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ISSN 0096 3925, Moscow University Biological Sciences Bulletin, 2010, Vol. 65, No. 2, pp. 78–83. © Allerton Press, Inc., 2010.
Original Russian Text © O.A. Astakhova, I.R. Beme, 2010, published in Vestnik Moskovskogo Universiteta. Biologiya, 2010, No. 2, pp. 40–45.
The Typological Organization and Microgeographical Variation
of Chaffinch (Fringilla coelebs L.) Songs in the Population
of the Curonian Spit
O. A. Astakhova and I. R. Beme
Vertebrate Zoology Department, Biological Faculty, Moscow State University, Moscow, 119991 Russia
e mail: chaffinch@bk.ru
Received December 25, 2007
Abstract—The song repertoire of many sparrow species (Passeriformes) has not one but several song types.
In the investigated chaffinch population (Fringilla coelebs L.), we revealed 22 types of songs, of which four
turned out to be rare (they were found in 1–2 points), and five song types were combined (they consisted of
phrases of different song types). The average size of the chaffinch repertoire in the Curonian Spit was repre
sented by 2–3 song types (max of 6 song types and min of 1 song type). The variation in song types considered
by us can speak for genetic heterogeneity.
Key words: chaffinch (Fringilla coelebs L.), song, song types, variation in song types.
DOI: 10.3103/S0096392510020082
the biostation: the Morskoi settlement, Rybachii set
tlement, Lesnoi settlement; the distance between each
of them averaged 10 km (Fig. 1). The parcelling
approach was applied to the territorial distribution of
individuals [9], and record points were mapped.
Singing chaffinch males (N = 102) were registered
in the period from April 15 to May 10, 2005. A Pana
sonic RQ SX95F tape recorder and Philips SBC
ME570 condenser microphone were used to record
songs. The repertoire of chaffinch males was recorded
in different points of the territory (the environs of the
Fringilla biostation (on average, 20 songs from each
male). In sum, approximately three thousand five
hundred songs were investigated.
Subsequently, song sonograms were analyzed using
the Avisoft SaSLab Light program. In sum, seven
main frequency and time parameters were measured:
structure, duration, and number of elements in a song
type; duration of elements in a trill; maximal, mini
mal, and medium frequency of a song, and intervals
between songs. The chaffinch song is traditionally
divided into three parts: whistle sounds (the introduc
tion), trill sounds (the medium part), and a flourish
[10, 11]. Separate phrases consisting of elements sim
ilar in form (syllables) can be distinguished in sono
grams within each part (Fig. 2). Song types were des
ignated by Latin letters. The songs that had two or
three similar parts were referred by us to single type
songs.
For example, the songs A1 and A2 relate to the type A
(Fig. 3a), and I1 and I2 similar in two phrases relate to
In sparrow birds, singing is the immanent part of
their vital activity that determines and structures the
preparation and course of the reproductive cycle.
A bird song has the most important biological func
tions. It is not only the instrument for attracting
females but also the instrument for browbeating a rival
and determining the limits of a nest territory in a pop
ulation [1–3]. Detailed investigation of the song rep
ertoire in many species has shown that an individual
can have not one but several song types [4–9].
The specific bird songs differ for individuals relat
ing to different geographical populations; there is also
an individual song variation in males relating to the
same population [7, 8].
The chaffinch (Fringilla coelebs L.) is the classical
object of investigation of a vocal repertoire [10–12]
and geographical variation in songs [9, 13–15].
The goal of this research is to ascertain the typolog
ical organization and to reveal the microgeographical
variation in chaffinch songs in the populations of the
Curonian Spit. For this purpose, the repertoire and
song types of chaffinches living in the Curonian Spit
have been revealed using qualitative and quantitative
analysis. Several development properties of song cul
ture in the species of the population have been noted.
MATERIAL AND METHODS
The main work was carried out in the Curonian
Spit in the environs of the Fringilla biostation but
chaffinch songs were also recorded in other regions of
78
THE TYPOLOGICAL ORGANIZATION AND MICROGEOGRAPHICAL VARIATION
79
The Curonian Spit
Morskoi
Rybachii
Slavsk
Dunes
Lesnoi
Svetlogorsk
Neman
Zelenogradsk
Pionerskii
Krasnoznamensk
Polessk
Guryevsk
Svetlyi
Baltiysk
Chernyakhovsk
Nesterov
Gusev
Kaliningrad
Pravdinsk
Bagrationovsk
Mamonovo
Fig. 1. The map of the Curonian Spit.
kHz
20
15
10
5
Whistle elements
1 phrase
1
2
3
4
1
0.2
0.4
0.6
0.8
2
3
4
1.0
Flourish
Trill elements
2 phrase
5
1
1.2
2
3
1.4
1
2
3
4
1.6
1.8
4
2.0
5
2.2
s
Fig. 2. Structure of the chaffinch song.
the type E. However, most song types were similar in
all three phrases (Fig. 3.2).
