Sexual Selection and Body Size in Male Red-Winged Blackbirds

Sexual Selection and Body Size in Male Red-Winged Blackbirds
Author(s): William A. Searcy

Source: Evolution, Vol. 33, No. 2 (Jun., 1979), pp. 649-661
Published by: Society for the Study of Evolution
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Evolution, 33(2), 1979, pp. 649-661
SEXUAL SELECTION AND BODY SIZE IN
MALE RED-WINGED BLACKBIRDS
WILLIAM A. SEARCY'
Department of Zoology, University of Washington, Seattle, Washington 98195
Received March 16, 1978. Revised September 18, 1978
In most species of birds adult males are
larger than adult females. The two major
hypotheses proposed to explain this pat-
tern of sexual size dimorphism are appro-
priately termed the sexual selection and
competition avoidance hypotheses. The
sexual selection hypothesis states that
large size gives males an advantage in in-
trasexual competition for females or in
epigamic displays, activities not engaged
in by females. Males will thus be larger
than females, and the difference should be
greater among polygynous and promiscu-
ous species where there is greater vari-
ability in male reproductive success and
thus stronger sexual selection (Darwin,
1871; Huxley, 1938; Fisher, 1958). The
sexual selection hypothesis, then, explains
the relationship between pronounced sex-
ual dimorphism in size and nonmonoga-
mous mating systems in birds (Darwin,
1871; Selander, 1958, 1972; Amadon,
1959; Orians, 1961), a relationship partic-
ularly evident among the Icteridae (Selan-
der, 1958).
The competition avoidance hypothesis
states that sexual size dimorphism is fa-
vored because it reduces competition be-
tween males and females of the same
species (Selander and Giller, 1963; Selan-
der, 1966, 1972; Johnston and Selander,
1973). If size affects the range of resources
used (e.g., larger individuals eat larger
food items), then those males differing
most in size from females would overlap
in resource utilization with the fewest in-
dividuals and thus have the most re-
sources available. This hypothesis may be
especially applicable to monogamous, ter-
1
Present address: Rockefeller University Field
Research Center, Tyrrel Road, Millbrook, New
York 12545.
ritorial species in which sexual selection
is weak and the male and female in a pair
may often be each other's worst competi-
tors. The competition avoidance hypoth-
esis does not explain the trend toward in-
creased dimorphism in nonmonogamous
species, which implies that sexual selec-
tion must be at least one of the causes of
dimorphism in families, such as the Icter-
idae, exhibiting this trend. The two hy-
potheses are not mutually exclusive, and
competition avoidance could explain the
small degree of size dimorphism in mo-
nogamous Icterids while sexual selection
accounts for the increased dimorphism in
nonmonogamous species.
Adult male red-winged blackbirds (Age-
laius phoeniceus) are about 20% larger
than females in wing length. Male red-
wings should be strongly affected by sex-
ual selection since there is a great range
of variation in mating success among male
redwings. Female redwings in Washing-
ton populations choose mates largely on
the basis of the quality of their territories
(Searcy, 1979) so the outcome of inter-
male competition for territories should
have a strong effect on the direction of
sexual selection. The ability to win ag-
gressive encounters should be of advan-
tage in competition for a territory, and
large males should have an advantage in
aggressive encounters. Thus large males
should presumably have an advantage in
competition for territories, which provides
a clear mechanism by which sexual selec-
tion could favor large size in male red-
wings.
If increased size dimorphism in non-
monogamous Icterids is caused because
large size in males is favored by sexual
selection, what selective forces act to limit
male size? Selander (1965) thought that in
649
650 WILLIAM A. SEARCY
the promiscuous great-tailed grackle
(Quiscalus mexicanus), large size made
males less efficient at foraging than fe-
males and thus exposed them to a greater
risk of starvation in winter. He also
thought that the conspicuous size of males
made them more vulnerable to predation.
Using estimates of sex ratios in roosts, Se-
lander was able to show that males suffer
greater mortality than females in winter.
Thus it seems probable that size in male
great-tailed grackles is set by a balance
between sexual selection favoring in-
creased size in the breeding season and
selection for survival ability favoring de-
creased size. Note, however, that it has
not been shown that large males actually
do better reproductively than small males,
nor has it been shown that small males
survive with greater probability than large
though the latter is a reasonable inference
from available data.
The present study is an attempt to dis-
cover the selective forces determining
male body size in a population of red-
winged blackbirds. I started by investi-
gating whether Selander's hypothesis
about size in male grackles could be ap-
plied to these redwings. Redwings possess
the advantage for such a study that male
reproductive success is proportional to
harem size (Orians, 1972; Holm, 1973)
which can be measured relatively easily.
Another advantage of redwings is the site
fidelity of territorial males (Nero, 1956;
Searcy, 1979), which allows reliable esti-
mates of survivorship.
