1858 Animal Behaviour, 36, 6
winged choughs, Corcorax melanorhamphos. Ibis, 120,
178-197.
Vehrencamp, S. L. 1977. Relative fecundity and parental
effort in communally nesting anis, Crotophaga sulciros-
tris. Science, N.Y., 197, 403 405.
Woolfenden, G. E. & Fitzpatrick, J. W. 1984. The Florida
Scrub Jay. Demography o f a Cooperative-breeding Bird.
Princeton, New Jersey: Princeton University Press.
Yom-Tov, Y. 1975. Synchronization of breeding and
intraspecific interference in the carrion crow. Auk, 92,
778-785.
(Received 22 January 1988; revised 27 April 1988;
MS. number: sc-423)
Do Female Blackbirds, Turdus merula, Decode Song
in the Same Way as Males?
Most studies of the decoding of bird song have used
males only. Several authors (for example Becker
1982) have pointed out that this is unsatisfactory,
not only because song may convey information to
both males and females, but also because the two
sexes differ in behaviour and may sometimes
occupy slightly different habitats. Differences in
habitat use may result in males and females
experiencing different degrees of song degradation.
If the difference is marked, sex differences in song
decoding may evolve. As an example, the range of
variation within which certain song features elicit a
response may differ in males and females, i.e. the
two sexes may have different song variation toler-
ances (Shiovitz & Lemon 1980). We report here on
experiments testing this possibility in blackbirds.
Blackbird song consists of a long-ranging motif
followed by a higher-pitched and short-ranging
twitter (Dabelsteen 1984). Both the motif and the
twitter song convey information to territorial males
and receptive females (Dabelsteen & Pedersen
1985, in press; Dabelsteen 1988). However, there is
a marked sexual difference in receiving conditions
of forest-living blackbirds. Males often hear song
in relatively open conditions, for example when
patrolling territorial borders or when singing,
whereas females usually hide in vegetation at or
near ground level. A change of certain song
features (for example frequency level) up to or
beyond their natural limits of variation is not
tolerated by males (Dabelsteen & Pedersen 1985).
Because of the inferior receiving conditions the
song variation tolerances could be wider in females
than in males. We tested this by presenting females
with motif songs whose frequencies had been
altered (cf. Dabelsteen & Pedersen 1985).
A recording of blackbird song lasting 2 min was
modified on a computer. All twitter parts were
removed, and the remaining motif songs were used
to construct six test tapes each lasting 3 min. The
first control tape had unaltered motif songs. The
remaining five tapes had motif songs with all
frequencies multiplied by one of the following
values: 0.75, 0-875, 1.1, 1.2, or 1.4. Three of the
modifications changed the frequency level up to
(0.75, 1.2) or beyond (1.4) its natural limits of
variation making it abnormal for motif songs.
Multiplication by 0.875 and 1.1 changed the level
within its natural limits of variation making it code
for decreased and increased arousal, respectively
(Dabelsteen & Pedersen 1985).
Ten females were made receptive to song stimu-
lation by long day conditions (16:8 light: dark)
and subcutaneous injections of 1 mg oestradiol-
valerat, and then tested in an anechoic chamber.
Each bird was tested in two experimental series
each consisting of six double tests. In a double test
the bird was confronted twice with one of the six
types of song. Two hours passed between the two
presentations in a double test. Each test consisted
of three 3-min playback sessions with two 3-min
pauses inserted between the sessions, plus a 3-min
post-playback period. For further details of pro-
cedure see Dabelsteen & Pedersen (in press).
The responses of the females were recorded on
an NEC PVC-9507 videotape recorder. Three
response types (R-I-3) and eight behaviours be-
longing to them were quantified: R-I: sexual
behaviours, duration of copulation displays; R-2:
nest-building behaviours, number of pecks and
scrape-turnings in straw deposited on the nest-site
platform, ae-sounds (a female vocalization) and
transport of straw between the floor, perches and
platform; R-3: other behaviours, number of orien-
tating head movements, jumps, and pecks at the
bottom of the cage (see description in Dabelsteen
1988). A behaviour was regarded as released by the
song if its duration or frequency exceeded that of
the spontaneous behaviour during the 3-min pre-
test period. If not, the score was set at zero. The
response types were released when one (R- 1) or two
(R-2 and R-3) of their behaviours were released.
Any response type released in a test scored one
point, i.e. the maximum score of a double test was
six. Within each experimental series the durations
and frequencies obtained or points scored for each
bird in each double test (N = 20) were expressed as
percentages of the highest value possible. A Fried-
man two-way ANOVA was used to test the overall
heterogeneity between the values obtained for the
six types of song.