RESULTS AND DISCUSSION
In the investigated chaffinch population, we
revealed 22 song types, of which four turned out to be
rare (they were found in 1–2 record points), and five
song types were combined (they consisted of phrases
of different song types). The average size of the
chaffinch repertoire in the Curonian Spit was repre
sented by 2–3 song types (max of 6 song types and min
of 1 song type). The repertoire and succession of perfor
mance of song types are exemplified by the song record
of one of chaffinch males recorded in point no. 14:
R R E2 R E2 E2 E2 E2 R R R R R R R R R R R R R
R R R R R E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2
R R E2 R R R R R R E2 E2 E2.
Only two song types (R is 29 (56.9%), E2 is 22
(43.14%), and n = 51) performed with an almost anal
ogous frequency were noted in its repertoire.
The ratio of song types in repertoires of different
chaffinch males is reflected in Table 1.
THE INDIVIDUAL VARIATION
IN SONG TYPES
The individual variation in chaffinch song types
was revealed, which can be traced when comparing the
most distributed song types in the population of these
birds in the Curonian Spit (A and C) (Table 2). The
individual variation is likely to have a different charac
ter for other song types found by us in the population
(they number more than 20).
For both song types, the greatest differences are
discovered in the duration of a song and number of syl
lables. The minimal and maximal frequencies of songs
vary insignificantly.
MICROGEOGRAPHICAL VARIATION
When the song types from different settlements
were analyzed, it turned out that there was a small
microgeographical variation. For example, the songs
of type G differed in length and number of syllables. In
addition, some difference in values of maximal fre
quency was discovered, which was not revealed when
MOSCOW UNIVERSITY BIOLOGICAL SCIENCES BULLETIN
Vol. 65
No. 2
2010
80
ASTAKHOVA, BEME
(a)
kHz
20 Whistle elements
15
10 1 2 3 4
5
0.2
kHz
20 Whistle elements
15
10 1 2 3 4 5
5
0.2
(b)
kHz
20
15
10
5
1
3
2
0.4
0.6
2
1
0.4
0.8
Song type A5
2
4
3
1.2
1.4
3
0.4
0.6
0.4
6
1.2
1
2
3
1
7
1.4
4
1
2
3
6
1.2
0.6
0.8
Song type I3
4
1.0
1.8
s
Flourish
5
0.8
1.0
Song type I2
2
1.6
1
1.4
Trill elements
3
2
5
Trill elements
4
s
Flourish
0.8
1.0
Song type A6
Whistle elements
0.2
1.0
4
3
0.6
2
1
1
5
4
Whistle elements
1
Flourish
Trill elements
0.2
kHz
20
15
10
5
Trill elements
1.6 s
Flourish
1
5
2
1.2
3
1.4
s
Fig. 3. Principles for distinguishing single type songs recorded in different chaffinch populations. (a) Song type A (variants A5 and A6).
(b) Song types I (variants I2 and I3).
comparing the single type songs inside a settlement
(Table 2).
Analyzing the properties of performance of single
type songs in several settlements by the example of the
type M, we see that the same song type can occur both
in different individuals in one micro settlement and in
other settlements. Moreover, the differences inside a
settlement are comparable with the differences in song
types between settlements (individual and microgeo
graphical variation). Three parts of songs of these
types are perceived orally in different ways: the first
two phases of M1, M2, M4 sound as the introduction
to a song (a series of whistle sounds), meanwhile, in
the song types M5, M11, M6, these elements are in the
middle of a song and heard as a trill. It is of interest to
note that the introduction to a song (whistle sounds) in
the song type M16 (Fig. 4) is doubled by two analo
gous phrases; moreover, the last element of these
phrases sounds as an acute gritting sound, rather than
a thin whistle sound. Nevertheless, it is included in the
first part of the song that is represented in most types
by sounds of a thinner tone (the introduction). In
addition, the trill elements of this song type differ in
form and, to a certain extent, in tune (the phrase that
is behind the flourish): in some types, the emphasis is
placed on the upper bend of trill elements (M2, M5,
M11), in other types, it is placed on the lower bend
(M1, M4, M16).