METHODS
Fieldwork for this study was done at
the Turnbull National Wildlife Refuge in
eastern Washington State. The refuge is
located in a region of ponderosa pine (Pi-
nus ponderosa) parkland interspersed with
shallow lakes. Red-winged blackbirds
nest in emergent vegetation (mainly cat-
tails [Typha latifolia] and bulrushes [Scir-
pus sp.]) along the margins of the lakes.
Fieldwork was conducted late March
through June in 1974, 1975, and 1976, and
May 6-9, 1977.
I estimated body size by wing length,
generally considered the best measure of
size in birds (Hamilton, 1961; James,
1970). Wing length correlates closely with
fat-free weight at least in some birds (Con-
nell et al., 1961), and, unlike weight, wing
length does not fluctuate greatly either
daily or seasonally. Wing length was mea-
sured in 1974 as the chord of the wing,
but in 1975 and 1976 the wing was flat-
tened when measured because this pro-
cedure gave a more repeatable value.
Males were captured using decoy traps
(Smith, 1972) and mist nets. Captured
males were measured, banded with
unique combinations of colored leg bands,
and released. Weights were measured
with a 100 g spring scale. Weights of ter-
ritorial males declined by 0.09 g per day
in 1974 (Pearson's r = -.327, P < .05)
and by 0.11 gperdayin 1975 (r = -.371,
P < .05), but there was no significant de-
cline in 1976. I will refer to the actual
weight at the time of measurement as
"weight" and to the weight of a bird cor-
rected back to the day the first individual
was captured, using the assumption that
each individual lost weight at the average
rate, as the "corrected weight."
Time budgets of territorial males were
estimated once in 1976 (March 22-April
5) and at three different stages of the 1975
season (March 25-April 11, May 7-May
24, and June 4-June 8). Estimates were
made by visiting territories at 15 min in-
tervals through eight h periods in the mid-
dle of the day and noting the owners' ac-
tivities.
Harem sizes were calculated as the
number of females needed to account for
the nesting activity on a territory. It was
assumed that a female would not renest
sooner than seven days after her first
clutch was destroyed or 20 days (including
the time spent feeding fledglings) after her
last young fledged if her first nest was suc-
cessful. Nests started late in the season
were attributed to a female who had nest-
ed earlier on the same territory whenever
a long enough period intervened between
the disappearance of all eggs or young
from the first nest and the appearance of
eggs in the second. Thus my estimate of
SELECTION IN RED-WINGED BLACKBIRDS 65 1
TABLE 1. Correlations of harem size with wing
length.
r,
= Spearman rank correlation coefficient.
N
r,
P
1974 42 -.210 >.05
1975 34 .075 >.05
1976 41 .319 <.05
the number of females settling on a terri-
tory is a minimal one because there are
undoubtedly cases where the owner of the
first nest failed to renest on the territory
and a new female moved in to start the
second nest. However, I am not able to
correct for these occurences because the
females were not marked.
RESULTS
Tests of Selander's hypothesis.-One
way to test Selander's hypothesis is to
compare survival rates of males and fe-
males. If large size carries a price in de-
creased survival, then male redwings
should have lower survival rates than fe-
males; however, this is a weak test since
males and females differ in other charac-
teristics besides size which could affect
survival. Haigh (1969) sexed 58 redwing
nestlings found dead in the nest: 39 were
males and 19 were females. If the primary
sex ratio (at hatching) was 1 to 1, then
these data indicate that males are more
likely to die as nestlings than are females
(X2
= 6.90, P <
.01). However, Haigh's
best estimate of the primary sex ratio,
based on the sexes of fledglings in broods
where none of the nestlings died, was 44
males to 33 females, or 1.33 to 1. If this
estimate of the primary sex ratio is used,
then Haigh's data do not indicate that
male nestlings are significantly more likely
to die
(X2
= 2.45, P > .10). Fankhauser
(1971), using banding records of 400 fe-
males and 953 males, found nearly iden-
tical mortality for the two sexes as adults:
57.9% annual mortality for males and
56.7% for females.
Stronger tests of Selander's hypothesis
can be made by comparing survival rates
and mating success of large and small
males. Selander's hypothesis predicts that
large males should have larger harems
than small males but lower survival rates.
The first prediction was tested by calcu-
lating Spearman rank correlation coeffi-
cients between male size and harem size
for each of the three seasons (Table 1).
Only in 1976 was a significant positive
correlation found.
If small males have a higher survival
rate than large males, then the mean wing
length of the males surviving from one
year to the next should be lower than the
mean wing length of the nonsurvivors.
This prediction was tested by dividing the
territorial males which were measured in
a year into those that were seen the next
year and those that were not and compar-
ing the mean wing lengths of the two
groups. The 17 males that survived from
1974 to 1975 had an average wing length
of 127.9 mm while the 25 nonsurvivors
had an average of 128.8 mm. The differ-
ence is not significant (t = 1.02, P > .10
by a one-tailed test). For the 1976-1977
interval, the mean wing length of the 20
survivors (131.0) was again smaller than
the mean wing length of the 21 nonsur-
vivors (131.6) but not significantly so (t =
.709, P > .20 by a one-tailed test). How-
ever, for the 1975-1976 interval, the 25
survivors had a larger mean wing length
(130.8) than the 10 nonsurvivors (130.2)
though again the difference was not sig-
nificant (t = .625, P > .50 by a two-
tailed test).
