barber and Evans (1995), also
working on Yellow-headed Blackbirds, similarly noted
core
8IGD
that
the survival of marginal offspring rose in nests
marginal
30re
*nd
with hatching failure. Others report the same patterns
in other species (Smith 1988, Stouffer and Power 1990,
30~~~~~~1
Forbes et al. 1997, 2001). These results support the
~~~~25 ~
~
~ ~ 3
prediction of compensatory survival from insurance
theory
(Forbes and Lamey 1996). Our experimental
29
14
clutch manipulations parallel this pattern. The survival
C
20
of marginal offspring rose in nests where clutches were
Cl
6
experimentally reduced and fell where clutches were
obs9
aac21
chi
experimentally enlarged. The effects upon core off4
-.2.
4
spring, although in the same direction, were much more
15
modest (Fig. 5). Interestingly, the survival of marginal
9
nestlings in experimentally reduced broods approached
0
the survival of core offspring. Again, we find evidence
that marginal offspring survival is strongly contingent
0
upon developmental uncertainty, whereas core offspring, buffered by the phenotypic handicap imposed
5
upon marginal offspring, are only weakly affected.
Progeny choice.-Under progeny choice, parents use
initial parental optimism to create an expanded array
of incipient progeny from which the lowest quality in1993 1994 1995 1996 1997 2000
1993 1994
dividuals are culled and the highest quality individuals
FIG. 7. Coefficient of variation in the mass of 8-d-old are retained (Buccholz 1922, Kozlowski and Stearns
core and marginalYellow-headedBlackbirdnestlingsin syn- 1989). For example, individuals with congenital dechronizedand asynchronousbroodsin 1993-1997 and 2000.
Sample sizes (numberof broods) are shown above the bars. fects may be replaced by healthy progeny, for which
the crucible of sibling competition may serve as a tool
of identification (Forbes and Mock 1998). One path by
handicap is also contingent upon developmental un- which offspring quality can be enhanced is to eliminate
the most expensive progeny when resources are scarce.
certainty.
Male Yellow-headed Blackbirds fledge at masses that
In an important paper, Pijanowski (1992) established
that hatching asynchrony for the purpose of adaptive are roughly 150% those of females and thus are clearly
brood reduction could be favored even if it exacts a more expensive to rear than females. This is also the
survival cost for marginal offspring relative to core case in Red-winged Blackbirds and Great-tailed Grackoffspring in "good" years. There are two key prop- les, (Quiscalus quiscalus) (Fiala and Congdon 1983,
erties of Pijanowski's model: the handicap (1) ensures Teather and Weatherhead 1989). We observed that the
swift elimination of the marginal offspring in bad normally female-biased sex ratio was exaggerated duryears; and (2) is reversible, at least in part, in good ing times of stringency, especially so in synchronized
years. Our results provide substantial support for the broods. By thwarting the normal relief of partial brood
Pijanowski model. Asynchrony resulted in earlier loss, synchronization in 1993 appeared to elevate the
brood reduction by ensuring that marginal, but not core, costs of rearing nestlings that, in retrospect, were
offspring perished. And as we have shown, the phe- doomed from hatching. By shifting male nestlings from
notypic handicap was partially reversed by improving the marginal brood, where they are swiftly removed in
ecological conditions. That there was still a residual times of shortfall, to the core brood, where they are
mortality cost, even in the best years, of marginal off- not, experimental synchronization elevated the costs of
spring status is potentially explained by the Pijanowski brood rearing. Evidently, this surtax of reduced brood
model as the cost of a brood reduction strategy, which reduction efficiency made it even less likely that any
is balanced against the benefit of efficient brood re- of the more expensive males would survive to fledging
duction in bad years. But was brood reduction indeed in synchronized broods.
efficient?
Although offspring quality can vary independently
Insurance.-Our results also provide strong support of offspring number (as in the case of congenital defor the insurance component of parental optimism, by fects), differential costs of rearing males and females,
which marginal offspring serve as replacements for density dependence, and stochastic resources blur the
core progeny that fail early and unexpectedly. The sur- distinctions between resource-tracking and progeny
vival of marginal offspring rose sharply in broods in choice: there are no sharp lines to be drawn. Similarly,
which one or more eggs failed to hatch, whereas hatch- whether an insurance offspring will be required at any
35

2540
SCOTT FORBES ET AL.
given time will be contingent upon prevailing resources

and the extent of losses within the core brood. In 1993,
even three nestlings were too many for nearly all parents, particularly if one or more of the nestlings were
males. Thus, for parents laying a clutch of four, with
the most common pattern of two core and two marginal
eggs, the entire marginal brood was rendered redundant
if both core offspring hatched.