When comparing single type songs from regions
remote from each other, one can reveal the differences
in their frequency and time parameters. Such differ
ences were attributed by us to the microgeographical
song variation. The geographic variation in songs of
different sparrow species and formation of dialects
that are perceived orally as “local melodies” were dis
cussed in many works [14, 16–18].
MOSCOW UNIVERSITY BIOLOGICAL SCIENCES BULLETIN
Vol. 65
No. 2
2010
THE TYPOLOGICAL ORGANIZATION AND MICROGEOGRAPHICAL VARIATION
The microgeographical variation was defined by us
as the existence of variants (variation) of single type
songs fixed in different territories, the distance
between each of which averaged 10 km (Fig. 1). The
microgeographical variation in song types is feebly
marked. Possibly, this may be explained by the high
density of the chaffinch population in the Curonian
Spit and by the fact that males learned songs in their
“communicative” groups well (without strong devia
tions in the frequency and time parameters). Despite
the similarity in sonograms of single type songs, some
differences in the frequency and time parameters were
nevertheless revealed. Table 2 shows the results.
81
Table 1. The ratio of chaffinch (Fringilla coelebs L.) song
types in the environs of the Fringilla biostation
Quantity of
Number
song types in
of a male
a repertoire
Percentage ratio
of song types
1
2
C – 35%, G – 65%
2
2
C2 – 51%, H1 – 49%
3
2
M – 43.5%, N – 56.5%
4
1
N1 – 100%
5
2
E1 – 21%, O – 79%
6
1
C3 – 100%
THE MIXED SONG TYPE
7
1
C4 – 100%
The cases of composing new song types from the
combination of phrases of different males recorded
in different regions of the Curonian Spit were noted
(Fig. 4). The similar cases had earlier been described
only for the birds raised under the experimental con
ditions, when a caught male was daily played different
songs of two wild males in the sensitive period. As a
result, in some period, the laboratory bird formed a
song joining the phrases of two wild males [12]. The
cases noted by us are also likely to be an evidence for
the nonstandard learning of song types in the period of
their crystallization.
As the works of M. Beecher have shown [19], the
formation of a song in the birds brought up in a labo
ratory has a whole series of differences. In the song
sparrow (Melospiza melodia), the male repertoire con
sists of 5–10 song types. Nestlings brought up under
natural conditions learn a song of only one male (most
often, that of their father), although they also hear
other singing birds; under experimental conditions,
8
1
O1 – 100%
9
6
O2 – 18%, O3 – 13%, P – 5%,
C5 – 16%, I2 – 45%, A4 – 3%
10
5
A5 – 6%, A6 – 6%, O3* – 35%,
O4 – 12%, I3 – 41%
11
2
E2 – 56.9%, R – 43.14%
12
2
T – 58%, S2 – 42%
nestlings mix melodies of many males in their song.
The similar mixing of song types was noted in one
chaffinch male in the Curonian Spit. The territory of
this male was located at the boundary of the popula
tion, and its repertoire was revealed to include the
maximal number of song types: 6.
kHz Whistle
20 elements
15
10 1 2 3 4
5
kHz
20 Whistle elements
15 1 ph.
2 ph.
10 1 1 2 3 4 5 6
5
Trill elements
1
2
3
4
Notes: The repertoire of chaffinch songs was taken into consider
ation when selecting songs of one male on the average n > 20.
* Single type songs in different record points were noted by
numbers in the ascending order.
Flourish
5
1 2 3 4 5 6
7
0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 s
Song type E2
kHz Whistle elements
(The type E)
20
15
10 1 2 3 4 5
5
0.2
0.4
0.6
0.2
0.4
Trill elements
1
0.6
1 phrase (The type E)
2
0.8
3
2 phrase (The type M)
4
1
3
4
5
0.8 1.0 1.2
Song type M1
6
7
1.4
1
1.6
2
3
1.8
s
Flourish
(The type M)
Trill elements
1
2
Flourish
2
1.0 1.2 1.4 1.6
Mixed song type EM
3
1
4
1.8
2
2.0
3
2.2
2.4 s
Fig. 4. Sonograms of combined song types recorded in different regions of the Curonian Spit. Noncombined song types (the
record in the environs of the Fringilla biostation: song type E2, song type M1), mixed song type EM (Lesnoi settlement).