Similar data were gathered by D. M.
Johnson and associates (pers. comm.).
These authors, working at a winter roost
in Houston, Texas, took wing measure-
ments of live redwings captured in traps
and of redwings found dead at the roost.
The average wing length of 24 live adult
males was 127.9 versus an average of
126.7 for 279 dead adult males. The dif-
ference is in the opposite direction from
that predicted by Selander's hypothesis
but is not significant (P
>
.20). The av-
erage wing length of 60 live first year
males was 122.2 versus an average of
122.0 for 154 dead first year males. Again,
the difference is in the opposite direction
from that predicted but is not significant
(P
> .20).
To summarize, the prediction that mat-
ing success would increase with size was
upheld only one year in three. No support
was found for the prediction that adult
females would have higher survival than
adult males, nor for the prediction that
small males would have higher survival
than large males. The evidence on differ-
ential mortality among the sexes as nest-
lings was somewhat ambiguous.
A model of male size in redwings.-I
have argued that males are larger than fe-
males in redwings because of sexual selec-
tion, and yet males do not seem to pay a
price for large size in increased mortality.
What, then, acts to limit size in male red-
wings? To answer this question, it is con-
venient first to construct a model of opti-
mal size in the absence of sexual selection.
If sexual selection were not affecting size,
optimal size would be that size where sur-
vival and parental investment are maxi-
mized. Survival should be influenced both
by predation and energy balance, but I
will neglect predation because the direc-
tion of the effect of predation is not
known, that is, it is not known whether
predation selects for larger or for smaller
size in redwings. Since there is evidence
that large male redwings are not more
likely to die than small ones, predation
must not select strongly against large size
anyway. The amount of parental invest-
ment which can be made also depends on
energy balance.
Energy balance is most favorable at the
size at which an animal has the most
energy left over after its maintenance re-
quirements have been met to expend on
parental care, energy storage, etc. Both
the efficiency of foraging and the cost of
maintenance will change with body size.
Let us define the profit of foraging (in
units of energy/time) as a function f of
size (s) and the cost of maintenance (also
in energy/time) as a function g of size.
Then the optimal size in terms of energy
balance is that size where f(s)
-
g(s) is
maximal. This is not quite correct because
a small animal can do more with a given
amount of energy than can a large one.
Let us suppose that the most crucial activ-
ity of an animal has a cost, again depen-
dent on size, of h(s), in units of energy
per unit activity. Then the optimal size
is that size where
f(s)
( ) is maximal.
h
(s)
The units of this expression are
energy/time - energy/time unit activity
energy/unit activity time
An imaginary example may clarify this
formulation. Suppose that it is of great
importance to the reproductive success of
a male bird to be able vigorously to chase
rivals from his territory. Suppose further
that on average he must spend half his
daylight hours in territory defense, and
during the rest of the day he is able
to forage. Maintenance metabolism costs
him y kcal/h, and while foraging he makes
a profit of x kcal/h. If chases cost him
z kcal/m, then for a 24 h period with 12 h
of daylight, he will be able (without de-
pleting energy reserves) to afford
(6x
-
24y) kcal/day _ (6x
- 24y)
z kcal/meter z
meters/day of chases.
Since I am using maintenance metabo-
lism to mean the rate of energy expendi-
ture in the absence of any activity or en-
ergy storage, g(s) in a thermal neutral
environment is equivalent to basal meta-
bolic rate (BMR). The shape of the BMR
versus size curve is well known for many
taxa; for passerines, BMR in kcal/h is 4.78
W .726 where W is weight
in
kg (Aschoff
and Pohl, 1970; see Fig. 1). At tempera-
tures below thermal neutrality, mainte-
nance metabolism is equivalent to stan-
dard metabolic rate (SMR), which has a
different relationship to size at every tem-
perature. However, the general shape of
the SMR versus size curve at any temper-
ature below thermal neutrality should be
similar to the BMR curve, that is, the
SMR curve will rise as size increases but
with ever decreasing slope (Kendeigh,
1969).
SELECTION IN RED-WINGED BLACKBIRDS 653
9 (s
z
small SIZE large
FIG. 1. Hypothetical curves relating size to en-
ergy intake and maintenance energy costs. f(s) = en-
ergy profit per time foraging for individuals of size
s. g(s) = energy cost of maintenance for individuals
of size s.
BMR is a very general expression for
h(s) since the costs of many activities are
approximately constant multiples of BMR
(Hemmingsen, 1960; Berger and Hart,
1972; King, 1974). For example, Berger
and Hart (1972) found that flight in birds
cost 45.5 W
73
kcal/h, which is equivalent
to 9.3 x BMR (King, 1974). Hemmingsen
(1960) found that the maximal 'output of
energy/time in activity is approximately
20 x BMR for animals of a wide range of
f(s)
-
g(s) .
sizes. Thus
i()
is a general expres-
sion for the amount of activity which can
be afforded by an animal over a given
period of time with a given fraction of
that period devoted to foraging. Maxi-
mizing this expression also maximizes the
length of time an individual can survive
at temperatures within the thermal neu-
tral zone on the profit of a given period
of foraging.
