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The quantitative analysis of chaffinch song

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The quantitative analysis of chaffinch song types (
Fringilla coelebs L
.
) in populations of the European part of Russia
Astakhov
a
O.A. (dissertation) 2006.
•
The quantitative analysis of songs of birds is one of the basic ways of an estimation of structure song
patterns, their set in repertoir
e
s of individuals and a typological comparison
in song culture of populations. With a view of revealing individual and geographical features of song
types on different time
-
and
-
frequency parameters from experiments in local populations of different species of birds many bioacoustics had been lead
(had been carried out) calculations of their statistical data and the subsequent comparisons of values (Barber, 1959; Zablotskaya
, 1981, 1982; Marler and Tamura, 1962; Konishi, 1964; Kroodsma, 1985; Bremond, 1968).
First of all, in researches the characteristic of bird repertoir
e
s (number of song types, their general (common) phrases
or parts
, qualitative structure was mentioned at the certain volume of sample, frequency of occurrence and a
comparison of
song
patterns in repertoir
e
of separate individuals and in a population, as a whole) (Brown, 1985; Krebs, 1976; Kroodsma, 1976; Whitney, 1987; Dobson and Lemon, 1975; Payne et al., 1981; Falls and Krebs, 1975; Lemon and Chatfield, 1971). Chaffinch
(
Fringilla coelebs L
.
) –
one of the most widespread species of birds which were objects of studying song organizations (Hinde, 1958; Slater, 1980, 1983). As against many others, chaffinch
has harmon
i
ous, frequently short, precisely organized and in the greater degree song genetically determined in a time
-
and
-
frequency range (Thorpe, 1958; Marler, 1952; Nottebohm, 1969, 1967). The size of repertoir
e
of one individual is within the limits of 1
-
6 types of songs (Slater, 1981; Marler, 1956).
The problem of our work will consist in definition of the quantitative characteristic of repertoir
e
and the typological organization of chaffinch song (
Fringilla coelebs L
.
), revealing of individual and geographical features of song
patterns.
In different areas of the European part of Russia in spring
-
summer
period
of 2005
–
2006 years
. Records of singing males
have been made. For record of songs used tape recorder Panasonic RQ
-
SX95F, condenser microphone Philips SBC ME570. Further
,
sonogramms of
songs were analyzed with the help of computer program Avisoft
-
SaSLab Light
. In total it has been analy
z
ed about five thousand songs. Types of songs were mark
ed by L
atin letters. At the quantitative analysis measured both structure of song type as a whole, and its separate phrases (in three parts of songs –
started singing, a trill, a final stroke) on different frequency
-
time parameters: length (duration), maximal, minimal and average (median) frequency, quantity
(amount) of elements (syllables), length
(duration)
of syllables, intervals between syllables, intervals between phrases.
During statistical processing the primary data basically defined arithmetic
-
mean value and a standard (arithmetic
-
mean) deviation (with the help «Master of functions» of Microsoft Excel). At comparison frequency
-
time parameters are allocated (
distinguished)
a range
,
within the limits of which the difference of values was considered insignificant (
if values
difference
equal or smaller 0,5 К
Hz
in frequency and 0,02 sec
in length). The characteristic of song
types and repertoir
e
s of males
with the greater sample of songs (n> 14) was more significant.
Material and methods
Song repertoir
e
includes
song types, which is capable to reproduce male of
chaffinch (
Fringilla
coelebs L
.
) at singing. S
ong
t
ype –
individual (within the limits of one population) or geographical (in populations o
n different territories) song
variant of a species.
It is usually these song
patterns in its
individual becoming are precisely determined in repertoir
e
of an adult individual after a stage of crystallization (stabilization) of
species specific
song
. The repertoir
e
of young individuals frequently is established not completely
, because
after sensitive
period of training (learning) song types are still plastic in the structure, can change or not appear at all under influence of "communicative" group (Thorpe, 1958; Marler, 1956; Nottebohm, 1967).
On the basis of experiments in different local populations the statistical analysis of the size of chaffinch repertoir
e
(
Fringilla coelebs L
.
) has been lead
(has been carried out). Those individuals from which n>14 songs have been recorded were taken into account only. The primary data are shown in tables (tab. 1, 2). In northwest of the European Russia from the general (common) sample are allocated males
, recorded in different areas
of Curonian spit of Baltic Sea (settlement Sea
or Morskoy
, settlement Fishing
or Ribachiy
, area of biological research station "
Fringilla
", settlement Wood
or Lesnoy
) and having on the average n
> 20 reproduced songs (tab. 1).
In the central part of the European Russia at processing records of chaffinch
(
Fringilla
coelebs L
.
) also were taken into account males
with volume of sample of songs on the average n> 20. In the table repertoir
e
s of individuals from different areas (
Zvenigorod
, Moscow, Michurinsk) are shown (tab. 2). Results and discussion
Table 1.
The characteristic of chaffinch males repertoir
e (
Fringilla coelebs L
.
