Behav Ecol Sociobiol (1981) 8:183-188
Behavioral Ecology
Repertoire Sharing and Song-Post Distance
in Nightingales (Luscinia megarhynchos B.)
Henrike Hultsch and D i e t m a r T o d t
Abteilung fiir Verhaltensbiologie, Institut ffir Allgemeine Zoologie, Freie Universit/it Berlin, Haderslebener Strasse 9, D-1000 Berlin 41
Received April 18, 1980 / Accepted December 1, t980
Summary. The relationship was studied between song-
post distances and the extent o f vocal repertoire shar-
ing in 34 territorial nightingales settling in six h o m o -
geneously structured habitats. Repertoires were com-
pared on the basis o f shared song types and distances
were measured between nocturnal song posts o f first-
order and higher-order neighbours. O u r results
showed that male nightingales shared fewer song
types with very close and more distant neighbours
than with neighbours at intermediate distances. This
distribution is explained by the interaction between
repelling and attracting components, which depends
on the distance between song posts.
Introduction
Bird song m a y vary intraspecifically. In m a n y species
the degree o f diversity o f song patterns is closely relat-
ed to distance between songsters. This diversity is
often more p r o m i n e n t between populations (local dia-
lects) than between males o f the same population
(Marler and T a m u r a 1962; Thielcke 1969; Notte-
b o h m 1969; Gfittinger 1977; Jenkins 1977; Baker and
Mewaldt 1978).
Within populations similarity o f vocal repertoires
generally decreases with increasing distance between
territorial neighbours (Bertram 1970; T h o m p s o n
1970; Harris and L e m o n 1972; H o w a r d 1974;
K r o o d s m a 1974). However, blackbirds (Turdus meru-
la L.) and nightingales (Luscinia megarhynchos B.)
were f o u n d to follow this rule only above a certain
distance range, but apparently not when settling
closer than 100 m (Schmidt 1971 ; W o l f f g r a m m 1979;
Hultsch, paper presented at Int. Ethol. Conf., Van-
couver 1979). Recently a similar finding was reported
for the cirl bunting (Emberiza cirlus; Kreutzer 1979).
The study presented here examines the relationship
and Sociobiology
9 Springer-Verlag 1981
between sharing song repertoires and the distances
between song posts in nightingale habitats c o m p o s e d
o f contiguous territories.
Nightingales are particularly g o o d for this kind
o f study for the following reasons: (1) repertoires
can readily be compared, since song types o f males
(units of comparison) can easily be identified and
categorised and the same types show little variation
within and between individuals; (2) each male has
a large repertoire o f song types (about), which pro-
vides a wide range in the degree o f repertoire sharing
between individuals (Todt 1971); (3) the birds often
occur in long strips o f habitat where territories are
almost linearly arranged. This kind o f a r r a n g e m e n t
permits an easy assessment o f relationships between
song-post distances.
Methods
Sites. An analysis was made of vocal repertoires and the distance
between song posts for 34 nightingales (Luscinia megarhynchos
B.) in six habitats showing homogeneous habitat structure and
approximately linear arrangement of territories. The study areas
were: St. Miehel/Haute Provence (two sites; 0.53 km with seven
and 0.47 km with six birds), Selestat/Alsace (0.49 km with seven
birds) and Wannsee/Berlin (three sites; 0.52 km with five; 0.54 km
with five and 0.37 km with four birds). Habitat lengths were mea-
sured between nocturnal song posts of marginal males.
Recording. Recordings were made April-May 1978 and 1979 within
2 weeks after the first seasonal nightingale song in a habitat. We
used a Uher 4200 Stereo IC tape recorder and Sennheiser micro-
phones MKE 802 N. Only territorial males were chosen for analy-
sis. We identified individuals by their location and by the fine
structure of their songs. Territories were estimated on the basis
of the diurnal song-post distribution (Fig. 1). Analysis of the rela-
tionship between song repertoire sharing and distance between
males was done on the nocturnal song only, for the following
reasons: (1) In contrast to the daytime, males do not change their
song posts during the night, and the same post is used on several
successive nights. This facilitates ascertainment of spatial distances
between individuals. (2) During the night, the birds produce long
continuous sequences of songs, and song types are repeated less
0340-5443/81/0008/0183/$01.20
184