The shape of f(s) is unknown, but we
can make certain assumptions about it. It
is logical to assume that for a given type
of animal feeding on a given type of prey
(e.g., a blackbird feeding on emerging in-
sects), f(s) will go to zero below some size
(e.g., as the blackbird becomes too small
X- f(s)
CT)g(s)
z
, g(s)
g(s)
Sopt
small SIZE large
FIG. 2. Optimum size where selection maximizes
the difference between energy intake per time and
maintenance energy costs per time scaled to main-
tenance energy costs. f(s)
=
energy profit per time
foraging for individuals of size s and g(s) = main-
tenance energy costs for individuals of size s.
S0,t
is
the size at which f(s)Ig(s) - g(s)lg(s) is maximal. Sex-
ual selection shifts actual size above
S,,t,
thus de-
creasing f(s)/g(s) - g(s)lg(s).
to handle and swallow insects). Similarly,
f(s) must decrease above a certain size
(e.g., as the blackbird becomes too large
to walk among cattails). The exact shape
of the curve between these two sizes has
little effect on the argument, as long as the
curve is unimodal. I will assume the curve
has the shape graphed in Figure 1.
Probable shapes of typical f(s)lg(s) and
g(s)lg(s) curves are graphed in Figure 2.
The optimal size energetically is that size
where the difference between these two
curves is greatest, that is, at the peak of
f(s)lg(s). I will call this energetically opti-
mal size So,,t.
If sexual selection has moved the size of
male redwings above
S,pt,
then the larger
an individual is, the smaller
fs)
-
g(s)
is
g(s) '
for that individual. Thus counterselection
against large size could be due to large
males being more susceptible to starva-
tion, but as mortality does not increase
with size in male redwings, starvation
seems to be unimportant as a selection
pressure on size. Large individuals should
be able to perform less activity on the
energy gained in a certain period of for-
aging. Large males should thus perform
less parental care than small males, but
male redwings in Washington populations
contribute little parental care, and their
contribution has little effect on the suc-
cess of their broods (Searcy, 1979). Large
males should also not be able to perform
as much territory defense behavior as
small males. This suggests the hypothesis
that size is limited in male redwings be-
cause males are able to perform less and
less territorial defense behavior as their
size increases. This disadvantage should
increase exponentially as the amount of
territorial behavior goes to zero and so
should eventually cancel out the advan-
tage of large males in aggressive encoun-
ters. Further, males will be able to per-
form less epigamic display behavior as
size increases, which may also lower pair-
ing success. Weatherhead and Robertson
(1977) have found that the vigor and
duration of epigamic display does affect
male redwing pairing success, at least
in Ontario populations, and they believe
that the amount of epigamic display per-
formed is energy limited. Thus because
of increasing energy limitation of territory
defense and epigamic behavior as size
increases, sexual selection may not con-
tinually favor larger body size but, in-
stead, may favor a stable body size some-
what above
SPt.
Evidence for the hypothesis.-The hy-
pothesis that large males suffer an ener-
getic disadvantage which tends to cancel
out their advantage in aggressive encoun-
ters during competition for territory leads
to the prediction that mating success in
male redwings will not increase with size.
As we have already seen, this prediction
was borne out in two out of three years of
this study. Only in 1976 was there a pos-
itive correlation between harem size and
male wing length (Table 1).
The hypothesis assumes that some
males, at least, have difficulty getting
enough energy to perform sufficient epi-
gamic or territorial defense behavior. The
impressive amount of energy invested by
males in territory defense and display be-
havior makes this assumption credible.
Male redwings in eastern Washington set
up territories in early February and de-
fend them two to three months before any
females start to nest. During much of Feb-
ruary and March, snow may cover the
ground and lakes may be frozen so that
presumably many food sources are un-
available. Many males continue to defend
territories into July, thus defending for a
total of five months. Daily time invest-
ment rises from a brief period at morning
and evening to an average of more than
70% of daylight hours in May. Ridding
territories of intruding males may involve
many minutes of energy-costly aerial
chase. At peak display times, males dis-
play hundreds of times an hour, and
though individual displays may cost little,
the cost added over many months must be
substantial. How much of this display be-
havior is directed towards warding off ri-
vals and how much towards attracting fe-
males is not known; most displays
probably serve both functions simulta-
neously. For this reason, I will lump ter-
ritory defense and epigamic behavior to-
gether as "territorial behavior."
The fact, mentioned earlier, that in two
years the weights of territorial males de-
clined in April and early May is evidence
that males are under energy stress. The
decline was 0.09 g per day in 1974 (P <
.05) and 0. 1 1 g per day in 1975 (P < .05).
