Anita. Behav.

, 1990, 39, 1173-1178

Nest defence as shareable paternal care in red-winged blackbirds

PATRICK J. W E A T H E R H E A D
Department of Biology, Carleton University, Ottawa, Ontario KI S 5B6, Canada

Abstract. One of the costs to female birds of mating polygynously is having to share the parental care
provided by their mate. Nest defence, however, is one form of paternal care that females mated to the same
male should be able to share. In this study there was no difference in the defence of primary and secondary
nests by male red-winged blackbirds, Agelaiusphoeniceus. Nests that were provisioned by males were not
defended more aggressively than nests that were not provisioned. Nests that were successful had been
defended more aggressively by males than those that failed, indicating that male nest defence is valuable to
females. These results suggest that male nest defence is entirely shareable, in contrast to a recent study that
showed males preferentially defended nests of primary females (Knight & Temple, Condor, 1988, 90, 193-
200). Although the study design used here more realistically reflects natural nest defence (males had to
defend one nest at a time rather than having to choose which of two nests to defend), it does not explain
why males preferentially defended certain nests when forced to make a choice but not when defending nests
one at a time.

As with many behaviour patterns, mate choice
involves costs and benefits. We expect natural selec-
tion to favour females who choose mates in a way
that maximizes their net benefits, given whatever
constraints are in effect. Thus, attempts to under-
stand why female birds choose to mate polygyn-
ously, even though opportunities for monogamy
are available, have focused on both the costs and
benefits of polygyny to those females (e.g. Orians
1969; Weatherhead & Robertson 1979; Lightbody
& Weatherhead 1988) and on the constraints that
affect their choice (e.g. Alatalo et al. 1981). In the
case of polygynous passerines, two of the potential
costs to females arise from having to share the
food on their mate's territory and whatever par-
ental assistance he might provide. Male parental
care, such as feeding nestlings, is not easily shared
by females, since when a male is feeding at one nest,
he cannot be feeding at another. By contrast, nest
defence by males has generally been regarded as
readily shareable (Orians 1980; Patterson et al.
1980) because a male can defend all his nests
simultaneously when they are threatened by a
single predator. Therefore, sharing male nest
defence should not involve a cost to polygynous
mating. Recently, Knight & Temple (1988) con-
ducted an experiment with red-winged blackbirds,
Agelaius phoeniceus, that challenged this assump-
tion. Here, I present data from red-winged black-
birds that support the assumption that male nest
defence can be shared by females without any cost.
Knight & Temple (1988) presented model preda-
tors simultaneously at primary and secondary
(defined by order of nest initiation) nests within
males' territories. This experimental protocol
required males to choose which nest to defend.
Although both nests were defended, males were sig-
nificantly more aggressive in their defence of the
nests of primary females. The implication of this
result is that females that choose to nest in a terri-
tory where another female is already present do so
at the cost of receiving less aggressive defence of
their nests from the male. Although it is conceivable
that males may occasionally face the real world
dilemma simulated by Knight & Temple's (1988)
experiments (i.e. having predators threatening two
of their nests simultaneously), a more usual scen-
ario is likely to involve a single predator threaten-
ing each nest separately. Thus, I asked whether
males defend primary nests more aggressively than
secondary nests when defending them one at a time.
In addition, I tested whether males defend nests at
which they feed young more aggressively than those
at which they do not.
METHODS
Data were collected from May to July 1988 as part
of a broader study of the reproductive biology of
red-winged blackbirds being conducted at the
Queen's University Biological Station in eastern
Ontario. Study sites consisted of seven marshes