HABITAT OF A NIGHTINGALE POPULATION : 7 BIRDS A i

(ST, M I C H E L ; HTE, PROVENCE ; A P R I L '7~) A [55 125 24(1 305 385 530
i

B 70 185 250 33(~ 4 7 ;

C 115 180 260 405
__ ~ ~ - - ~ ~-~--...,, ,--
,.-J-" ,.---7 ~ ~^ D 155 145!290
~ - ~ j ~ , ~ " " < ~k.~\ ,--, ...... _

. . . . " " N - ') ,~., ~ J " ~ .,-.. " ~ DISTANCECm) I ~ . _ ~ _ _

A B . v . -~=._ , . __
E

Fig. 1. Song post distribution in a nightingale habitat. Distances (see: inset table) are measured between nocturnal song posts (circles
with stars) of the neighbours A, B, C, D, E, F, G. Diurnal song posts (filled circles) surround the nocturnal posts. Line at bottom:
habitat boundaries (shrub area with meadows beyond)

the frequency distribution of the number of element types by which

20C
10~
100 300
NUMBER OF CONSECUTIVE S O N G S
Fig. 2. Cumulative frequency distribution of occurrence of new
nightingale song types. A: nocturnal singing; B: diurnal singing;
this comprises successive bouts separated by long pauses (arrows).
Data recorded from one bird
frequently than during the day. Thus, the entire repertoire of a
male can be recorded in about 45 rain. We considered the repertoire
to be sampled if we recorded no new song type in a sequence
of 50 switches of songs (Fig. 2). In control tests during five succes-
sive nights no additional song types were detected. Tapes were
analysed by a Nicolet UA 500 Spectrum Analyser and a Kay
Electric 7029 A Sonagraph (frequency range 8-8000 Hz; wide band
setting). The classification of songs into types was based on a
catalogue of element types (Hultsch 1980a).
song types differed was bimodal; song types differed either by
0 3 element types or by many more than 3 element types. During
counter singing, mutual matching was performed using songs that
were classified by us as belonging to the same types.
To calculate the amount of song types shared by pairs of
males, we used the following coefficient of repertoire sharing (RS)
in a dyad (D):
Z
RSD= (:g+ ~ - Z
Z=number of song types shared by both individuals of the dyad
X = repertoire size of one member
Y=repertoire size of the other member of that dyad.
The values for RSD can range between 0 and 1. When RSD=O,
500 there is no sharing of songs, and when RSD=I, all song types
in both repertoires are shared. RSD is identical for both males
in the comparison of pairs, even if individuals have different sized
repertoires.
To standardize measurements of RSD between habitats we
expressed them as a percentage of maximum value within a habitat.
Each standardized RSD value was plotted against the distance mea ~
sured between nocturnal song posts for each dyad (Fig. 6). Ob-
served frequencies (f~) of dyads in the cells of a contingency table
were compared with expected frequencies (fe) calculated on the
basis of a null hypothesis which assumed that dyads were distribut-
ed by chance (Table 1). Data were analysed using the x2-test (signif~
icance level: P=0.01%).
Results

Repertoire Sharing. We characterized songs as belonging to the Song-Post Distribution

same song type when they differed in not more than three of
approximately ten element types in the first two sections of the Distances were measured between nocturnal song
song (sections e and/~; Fig. 3). We adopted this criterion because p o s t s o f 34 n i g h t i n g a l e s i n six h a b i t a t s ( F i g . 4). D i s -
 185