Males in 1976 were estimated to have lost
0.06 g per day, but the loss rate was not
significantly greater than 0 (r =-.188,
P > .05).
The assumption that males curtail their
territorial behavior for want of sufficient
energy implies that if males had more en-
ergy available, they would increase terri-
torial behavior. This prediction was tested
by an experiment in which males were
provided food on their territories and their
display rates compared to those of a con-
trol group of unfed males. Experimental
males were fed a mixture of cracked corn
and commercial dog food on elevated
trays on their territories. Control and ex-
perimental males held territories on the
same lake, and were chosen to yield a re-
TABLE 2. Results of feeding experiment. Averages
for the measures of territorial behavior are given for
the experimental group (N = 7) and control group
(N = 7). Feeding was begun between observation
rounds 1 and 2. * indicates a significant
difference
between the experimental and control averages at the
.05 level according to a one-tailed Mann-Whitney U
test.
Song Flight Minutes Song
number displays on rate
Round 1
experimentals 33.4 3.7 20.0 1.67
controls 34.4 3.4 19.5 1.75
Round 2
experimentals 45.3 2.1 18.8 2.37
controls 40.0 3.7 20.0 2.00
Round 3
experimentals 38.6* 3.3* 20.0* 1.93
controls 23.3 1.0 14.4 1.69
Round 4
experimentals 35.0 3.9* 19.8 1.79
controls 22.0 1.3 14.9 1.33
Round 5
experimentals 33.4* 3.0* 19.4 1.71*
controls 21.9 0.6 17.6 1.13
Round 6
experimentals 35.0 3.9* 19.9 1.76
controls 23.4 1.0 17.2 1.20
peating series of an experimental male, a
male which was neither fed nor observed
(a blank male), a control male, and a
blank male, down the lake shore. Displays
were recorded for 20 min periods during
the first two h after sunrise. One round of
observations was made on the seven ex-
perimental and seven control males before
feeding began, and five rounds during
feeding.
The results of the experiment are pre-
sented in Table 2. The numbers of songs
and flight displays are those given during
a 20 min period, the number of minutes
are those out of the 20 min observation
period that the male spent on his territory,
and song rate is the number of songs given
per minute on the territory.
There was very little difference in dis-
play rates between the experimentals and
controls before feeding began. Feeding
was begun between rounds 1 and 2, but
there was still no significant difference in
round 2, perhaps because many of the ex-
perimentals seemed not to have discov-
ered their food trays. In rounds 3, 4, 5,
and 6, the averages for the experimentals
are higher than for the controls for each
measure of territorial behavior, and half
the differences are significant. There is
good evidence that fed males spent more
time on their territories (round 3), and that
they sang more often when they were on
territory (round 5). The most consistently
significant difference between the experi-
mentals and controls was in flight dis-
plays, and it is perhaps significant that
flight displays are energetically more ex-
pensive than songs.
A possible objection to this experiment
is that the positive results occurred not
because the food provided energy but be-
cause the food attracted more intruding
males who had to be chased away or more
females to be courted. I calculated the av-
erage numbers of males and females
chased during the observation periods in
each round of observation. The only sig-
nificant difference between the experi-
mental and control averages occurred in
the number of males chased in round 1,
before feeding began. Thus this objection
can be dismissed.
I have provided evidence that males are
under energy stress and that the amount
of energy available to males limits their
territorial activity. Is there evidence that
energy stress is greater for large males
than for small ones? Such evidence could
consist either of a demonstration that large
males use up their energy reserves more
quickly than small males, or of a demon-
stration that large males limit their terri-
torial behavior more than small males. I
investigated both possibilities.
I determined whether large males had
proportionately smaller energy reserves
than small males at the time of weighing
by determining whether large males were
further below the expected weights for
their sizes. Expected weights were found
using data from J. E. Mosimann and F.
C. James (unpub. ms.) on geographic
variation in wing length and weight in
Florida populations of redwings. A power
function was fitted to the average weights
and wing lengths for 12 samples from dif-
ferent localities (Fig. 3). The equation
for the curve is w = 0.000381 (L2503),
where w is weight in grams and L is wing
length in mm (r = 0.919, P < .01). Al-
though Florida redwings are substantially
smaller than Washington redwings, the
curve fits the average corrected weights
and wing lengths from my Washington
samples impressively well. If we assume
that the redwing populations studied by
Mosimann and James did not differ in any
systematic way in the average percentage
of weight made up by stored fat, then this
curve can be used to predict, for given
wing lengths, weights with a constant per-
centage of the weight as stored fat. We
can thus use the percentage deviation of
the weight of individual males from the
weight predicted by this curve as a mea-
sure of the relative sizes of their energy
reserves.