MOSCOW UNIVERSITY BIOLOGICAL SCIENCES BULLETIN
Vol. 65
No. 2
2010
MOSCOW UNIVERSITY BIOLOGICAL SCIENCES BULLETIN
Trial point
Point 2
Point 8
Point 9
Point 11
Point 13
Rybachii settlement
C
C2*
C3
C4
C5
C6
G1
Vol. 65
2.46 В± 0.02
2.3 В± 0.21
2.72 В± 0.18
1.96 В± 0.12
1.75 В± 0.15
2.01 В± 0.21
2.074 В± 0.118
1.917 В± 0.11
2.025 В± 0.11
1.8721
2.0511 В± 0.16
1.5093
1.8605
2.36 В± 0.026
2.0114
Length of
a song, s
7.9 В± 0.49
8.23 В± 0.6
1.58 В± 0.077
8.65 В± 0.67
1.47 В± 0.09
1.54 В± 0.14
7.522 В± 0.36
7.8 В± 0.5
1.55 В± 0
1.7 В± 0.13
9.25 В± 0.6
7.88 В± 0.48
1.48 В± 0.14
1.75 В± 0.015
8.01 В± 0.37
1.447 В± 0.166
9.698 В± 0.41
8.096
1.722
1.627 В± 0.131
7.5 В± 0.6
1.5 В± 0.1
3.72 В± 0.094
3.76 В± 0.17
3.89 В± 0.125
4.36 В± 0.2
4.3 В± 0.07
4.056 В± 0.22
4.0996 В± 0.173
4.164 В± 0.213
4.134 В± 0.215
4.478
4.306 В± 0.3
4.306
7.579
1.894
4.2 В± 0.1
3.962
Medium
frequency,
kHz
4.306
7.6 В± 0.46
7.407
Maximal
frequency,
kHz
7.924
1.55
1.5 В± 0.1
1.722
Minimal
frequency,
kHz
17 В± 0
16.2 В± 1.8
18.2 В± 0.97
15 В± 1
13.7 В± 1.03
16.45 В± 1.37
17.25 В± 0.85
17.06 В± 1.16
16.7 В± 1.16
14
15 В± 1
13
13
18.3 В± 1.15
16
Number of
syllables in
a song type
0.067 В± 0.002
0.064 В± 0.005
0.076 В± 0.004
0.0632 В± 0.005
0.067 В± 0.004
0.066 В± 0.005
0.068 В± 0.003
0.05 В± 0.004
0.084 В± 0.028
0.11 В± 0.003
0.11 В± 0.006
0.099 В± 0.007
0.099 В± 0.007
0.12 В± 0.01
0.136 В± 0.024
0.15 В± 0.004
0.156 В± 0.012
0.165 В± 0.008
0.103 В± 0.003
0.0913 В± 0.05
0.128 В± 0.01
0.1155 В± 0.008
0.094 В± 0.009
0.13 В± 0.007
Length of syllables
(trills), s
Notes: The average value and standard deviation of parameters of song types that are taken from statistical calculations for all single type songs reproduced by chaffinch males in these
record points are indicated.
* The song types in the large volume repertoires were unitary; the difference in parameters >0.5 kHz in a frequency and >0.02 s in a length were considered to be the strongest
(they are in bold).
Fringilla biostation
Point 13
A7*
G3
Point 13
A6
Lesnoi settlement
Point 12
A5*
G2
Point 11
Point 3
A3
A4*
Trial point
A1
Song type
Number of a record
(designated
point
by a letter)
Table 2. The main frequency and time characteristics of song types (A, C, G)
82
ASTAKHOVA, BEME
No. 2
2010
THE TYPOLOGICAL ORGANIZATION AND MICROGEOGRAPHICAL VARIATION
CONCLUSIONS
During the investigation of the songs in the
chaffinch population in the Curonian Spit, their typo
logical organization as well as insignificant microgeo
graphical variation in single type songs have been
revealed. The chaffinch song is known to have a
genetic basis but the variation in song types in the pop
ulation is constantly being supplemented by learning,
improvisation, and errors in copying. The discovered
variation of song types can speak for the genetic heter
ogeneity of the population and microevolution pro
cesses taking place in it. This paper preliminarily ana
lyzes the typological organization of the songs of
chaffinches living in the Curonian Spit. A more
detailed description of the structure of each song
phrase, frequency, and time organization of songs will
be presented in the following works.
ACKNOWLEDGMENTS
We are deeply grateful to the workers of the
Rybachii biostation of the Zoological Institute, Russian
Academy of Sciences, in the Curonian Spit for help and
support in work and to professor G.N. Simkin for giv
ing valuable counsels when analyzing the material.
The research was financially supported by the Russian
Foundation for Basic Research (grant nos. 05 04
49173 a and 07 04 00609 a).
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MOSCOW UNIVERSITY BIOLOGICAL SCIENCES BULLETIN
Vol. 65
No. 2
2010
Автор
Olesya Astakhova
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