0003-3472/90/0601173 + 06 $03.00/0 9 1990 The Association for the Study of Animal Behaviour
1173
1174 Animal Behaviour, 39, 6
located within 10 km of the Biological Station. All
breeding males were individually colour-banded
and their territories were mapped. Marshes were
thoroughly searched for nests every second day in
order to locate all nests. Searching began in the first
week of May when the first nests were initiated.
Once a nest was located, its position was mapped
and an identification number was written on a piece
of flagging tape tied near the nest. Thereafter, each
nest was visited every second day until they failed or
the young fledged.
Nest defence trials were conducted at the begin-
ning of each visit to each nest. As observers moved
through the marshes looking for new nests, they
conducted trials at each nest that had been dis-
covered previously. To maximize the success at
detecting new nests, observers purposely used dif-
ferent routes through each marsh on sequential
visits. For the purpose of this study, the effect of
varying the route taken through each marsh was to
make the order in which nests were visited more
random.
Nest defence was quantified using a modified
version of a method I have used previously
(Weatherhead 1979, 1982, 1989), in which the
birds' response to the observer is used as a measure
of their nest defence. Once the observer had
approached to within 1 m of the nest, the male's
response was observed for the next 2 min and the
maximum response on a scale from 0 to 7 was
scored. I used identical categories to those used by
Eckert & Weatherhead (1987; page 37), where 0
was assigned to males that left their territories at the
beginning of a trial, 7 to males that physically
struck the observer, and intermediate values to
males whose reponses involved approaches and
vocalizations of various intensities. The rationale
for a 2-min trial (as opposed to one of longer
duration) is that for parents to be effective defending
their nests against real predators they must respond
as soon as a predator poses a threat. The use of
simple response categories rather than detailed
quantification of specific behaviour avoided bias
between observers. Data were collected by three
assistants, all of whom worked in all of the study
areas. Since the data were collected before I decided
to use them for the purposes of this paper, there
should be no bias with regard to the analyses herein.
Male red-winged blackbirds also provide some
food to nestlings. However, not all males feed
nestlings, and those that do only provide food at
some nests (Patterson 1979; Muldal et al. 1986).
Preferential feeding at some nests is a clear case of
differential allocation of male parental investment.
I wished to determine whether male nest defence
was correlated with feeding (i.e. more aggressive
defence of provisioned nests) as one might predict
from Knight & Temple's (1988) results. All nests in
which nestlings reached 5 or 6 days of age were
observed for 1 h on either day 5 or 6. Those males
that do feed nestlings usually do so in the latter part
of the nestling period (Muldal et al. 1986). Obser-
vations were made by one observer from outside
the male's territory. Male feeding was quantified as
the number of trips made to a nest. Subsequent
observations using videotape recordings at nests
have confirmed that male visits to nests nearly
always involved provisioning the young (personal
observations).
RESULTS
The total sample for this study consisted of 786
trials conducted at 132 nests of 52 males. Analysis
of nest defence through the nesting cycle for success-
ful (defined as fledging at least one young) and
unsuccessful nests produced two results (Fig. 1).
First, consistent with most nest-defence studies (see
Montgomerie & Weatherhead 1988), male response
became more aggressive through the nesting cycle
(successful: r 2 = 0"042, P < 0'001; unsuccessful: r 2 =
0'016; P = 0'005). Second, nests that were ultimately
successful were defended more vigorously than
those that failed (ANCOVA: slopes not signifi-
cantly different, F1.TS2 = 1"689, P=0.194; adjusted
means, F 1,783 5"99, P = 0"015).
=
A comparison of nest defence of primary and
secondary nests reveals no apparent difference (Fig.
2). I used two approaches to assess these data statis-
tically. First, I divided the nesting cycle into six
intervals beginning with the first day of incubation.
Within each interval, I used a Wilcoxon matched-
pairs signed-ranks test to compare responses of
males who had been tested at both a primary and
secondary nest within that interval. No difference
in response was found in any interval (Table I). To
take fuller advantage of the complete data set, I
compared the response at primary versus all other
nests through the nesting cycle using analysis of
covariance. The slopes of the two regression
equations were the same (F1,624 = 0.370, P = 0.544)
and no difference was found in the adjusted mean
responses (F1,625 = 0.409, P = 0.523).
 Weatherhead: Sharing nest defence by blackbirds 1175

0 I I I I I I I I I I I I
2 4 6 8 I0 12 14 16 18 20 22 24
Day
Figure 1. Mean (--FSE)male nest-defence score at nests that were eventually successful ( ) and at those that were
unsuccessful ( - - - ) relative to the day of the nesting cycle (day 0 is the day the first egg was laid).

a)
4
o
ul

g
a)