I divided the territorial males into those
above average and below average in wing
lengths and compared the percentage de-
viations of their actual weights from their
expected weights. For all three years,
large males were further below their ex-
pected weights than were small males. In
1974, 22 below average size males were
on the average 3.1% above their expected
weights and 20 above average size males
were 4.6% below their expected weights,
the difference being significant at the .001
level by a one-tailed Mann-Whitney U
test. In 1975, 19 below average size males
averaged 1.7% below their expected
weights versus 6.9% below for 16 above
average size males (P < .01). In 1976, 20
below average size males averaged 4.6%
below expected versus 8.3% below for 20
above average size males (P < .01). Thus
there is evidence that large males have
proportionately smaller energy reserves
than small males after two to three months
of holding territory.
I tested the prediction that large males
would perform less territorial behavior
using two different measures: display rates
and percentage of the day spent guarding
80
75
70
E 65
w 60
55
50
45
105 110 115 120 125 130
WING LENGTH (mm)
FIG. 3. Relationship of weight and wing length
in male redwings. x = mean from one of the Florida
localities (data from Mosimann and James, unpub.).
o
=
mean from one of the three years of this study.
Curve is fitted to Flordia data; with equation, w =
.000381(L2503), where w = weight in grams and L =
wing length in millimeters.
the territory. Table 3 shows the correla-
tions between wing length and three mea-
sures of display rates defined earlier: song
number, flight displays, and song rate.
The correlations tend to be negative in
1974 and 1975 (10 of 12 are negative), and
two are significant at the .01 level. For the
display rates observed in 1976, all three
correlations are positive but not signifi-
cant. Table 3 also shows the correlations
between display rates and condition of the
males, measured as percentage deviation
of their corrected weights from their ex-
pected weights. For 1974 and 1975, most
of the correlations are positive (10 of 12)
and in four cases they are significant at
the .05 level. The correlations for 1976 are
negative though not significant.
Table 4 shows the correlations between
wing length and estimates of the percent-
age of the day males spent guarding their
territories. No estimates were made in
1974. Estimates were made during three
TABLE 3. Correlations of display rates with body size and condition. Condition is measured as percentage
deviation of the measured weightfrom that predicted by the curve in Fig. 3.
r,
= Spearman rank correlation
coefficient. a = probability from two-tailed test; all other probabilities are from one-tailed test.
Song number Flight displays Song rate Song number Flight displays Song rate
vs. vs. vs. vs vs. vs.
wing length wing length wing length condition condition condition
1974 rs
-.115 -.079 -.172 .133 .128 .156
5/8-5/29 N 42 42 42 42 42 42
P .235 .311 .138 .107 .116 .073
1975 rs -.559 -.012 -.626 .498 -.091 .509
3/25-4/11 N 20 20 20 20 20 20
P .006 .480 .002 .013 .704a .011
4/12-5/6 rs
-.183 .102 -.164 .380 .048 .373
N 34 34 34 34 34 34
P .151 .566a .177 .014 .394 .015
5/26-6/3 rs
-.144 .229 -.210 .215 -.073 .237
N 28 28 28 28 28 28
P .232 .242a .142 .136 .714a .118
1976 rs .223 .156 .223 -.097 -.285 -.097
3/22-4/8 N 31 31 31 31 31 31
P .228a .404a .228a .604a .122a .604a
periods in 1975, and for two of these pe-
riods there was a negative correlation be-
tween wing length and percentage of time
spent guarding the territory, but the cor-
relations are not significant. In 1976, the
single set of estimates correlated positively
but not significantly with wing length.
Table 4 also shows the correlations be-
tween the percentage of the day spent
guarding the territory and condition. In
1975 there was a positive correlation in
two of three cases, and one correlation is
significant. In 1976 there was a negative
but not significant correlation.
Thus there is evidence, albeit weak,
that small males performed more territo-
rial activity than did large males in 1974
and 1975, the years in which body size
failed to correlate with harem size. There
is also evidence to support the hypothesis
that the condition of males in terms of en-
ergy stores affects the level of territorial
activity. The fact that large males per-
formed as much territorial behavior in
1976 as did small males may explain why
body size correlated positively with harem
size that year as large males would have
lost their disadvantage in territorial activ-
ity while retaining their advantage in ag-
gressive encounters.
Why were large males able to perform
as much territorial behavior as small
males in 1976 and not in 1974 or 1975?
The obvious hypothesis is that environ-
mental conditions were somehow better in
1976 than in 1974 or 1975 so that either
energy intake was up or maintenance costs
were down, or both, and a sufficient sur-
plus of energy was left for territorial be-
havior even for large males. This is a dif-
ficult hypothesis to support because
energy balance depends on a complex
blend of weather conditions and food
availability, and when during the season
the crucial periods are is unknown.
Weather records are available but infor-
mation on food availability is not. Table
5 shows average monthly temperatures for
February through May and the number of
days in each month with more than a trace
of snow cover for the nearest weather sta-
tion, the Spokane Airport, about 20 miles
from the study sites. Snow cover probably
lessens food availability. 1975 was clearly
the coldest year and also had substantially
more days of snow cover than either 1974
or 1976. On the other hand, 1974 had no
more snow cover than 1976, and three of
four months were warmer in 1974 than in
1976. As noted earlier, average weight of
TABLE 4. Correlations of percentage of time spent
guarding the territory with body size and condition
(measured as percentage deviation of measured
weight from that predicted by the curve in Fig. 3).
r.