I- . . . . 1S . . . . -I ~kHz , ~ , , ~ , ~ , ~ ~ , ,

| @

,/,t2_
-4 ......
JII
[
It
~I 'I vl vl -v- I~t
n

~llB ~N~amAM I ~ ILIMITR~ G O~ pI~I IRODK, R. J,

! 1 I

h
' li I~
tk
I 1 t

~,~ .,,2 .o.~.,. ~ ~, ,~t~tT~,~, co ~,.~ ,~oo~.. ~.

i r I I ~ I l 1 t t

| L
ill ,l|ilJ|,
lllllllfl~

~1 Iq ~ ~1 I'~ y

Fig. 3. Sonagraphic display of two song types (23 ; 86) recorded in the habitat shown in Fig. 1 and shared by nightingales A, C and D

tances between song posts of first-order neighbours Repertoire Sharing

(adjacent neighbours) ranged between 40 and 200 m
and formed two frequency peaks, one at 120 m and
another at 60 m. The latter was not found in Berlin
habitats. Thus, distances of about 60 m between
neighbours seemed to be correlated with a high den-
sity of males not occurring in Berlin.
We could not detect cues showing that the distri-
bution pattern of neighbours was generated by differ-
ences in the structure of their habitat. In five out
of six sites habitat structure appeared to be homoge-
neous not only in the occupied area but also on either
side of the habitat. This factor did not constrain males
from settling at distances greater than those observed.
We, and presumably the nightingales, too, were
able to hear a male singing at one end of the study
site while listening at the other end. Consequently,
all males within a site could be regarded as neigh-
bours. To discriminate between neighbours, males
separated by one or more territories are called 'higher-
order neighbours'.
The repertoire sizes of males ranged between 160 and
280 song types. Individual repertoires remained con-
stant during the study period.
For comparison of repertoires we calculated the
amount of repertoire sharing between particular pairs
of males. Figure 5 gives an example f o r RSo values
calculated for males settling in the habitat shown in
Fig. 1. Further evaluation provided evidence of a dis-
tinct relationship between repertoire sharing and
song-post distance of neighbours: males separated by
less than 90 m or more than 360 m shared relatively
fewer song types than those at intermediate distances.
This was true for all Haute Provence, Alsace and
Berlin sites (Fig. 6). According to repertoire sharing
we were able to characterize three distance zones for
all of these sites (Table 1).
In contrast to Haute Provence and Alsace habi-
tats, all Berlin habitats were unoccupied in zone I.
In zones II and III RSD data were distributed in
 186

I.-
< REPERTOIRE DISTANCE BETWEEN NOCTURNALSONG POSTS~m
I- SHARING (RSD
~ 1,5-
E X XX X XX (A) >75%
0
0T-
~O(~X
XX
X
~I~KX
X ( HP1)
(HP2) z
| |174174174174174
LU
D.
"o 1=

~C) 1,0
9 O0 (Sl)

(s2)
50-75%
|
9 "ee" " 9 (,%) | |
I.-
el
Z
l:1 < 5 0 % '~
I ' ' ' ' I
100 200
DISTANCE (m)

Z
5- @|174
UJ s
o 6
i,i 50-75%
ri-
LL
m @@
0~
@ |
100 200 <50% x,! |
DISTANCE (m)
Fig. 4. Distribution of distances measured between the nocturnal |
song posts of adjacent neighbours. Top: distances plotted against
density of males within three Berlin sites (Bb B2, B3), two Haute
Provence sites (HP1 =habitat shown in Fig. 1, HPa) and one Alsace Fig. 6. Relationship between dyadic repertoire sharing (standard-
site (A). Bottom: same distances superimposed ized RSD) and nocturnal song-post distances of nightingales within
a habitat (first-order and higher-order neighbours). Circle diame-
ters and numbers indicate frequency of dyads; circles with dots
on top (upper distribution) belong to Haute Provence/Alsace habi-
Number of song types shared (A&B~ A &Cj ....... ) tats (1 dot: HP1; 2 dots: HPz; 3 dots: A). Circles with dots
on bottom (lower distribution) belong to Berlin habitats (1 dot:
' Totalof
St.IA~B~C...) B1; 2 dots: B2; 3 dots: B3). x): values<25% were not observed