2 4 6 8 I0 12 14 16 18 20 22 24
Day

Figure 2. Mean (__+sE) male nest-defence score at primary ( ) and later ( - - - ) nests. Primary nests were those of the
first female to nest in a male's territory.
1176 Animal Behaviour, 39, 6
Table |. Wilcoxon matched-pairs signed-ranks tests comparing male
nest defence at primary and secondary nests at different stages of the
nesting cycle

Sample P
Stage* size Z (two-tailed)

Early incubation (day 0-3) 18 - 1.726 0.084

Mid-incubation (day 4-6) 19 - 0.852 0.394
Late incubation (day 7-10) 22 - 1.507 0.132
Early nestling (day 0-3) 20 -0.070 0.944
Mid-nestling(day 4-6) 14 -0.089 0.929
Late nestling (day 7-10) 12 - 1.540 0.124

*In the incubation period day 0 is the first day of incubation. In the
nestlingperiod day 0 is the day the first egg hatched.

In the preceding analysis it is possible that the
lack of difference between the defence of primary
and secondary nests is a consequence of how the
data were collected. Because male red-winged
blackbirds defend relatively small territories, nest
densities can be quite high (Weatherhead &
Robertson 1977). Consequently, a male's response
at one nest may not be independent of his response
at the previous nest tested. However, primary and
secondary nests were initiated on average 4.6 days
apart. Therefore, paired comparisons for a given
nesting stage (Table I) involve trials conducted on
different days in most cases. The time between trials
was even greater for the comparison of primary
versus all other nests.
A second potential confounding factor that
could arise as a result of the methods I have used
concerns Knight & Temple's (1986) hypothesis that
birds respond more strongly to a perceived nest
threat on successive exposures if the 'predator' has
left the nest unharmed on previous appearances. If
such positive reinforcement occurred, it should re-
sult in secondary nests being defended more aggres-
sively than primary nests since males defending
secondary nests have had more exposure to the nest
threat. That was clearly not the case. However, to
assess this quantitatively, I analysed whether males'
nest-defence scores increased as a function of either
the number of previous visits to the nest at which
they were being tested or the number of times they
had been tested previously at any nest. I restricted
the analysis to secondary nests so that each male
would be considered only once and to provide sub-
stantial variation in the number of previous trials.
For a given nest, at a given stage, the number of
previous trials at that nest depended on when the
nest had been found relative to clutch initiation.
Five nesting stages were selected for analysis. Prior
visits, either at the nest being tested or at any nest
belonging to a male, did not affect nest-defence
scores (Table II). This result is consistent with a
similar analysis of song sparrow, Melospiza
melodia, nest defence (Weatherhead 1989).
Feeding observations were made at 78 nests, only
15 of which received any provisioning by males
during the time they were observed. Using male
nest-defence scores on the day of the feeding obser-
vation or the day before, I found no difference in
the defence of provisioned and unprovisioned
nests (Mann-Whitney U-test, P=0.438). For the
15 males that provisioned nestlings, I found no cor-
relation between the number of feeding visits and
nest defence score (Spearman rank correlation, rs =
0"185, P=0.438). Therefore, there is no evidence
that males defend nestlings more aggressively if
they feed them.
DISCUSSION
In this study, males defended nests more aggress-
ively through the nesting cycle as predicted by
parental investment theory (Weatherhead 1979;
Andersson et al. 1980). Also as predicted if nest
defence is a form of parental investment, nests that
were defended more aggressively were more likely
to be successful. However, I found no evidence that
males defended nests of their primary females more
aggressively than nests of females settling later.
Also, there was no evidence that males defended
nestlings that they fed more than nestlings they did
 Weatherhead: Sharing nest defence by blackbirds
Table II. Linear regression analysis of defence score relative to the number of previous
trials at a nest and relative to the total number of previous trials in a male's territory*
Nest Previous trials at the nest:~
stager
(day) r2 P Range
6-7 0.178 0.072 0-2
10-11 0.019 0.537 I-4
14-15 0.011 0.683 2-6
18-19 0-000 0.940 2-8
22-23 0'043 0.458 3-11
*Onlytrials at secondary nests are included.
tDay 0 is the day the first egg was laid.
:~Rangeis the least and greatest number of previous trials within the sample; N is the
number of males tested.
not feed. Thus, the results of this study lead to the
conclusion that male nest defence is shareable.