= Spearman rank correlation coefficient. a =
probability from two-tailed test; other probabilities
are from one-tailed test.
With wing With
length condition
1975
March 25-April 11
r,
-.279 .414
N 26 26
P .084 .018
May 7-May 24
r,
.122 -.226
N 33 33
P .498a .208a
June 4-June 8 rs -.216 .143
N 26 26
P .145 .243
1976
March 22-April 5 rs .189 -.222
N 29 29
P .328a .248a
territorial males declined significantly in
1974 and 1975 but not in 1976, but the
difference in rate of decline is not signifi-
cant (Analysis of Covariance, F = 0.393,
P > .05). The average weight of males
was actually lower in 1976 than either
1974 or 1975, but again the differences are
not significant (1974 vs. 1976, t =
0.27,
P > .05; 1975 vs. 1976, t = 1.06, P >
.05). There is no clear evidence, then, that
1976 had the best environmental condi-
tions of the three years.
Selection for survival ability. -Does se-
lection for survival ability have any effect
on size? I have already shown that sur-
vival does not exercise any directional se-
lection on size, at least in adult males, but
it is still possible that survival has a sta-
bilizing effect on size. To investigate this
possibility, I divided the redwings cap-
tured each year into categories of the one-
fourth with the shortest wing lengths, the
one-fourth with the longest wing lengths,
and the one-half in the middle, and cal-
culated the percentage survival of each
group (Table 6). In each year the middle
group had the highest survival, but even
if all three years are pooled and the small
TABLE 5. Weather data, collected by the National
Oceanic and Atmospheric Administration Station at
the Spokane Airport.
Number of
Average days with
temperature more than a
(degrees trace of
centigrade) snow cover
1974 February 1.9 2
March 3.6 0
April 8.0 0
May 10.1 0
1975 February -4.1 28
March 1.1 4
April 5.4 2
May 11.5 0
1976 February 0.1 2
March 1.7 0
April 7.3 0
May 12.5 0
and large groups combined, the difference
in survival of the middle group versus the
combined large and small group is not sig-
nificant (X2 = 2.18, P > .10). Obviously,
more data on survival versus wing length
are needed before it can be decided wheth-
er survival selection has a stabilizing effect
on size.
DISCUSSION
The view of body size in male redwings
which has been developed here is that
size, in the absence of sexual selection, is
that which optimizes an individual's en-
ergy balance. Sexual selection has caused
the evolution of a large average size be-
cause large males have an advantage in
aggressive encounters which leads to in-
creased success in inter-male competition
for territories and thus to increased mating
success. However, large males suffer a
disadvantage in competition for territories
in that they cannot, because of energy lim-
itation, perform as much territory defense
behavior as small males. Large males are
also unable to perform sufficient epigamic
behavior and thus may attract fewer fe-
males than smaller males with equally
good territories. Thus at some size, there
is no further selection for size increase,
and size is stabilized by the action of sex-
ual selection alone.
TABLE 6. Percentage survival of body size cate-
gories. Categories: the quarter of the males mea-
sured that year with the largest wing lengths, the
quarter with the smallest wing lengths, and the
half in the middle. N = number of males in the size
category.
Largest Median Smallest
Percentage returning 36% 48% 30%
in 1975 from 1974 N = 11 N = 21 N = 10
in 1976 from 1975 N = 9 N= 17 N=9
in 1977 from 1976 N
=
10 N =
21 N
=
10
Total 47% 59% 45%
N = 30 N = 59 N = 29
I have provided evidence that the
amount of territorial behavior performed
is energy limited and that large males are
more severely limited both in energy stores
and in the amount of territorial behavior
performed than are small males. Thus it
seems that large males do have a disad-
vantage which would tend to negate their
presumable advantage in aggressive en-
counters during competition for territo-
ries. However, it is not clear whether
in the long run the advantage and dis-
advantage exactly cancel each other so
that large males do not experience any net
sexual selection advantage. Large males
may experience a slight mating advan-
tage, perhaps only expressed in occasional
good years, which is balanced by a slight
survival disadvantage, perhaps experi-
enced in the nestling stage. Alternatively,
the observed mating advantage of large
males in 1976 might be balanced by a mat-
ing disadvantage in particularly bad years
while survival has no effect on size.
If there is stabilizing selection on size
due to medium-sized individuals surviv-
ing better than extreme-sized individuals,
this would be somewhat destructive to the
view of body size just presented. If my
model of optimal size is correct, large
males could possibly be at a disadvantage
in survival relative to medium-sized
males, but small males, if anything,
should be at an advantage. One way to
reconcile stabilizing selection for survival
with my model is to postulate differentf(s)
curves for winter and spring. This is ac-
tually possible since redwings feed largely
on insects in the spring and on grain in
the winter. Larger size might be advan-
tageous to males feeding on grain in win-
ter flocks because it might be possible for
large males to monopolize the best feeding
patches. Thus the optimal size energeti-
cally could be larger in winter than in
spring so that medium-sized males could
be above the optimum in spring but near
it in winter, when stabilizing selection
could occur. This is entirely speculative,
of course; there is not even firm evidence
that selection for survival has a stabilizing
effect on size.