A 29 81 86 34. 42 33 (248)

B .06 ~ 31 76 69 47 32 (253) the same way in all habitats. This suggests that the
<~ 127)
m- peak of song sharing was related to absolute distance
9.-. .22 .07 ~ ;'J rather than to distance measured in terms of numbers
~< C (I001 (32) ~ 6g 55 54. 4.0 (196)
of territories between neighbours.
"~ D ,20 .'18 J9 N 28 80 87 [24.s
"o (911 (82) (861
"6
,~ .08 .17 .15 .07
Discussion
E 4.0 62 (212)
::3 (36) (77) (68) (32)
The distribution of distances measured between noc-
I .09 .10 34 .19 .09
70 (260) turnal song posts of first-order neighbours formed
cn~ F (411 (/-S) (64) (861 {411
r'.-" a clear peak at 120 m. It occurred independently of
.07 .06 .10 .21 .15 .16 habitat diversities in all sites studied so far. We as-
G (255)
(32) (27) (45) (96) (68) (73) sume that settling within 120 m may result f r o m a
Fig. 5. Repertoire sharing between seven neighbouring nightingales set of factors a m o n g which song plays an important
(habitat HP1; see Figs. 1, 4). Above major diagonal: number of role, namely by attracting and repelling conspecifics.
song types shared by two individuals; below major diagonal: dyad- A balance between these factors might support both
ic repertoire sharing (RSD); in brackets: sharing expressed as per- efficiency of communication as well as establishment
centage of the maximum R S D value found in the population (here:
dyad AC); right column: total of song types in the repertoire of territories large enough for sufficient food supply
of each bird (A, B, C, etc.) and reproductive success.