Before considering the implications of my results, I
consider possible reasons why my results may have
differed from those of Knight & Temple (1988).
Knight & Temple (1988) quantified nest defence
in much more detail than I did. However, method-
ological differences are unlikely to account for
differences in results. The value of nest defence to
females must be the protection of her offspring.
Both my approach and that of Knight & Temple
(1988) to quantifying nest defence showed that
nests that were successful had been defended more
aggressively. Thus, both methods assessed nest
defence in a way that reflected its apparent purpose.
One reason why Knight & Temple (1988) could
have found that males preferentially defended pri-
mary nests is because they were more valuable to
the males. By testing primary and secondary nest
simultaneously, primary nests would always be
closer to fledging at the time of the test. Because a
nest closer to fledging is of higher reproductive
value to the parent, the nest should be defended
more vigorously (Montgomerie & Weatherhead
1988). However, Knight & Temple (1988) found
that the difference in the intensity of nest defence at
primary and secondary nests was independent of
the time interval separating initiation of those
nests. Therefore, their results are apparently not
attributable to the males varying their response
according to nest value based on nestling age.
My results seem potentially internally inconsis-
tent in that nest defence (a form of parental invest-
ment) was not correlated with nestling feeding
(another form of parental investment). However,
1177
Previous trials in territory~
N r2 P Range N
19 0.050 0.359 2-14 19
22 0.128 0-102 4-14 22
18 0'006 0.761 7-16 18
15 0.115 0.217 10-21 15
15 0.110 0.227 11-24 15
male feeding is so limited in this population that it is
probably a poor index of parental investment.
The probable explanation for the difference
between this study and that of Knight & Temple
(1988) is experimental design. Knight & Temple
forced males to choose which nest (primary or
secondary) to defend. I allowed males to defend
each nest at whatever level they wanted, uncon-
strained by conflicting demands. As long as my
assumption is correct that males usually have to
defend nests one at a time, then nest defence should
be shareable by females within a harem. Even if my
interpretation is correct regarding the sharing of
nest defence, Knight & Temple's (1988) results
remain interesting by showing that males value
nests of primary females over nests of secondary
females, even if not enough to defend them differ-
ently when defending them one at a time. Knight &
Temple (1988) suggested that males may value
nests of primary females more if those females are
of higher quality or if their nests have a higher
chance of success. Why such differences should
only cause differences in nest defence when nests
are threatened simultaneously but not sequentially
remains unexplained and therefore an interesting
problem for further research.
Finally, my results have important implications
for polygynous mate choice. Female red-winged
blackbirds that mate polygynously can expect a
male to defend their nests with as much vigour as he
defends other nests in his territory, and the vigour
of that defence should not decrease with the
number of females nesting in the territory. There-
fore, nest defence is a substantiallydifferent form of
parental care than feeding behaviour and should
1178 Animal Behaviour, 39, 6
present no hindrance to females choosing to mate
polygynously. In fact, as a shareable male quality,
nest defence becomes a male trait u p o n which mate
choice could be based. Since nests that are defended
most vigorously are most likely to be successful,
males likely to be good defenders should be pre-
ferred by females. Also, because nest defence is cor-
related with epaulette size (Eckert & Weatherhead
1987), females have some potential to recognize
males likely to be vigorous nest defenders at the
time mate choice occurs. Consistent with these
expectations, Yasukawa et al. (1987) and Knight &
Temple (1988) found positive correlations between
harem size and nest defence in red-winged black-
birds. Such a correlation would not be expected if
nest defence was not shareable.
ACKNOWLEDGMENTS
I am grateful to Drew Hoysak, Kit M u m a and
Kevin Dufour for assisting with data collection,
Drew Hoysak for helping with data analysis and
Bob Montgomerie for commenting on the manu-
script. Financial support was provided by the
Natural Sciences and Engineering Research
Council of Canada.
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(Received 1 June 1989; initial acceptance
19 August 1989;final acceptance 27 September 1989;
MS. number: A5575)