One selection pressure which has not
been considered is predation. Predation
was ignored because it is not known
whether large or small size would be fa-
vored under predation pressure. Plausible
arguments can be made for both alterna-
tives. Large males may be more conspic-
uous to predators and may be more ex-
posed to predation because they must feed
longer both to grow and to sustain them-
selves. On the other hand, large size may
make males difficult for certain predators
to capture and handle. The evidence
shows that there is no strong directional
selection due to mortality so predation
cannot be exerting a strong directional
pressure on size unless it is nearly can-
celled out by some opposing pressure.
Although I started by discussing sexual
size dimorphism, I have largely neglected
female size. Females should have an f(s)
curve similar to that of males as the diets
of the two sexes are similar, at least in
the spring (Orians, 1979). The g(s) curve
should also be similar to that of males.
The strongest selection on females prob-
ably is to maximize the number of young
they can feed (Lack, 1954, 1966). Thus
females should have that size which maxi-
mizesf(s) ,
)
, where h'(s) measures the
h'(s)
energy cost of feeding an average off-
spring for a female of size s, assuming
that
this energy cost depends on the sizes of
the offspring at independence and that the
size of offspring is correlated with the size
of the mother. h'(s) may have a similar
shape to g(s) because a large fraction,
perhaps the majority, of the energy pro-
vided to the young is used by them
in maintenance metabolism rather than
growth (Ricklefs, 1974). Thus the size of
female redwings may be near the ener-
getically optimum size for both sexes.
Sexual selection selects for a larger size in
males thus creating sexual size dimorphism.
The hypothesis that energy limitation of
territory defense and/or epigamic behavior
limits male size can be applied to other
species. Males of nonmonogamous species
characteristically devote more time and
energy to display than males of monoga-
mous species (Selander, 1972), and thus
energy limitation of display is more likely
to affect nonmonogamous species. For
example, leking male grouse, for a period
of months in late winter-early spring,
spend at least several hours each morning
and evening in display, and some display
all night (Wiley, 1973). Besides display-
ing, males must expend energy on bound-
ary clashes and copulation attempts.
Kruijt et al. (1972) noted that a common
cause of loss of territory in black grouse
(Lyrurus tetrix) is apparent ill health of
the male, which might very likely be
brought on by energy depletion. Collias
and Collias (1967) found in the polygynous
village weaverbird (Textor cucullatus) that
males must build and display four nests
for every one accepted by a female, and
that building one nest requires 30 to 40
km of flight, obviously a great expenditure
of energy. It is possible that in these
species, as in redwings, large males ex-
perience a more severe energy limitation
of territory defense or epigamic behavior
than small males, and thus that sexual se-
lection may not continually favor larger
male size but instead may favor a stable
size somewhat above female size.
SUMMARY
Male red-winged blackbirds are about
20% larger than females. Selander (1965)
proposed that size in male Icterids is set
by a balance between sexual selection fa-
voring large size and survival selection fa-
voring small size. The trend towards
greater size dimorphism in nonmonoga-
mous Icterid species indicates that large
size in males has evolved because of sexual
selection. However, in a population of
redwings in Washington State, large
males had a higher pairing success than
small males only one year in three, and
there was no evidence of counterselection
against large size due to higher survival
of small males.
The hypothesis is proposed that coun-
terselection against large size is due to
large males having less energy left after
maintenance requirements are met to ex-
pend on territory defense and epigamic
behavior. Thus large males suffer an en-
ergy disadvantage in competition for
mates which tends to balance their pre-
sumable advantage in aggressive encoun-
ters. Evidence for the hypothesis is: (1)
males lose weight in the spring, indicating
they are stressed for energy; (2) males pro-
vided with food on their territories dis-
played more than a control group, indi-
cating that territorial behavior is energy
limited; (3) small males have larger energy
reserves proportional to their size than do
large males; and (4) large males perform
less territorial behavior than small males.
Thus size in male redwings may be sta-
bilized by the action of sexual selection
alone.
ACKNOWLEDGMENTS
I am very grateful to Gordon Orians for
his help in planning this study and for his
advice on preparation of the manuscript.
I am also grateful to Sievert Rohwer,
Thomas Schoener, W. James Erckmann,
Steven Fretwell, Daniel Johnson, and two
anonymous reviewers for helpful criti-
cisms of the manuscript. Frances James
and Daniel Johnson kindly allowed me to
cite unpublished data. Margaret Searcy
was of immense help with fieldwork and
with preparation of the figures. I thank
Jon M. Malcolm, Donald White, and the
staff of the Turnbull National Wildlife
Refuge for their cooperation with the
study. Financial support was provided by
the National Science Foundation, grant
BMS75-14937.
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