Both the duration of the copulation display and
the total reproductive response, as expressed by
points scored, varied significantly between the six
 Animal Behaviour, 36, 6
I00
o
50 a 1 b
"6 0.750.875 I.I I-2 1.4 0.750.875 I'1 1.2 1.4
I00
c)
50
0-75 0.875 I-I 1"2 I-4 0-75 0-875 I-2 1.4
Motif type
Figure 1. Responses of female blackbirds to playback of
unmodified motifs (closed histograms) and motifs
whose frequencies were multiplied by 0.75, 0.875, 1.1,
1.2, or 1.4 (open histograms). (a) Means for the
duration of sexual behaviour. (b) Overall means for the
frequencies of nest-building behaviours. (c) Overall
means for the other bebaviours. (d) Means for points
scored (see text for explanation). SESare indicated for
sexual behaviour and for points.
types of song (P<0.01 and <0.001, respectively;
Fig. 1). A pairwise comparison using the Wilcoxon
critical range method (Colquhoun 1971) showed
that unaltered motif songs and motif songs modi-
fied by the factor 1.1 both scored more points than
the 0.75 modification (P < 0-05), and that the 1.1
modification released more copulation displays
than the 0.75 modification (P<0.01) and the 1.4
modification (P < 0.05). Similar trends were found
for the other behaviour patterns, and orientating
head movements and jumps varied significantly
between the test songs (P<0.01 and <0.05, re-
spectively). Orientating head movements were
released less frequently by the 0-75 modification
than by unaltered motif songs (P<0.01) and the
modifications 1.1 (P < 0.05) and 1-4 (P < 0-05), and
the unaltered motif songs released more jumps than
the 0-75 modification ( P < 0.05).
The variation in females' responsiveness to
modifications 0"75, 0.875 and 1-1 was expected
from the way the coding of the frequency para-
meters was changed, and it is consistent with the
results of the previous experiments with males
(Dabelsteen & Pedersen 1985). However, unlike
those of the females, the males' responses were also
reliably reduced by modifications 1-2 and i -4. As all
modifications resulted in frequencies well within
the hearing range of passerines, such a difference in
1859
the responsiveness of the two sexes is difficult to
explain simply by the laboratory conditions of the
female experiments. We did not test the male and
female results directly against each other; however,
the difference in responsiveness may be real. High-
pitched motif songs may resemble twitters in
frequency, modulations, etc. (e.g. Dabelsteen
1984). Therefore, one possibility is that relatively
strong female responses were obtained with songs
whose frequencies were elevated by factors 1.2 and
1.4 because the females somehow perceived the
high-pitched motif songs as powerful twitters. A
second interpretation is that the decoding mecha-
nism of female blackbirds is less critical than that of
males as regards the upper part of the spectrum.
The two interpretations are not mutually exclusive,
and they both make sense as proximate explana-
tions since twitters in another experiment released
strong female responses (Dabelsteen & Pedersen, in
press).
We suggest habitat-induced variations in sound
degradation as one of the forces ultimately respon-
sible for the sexual variation in the decoding
mechanism of blackbirds. Females should be less
discriminatory than males when using high song
frequencies as identifying cues, because frequencies
higher than 3-4 kHz may degrade more when
passing through dense vegetation to females than
when passing the more open communication chan-
nel to other males (Wiley & Richards 1982). Other
forces, e.g. sexual selection, may tend to make
females more discriminating towards song struc-
ture. This may also apply to the blackbird, mani-
festing itself in the song variation tolerances for
song features other than the frequency level.
We thank M. Calford, O. N. Larsen and H. Lind
for their comments on the manuscript. The study
was made possible by a Niels Bohr fellowship given
by the Royal Danish Academy of Sciences and
Letters and the Tuborg Foundation, and by a
fellowship given by the Carlsberg Foundation, and
by the Danish Nature Research Council (grant no.
11-4153).
TORBEN DABELSTEEN*
SIMON BOEL PEDERSENt
* Institute o f Population Biology,
Copenhagen University,
Universitetsparken 15,
DK-2100 Copenhagen O, Denmark.
t The Acoustics Laboratory,
Technical University,
DK-2000 Lyngby, Denmark.
References
Becker, P. H. 1982. The coding of species-specificcharac-
teristics in bird sounds. In: Acoustic Communication in
1860 Animal Behaviour, 36, 6
Birds. Vol. 1 (Ed. by D. E. Kroodsma & E. H. Miller),
pp. 131-181. New York: Academic Press.
Colquhoun, D. 1971. Lectures on Biostatistics. Oxford:
Clarendon Press.
Dabelsteen, T. 1984. An analysis of the full song of the
blackbird Turdus merula with respect to message
coding and adaptations for acoustic communication.
Ornis Scand., 15, 227-239.
Dabelsteen, T. 1988. The meaning of the full song of the
blackbird Turdus merula to untreated and estradiol-
treated females. Ornis Seand., 19, 7-16.
Dabelsteen, T. & Pedersen, S. B. 1985. Correspondence
between messages in the full song of the blackbird
Turdus merula and meanings to territorial males, as
inferred from responses to computerized modifications
of natural song. Z. Tierpsychol., 69, 149-165.
Dabelsteen, T. & Pedersen, S. B. In press. Song parts
adapted to function both at long and short ranges may
communicate information about the species to female
blackbirds Turdus merula. Ornis Scand.
Shiovitz, K. A. & Lemon, R. E. 1980. Species identifica-
tion of song by the indigo bunting as determined by
responses to computer-generated sounds. Behaviour,
74, 167-199.
Wiley, R. H. & Richards, D. G. 1982. Adaptations for
acoustic communication in birds: sound transmission
and signal detection. In: Acoustic Communication in
Birds Vol. 1 (Ed. by D. E. Kroodsma & E. H. Miller),
pp. 131-181. New York: Academic Press.
(Received 20 January 1988; revised 27 April 1988;
MS. number: sc-422)