The phenotypic handicap and brood
reduction efficiency
We have argued that the phenotypic handicap is reversible under both the resource-tracking and insurance
functions of parental optimism. But is the phenotypic
handicap even necessary? Clark and Wilson (1981)
suggested otherwise. Small initial differences among
contemporary brood mates widen as the chicks grow
older, potentially providing the competitive asymmetries required for brood reduction to function. Indeed,
in both synchronized and asynchronous broods in our
study, the quantity of resources invested in the victims
of brood reduction was only a fraction of that invested
in core chicks prior to fledging (Fig. 1). This economical expenditure of resources is attributable to the joint
effects of reduced provisioning of marginal nestlings
and an early age at death, generally before the period
when provisioning costs rise sharply. Whether efficient
brood reduction can arise from initial synchrony is,
however, another question. In our study, asynchrony
shaved the costs of rearing the doomed chicks by ensuring that more of the cheaper marginal nestlings perished, particularly in the bad year of 1993, when such
economies were clearly most valuable. We note parenthetically that, although the mortality of marginal
offspring was higher in synchronized broods, there
were more marginal offspring in asynchronous broods
and, hence, in absolute terms, more of these died.
Would parents have been better off without marginal
offspring? That is, was asynchronous hatching costly?
The results of our synchronization experiment show
that the answer is clearly no. Synchronizing broods
resulted in lower, not higher, survival rates of both core
offspring in 1993 and 1994. Moreover, synchrony exacted a cost of reduced chick growth when the food
supply was short, as the mass of synchronized chicks
fell below that of both core and marginal offspring in
asynchronous nests. In 1994, when food was relatively
abundant, the mass of synchronized chicks was intermediate between that of core and marginal chicks in
asynchronous broods. Of course, we cannot rule out
the possibility that synchronized broods would have
outperformed asynchronous broods under even better
feeding conditions.
Our results show that the phenotypic handicap plays
a central role in keeping initial parental optimism affordable. Parents must play a balancing act between
having too few progeny when ecological conditions are
good and too many when they are bad (Forbes 1991).
By typically laying four eggs, Yellow-headed Blackbird parents will usually start with at least three hatchlings, ensuring that opportunities for rearing larger
broods are not missed when ecological conditions are
favorable. However, the presence of redundant progeny
is potentially costly, because overcrowding may ensue
under conditions of resource shortfall. This study, in
conjunction with the earlier work of Forbes et al. (1997,
2001) on Red-winged Blackbirds, shows that the buffer
created by the phenotypic handicap keeps such costs
manageable for core offspring, which become relatively more valuable during periods of stringency. Simultaneously, the value of marginal offspring falls,
particularly if a nestling is an expensive male. With
this built-in fail-safe mechanism, parents can afford to
be optimistic about their initial clutch size.
ACKNOWLEDGMENTS
This work was funded by NSERC (NaturalScience and

EngineeringResearchCouncil of Canada)researchgrantsto
L. S. Forbes. We thankN. Atmuri,L. Earle, 0. Griffith,S.
Jayakumar,A. Riedel, I. Schellenberg,K. Skibo, D. Swedlo,
S. Thornton,M. Winters, and P. Zary for field assistance,
Douglas Mock for numerousandproductivediscussions,and
CharlesBrown and four anonymousreviewersfor constructive commentson the manuscript.
LITERATURE
CITED
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next? Oikos 76:613-620.
Amundsen, T., and T. Slagsvold. 1998. Hatching asynchrony
in Great Tits: a bet-hedging strategy? Ecology 79:295-304.
Anderson, D. J. 1990. Evolution of obligate siblicide in boobies. 1. A test of the insurance-egg hypothesis. American
Naturalist 135:334-350.
Barber, C. A., and R. M. Evans. 1995. Clutch-size manipulations in the Yellow-headed Blackbird: a test of the individual optimization hypothesis. Condor 97:352-360.
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Multiple Incentives For Parental Optimism and Brood Reduction in Blackbirds

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Multiple Incentives For Parental Optimism and Brood Reduction in Blackbirds

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Multiple Incentives for Parental Optimism and Brood Reduction in Blackbirds

Author(s): Scott Forbes, Richard Grosshans and Barb Glassey

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Ecology, 83(9), 2002, pp. 2529-2541