Table 1. Repertoire sharing (RSD) in three distance zones (I, II, III); columns HP/A refer to Haute Provence and Alsace, columns
B to Berlin habitats
Repertoire Distancezones
sharing (RS~)
I (<90 m)
HP/A B HP/A
fo L L L L
>50% 2 6 - -
< 50% 8 4 - -
fo=observed frequencies of dyad distributions (see Fig. 6);fe=expected frequencies of dyads; for null hypothesis: random distribution
of dyad frequencies.Significanceof differencebetweenfo and f,:P < 0.01% for both HP/A and B distributions (ZZHp/A= 34, dfz ; )~2B= 21.4,
dfl)
Repelling Components
During nocturnal singing, nightingales settling close
to each other perform most of their songs alternately,
thus vocal patterns do not overlap in time (Hultsch
1980b). However, when they match each other they
give the response as soon as they detect which particu-
lar song type a neighbour has started to perform.
This results in temporal overlap of songs with mutual
masking of central song sections. The more song types
neighbours share, the higher the probability is that
they will match each other during counter singing.
The decrease in repertoire sharing over short distances
( < 90 m) might arise as a result of two strategies:
males might avoid settling too close to neighbours
with which they share many songs (repulsion hypothe-
sis). On the other hand, very close neighbours may
avoid performing song types that are shared (conceal-
ing hypothesis).
A study of blackbirds supports the first hypothesis
(Todt 1981): they were found to avoid singing on
song posts where playback songs frequently matched
the birds' own songs, with high amplitude, and with
a short delay (close to 1 s). Amplitude as well as
the timing of matching responses vary with distance
between songsters. Thus neighbours sharing many
song types couId prevent loud masking of their central
song sections (/?-sections, constituted by non-repeti-
tive units) by increasing the distance between their
song posts. At present we do not know whether the
assumed repelling effects are achieved via vocal inter-
actions (e.g. countersinging) or just by estimating the
amount of repertoire sharing auditorily.
We have no evidence to support the concealing
hypothesis. To examine it we plan to test the singing
behaviour of 'potential concealers' via removal of
their zone I neighbours, followed by playback of new
song types.
187
II (>90 m; <360 m) III (>360 m)
B HP/A B
L fo L fo L fo f~
32 22 19 15 1 7 0 4
3 13 1 5 10 4 6 2
Attracting Componem's
Our results have shown that males occupied territories
relatively close to one another, even though there
were large regions of apparently suitable habitat on
either side of the occupied areas. Particularly, males
sharing much of their repertoires (RSD > 50%) settled
closer than 360 m (Fig. 6). It is doubtful whether this
can be completely explained by the decrease in repel-
ling effects. In many species song learning and site
tenacy may combine to produce a tendency for neigh-
bours to share songs. The adaptive implications of
song sharing within populations were first discussed
by Nottebohm (1972) and recently reviewed by Trois-
man ( 1 9 7 8 ) a n d Trainer (1980). Male nightingales
learn much of their repertoire in early life, and visual
contact with a tutor is important (Todt et al. 1979).
This suggests that young birds predominantly learn
the song types of their father or adjacent neighbours.
Nightingales are found to return to breed very close
to the place where they were born (Hilprecht 1965).
This may result from both a preference for a specific
site (natal or former breeding site) and a preference
to settle within earshot of conspecifics, in which a
recruit detects similarities in repertoire. At present
we lack sufficient data to decide which of these prefer-
ences is most important.
As a test, we plan to broadcast local song types,
thereby labelling suitable sites at the time when males
return from their winter quarters and before they
select their territories. If returning birds make their
choice because of similarities in repertoire rather than
a preference in site it should be possible to attract
them, thereby dislocating the population in a predict-
ed direction.
Close Neighbours
In addition to the peak of song-post distances between
first-order neighbours at 120 m, there was also a
188
smaller peak at 60 m (Fig. 4). P r e s u m a b l y the shorter
distances were a result of some birds being able, due
t o low repertoire sharing, to settle b e t w e e n males
120 m apart. The i n t r u d i n g birds m i g h t be y o u n g e r
nightingales that have i m m i g r a t e d f r o m other p o p u l a -
tions.
This c o u l d be the case because: (1) A l t h o u g h
nightingales do n o t develop clear-cut local dialects,
n e i g h b o u r i n g p o p u l a t i o n s differ in their vocal reper-
toires (Todt, u n p u b l i s h e d data). (2) B a n d i n g data pro-
vided evidence that first-year males are m o r e likely
to m o v e into n e i g h b o u r i n g h a b i t a t s t h a n older ones
(22% of r e c a p t u r e d y o u n g individuals). (3) Older
males are the first to r e t u r n f r o m their winter q u a r t e r s
(Hilprecht 1966). I f these older birds space their terri-
tories a b o u t 120 m a p a r t (via c o m p o n e n t s discussed
above), recruits s h o w i n g low repertoire s h a r i n g m i g h t
have a chance to settle in prospective h a b i t a t s n o t
only o n the periphery.
It should be t a k e n into a c c o u n t that o u r results
are based o n a n analysis o f static relationships. The
relevance o f social i n t e r a c t i o n s has n o t yet been exam-
ined. T h e results o u g h t to be c o m p l e t e d by investigat-
ing the d y n a m i c s of vocal c o m m u n i c a t i o n d u r i n g
e s t a b l i s h m e n t of territories, p a r t i c u l a r l y by s t u d y i n g
the roles that shared a n d n o n - s h a r e d song types play
in territorial interactions.
Acknowledgements. We thank J.R. Krebs for reading the manu-
script. His comments and suggestions were greatly appreciated.
This work was supported by the Deutsche Forschungsgemeinschaft
(To 13/9).
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