) in populations of northwest of the European Russia (
Curonian spit, Kaliningrad region
)
Area of biological research station "
Fringilla
"
Number of a point (male)
record
Volume of sample
songs
(
n)
Number of song types in repertoire (designation)
Number of a point (male)
record
Volume
of sample
songs
(
n)
Number of song types in repertoire (designation)
Number of a point (male)
record
Volume
of sample
songs
(
n)
Number of song types in repertoire (designation)
1 (4)
43
2
(C G)
8
20
1
(C 3)
13
55
6
(O3*,O5 I4, I5 A5,A6,A7 C6 OI 5 OC6)
2
43
2
(C*2 H1)
9
2
1
1
(
C 4)
14
51
2
(E 2 R)
4
20
2
(H2 L)
10
15
1
(O 1)
16 (2)
43
2
(
T S2)
5
23
2
( M N)
11 (1)
38
6
(O2 O3 P C5 I2 A4)
17
14
3
(D1 D2 U)
6
31
1
(N1)
12
17
5
(A5 A6 O3* O4 I3)
18
20
2
(F2 C7)
7
19
2
(E1 O)
Settlement Fishing (Ribachiy)
1 (4)
19
3
(
I J K)
2 (2)
15
1
(O 17)
3 (2)
32
3
(M15 M16 T3)
Settlement Sea (Morskoy)
1
21
1
(
D 1)
5
29
2
(
J4 O6)
13
21
2
(J4 F4)
3
22
2
(
Т3 С)
8
19
2
(C*13 Q)
Settlement Wood or Lesnoy
1
38
2
(М 2 С 8
)
2 (2)
13
1
(
C* 9)
5 (2)
13
2
(S3 C*10)
2 (1)
2
3
2
(R
3 J2
)
3
15
2
(M 3 T 2)
The note: the characteristic of chaffinch repertoir
e
was taken into account at volume of song
sample from one male
n> 14; song
types were mark
ed by latin letters; songs of one type in different points of record were marked by numbers in ascending order.
Table 2.
The characteristic of chaffinch male repertoir
e
(
Fringilla coelebs L
.
) in populations of the central part of the European Russia
Zvenigorod
(Moscow region)
Number of
a point (male)
record
Volume of
sample songs
(
n)
Quantity
of
song types in repertoire (designation)
Number
of a point (male)
record
Volume of sample
songs
(
n)
Quantity of
song types in repertoire (designation)
Number
of a point (male)
record
Volume of sample
songs
(
n)
Quantity of song types in repertoire (designation)
1
21
3
(W3 V1 I10)
10 (2)
14
1
(A3)
18 (2)
15
1
(SNE3)
4
14
2
(NIE2 I3)
10 (4)
14
2
(
F I5)
19
21
2
(
V J2)
7
25
2
(A U)
11
17
2
(B NIE)
20 (2)
27
1
(N2)
8 (1)
20
1
(N)
12
17
1
(SNE)
20 (3)
17
1
(U3)
8 (2)
20
1
(C
#
)
15 (1)
15
2
(F I7)
22
48
3
(D2 C3 W2)
10 (1)
19
1
(C
#
2)
17
14
2
(I8 SNE4)
24 (1)
34
2
(N4 S2)
Moscow 1
25
2
(
I12 C
#
5)
8 (1)
19
3
(J J* J8)
9
41
2
(D3 NIE3)
2
15
2
(C
#
6 J7)
8 (2)
27
1
(C
#
10)
10 (2)
16
2
(C
#
12 A5)
Michurinsk (Tambov region)
1
23
2
(BF Y)
12
55
2
(N5 F3)
20 (2)
36
3
(Y3 B7 J12)
3
28
2
(CM TS)
13
20
1
(C сокр.)
21
19
2
(A6 C
#
22)
5
18
2
(J9 TS2)
14
57
2
(
BF5 C
#
19)
22
54
2
(TS4 G)
6
18
4
(Y2 Y*2 CT J5)
15
16
2
(B4 CT2)
23
28
3
(M1 OE A7)
The note: the characteristic of chaffinch repertoir
e
was taken into account at volume of song
sample from one male n> 14; song
types were mark
ed by latin letters; songs of one type in different points of record were marked by numbers in ascending order.
On the basis of the primary data arithmetic
-
mean value and a standard deviation
of number of males
song types have been calculated. In result appeared, that the average size of chaffinch repertoir
e
(
Fringilla coelebs L
.
) in different regions of the European Russia included approximately 2 types of songs (
t
ab. 3).
Table 3
. The average size of chaffinch repertoir
e
(
Fringilla coelebs L
.
) in different areas
of
t
he European Russia
Northwest of European Russia (Curonian spit)
_
x ±
õ
Central part of European Russia
_
x ±
õ
Area of research biostation «
Fringilla
»
2,5 ±
1,67
Zvenigorod
(
Moscow region
)
1
,68 ±
0,67
Settlement Fishing (Ribachiy)
2,3
3
±
1,5
Moscow
2 ±
0,63
Settlement Sea (Morskoy)
1
,83 ±
0,4
Michurinsk
(
Tambov region
.)
2,1 ±
0,66
Settlement Wood (Lesnoy)
1
,8 ±
0,45
Total
2,115 ±
0,35
Total
1,93 ±
0,22
The note:
the volume of sample of songs from one male
in the center of the European Russia has on the average made 24,7 ±
11,3; average volume of song sample from one male
in northwest of the European Russia 26,8 ±
12,25.
The maximum quantity of song types
,
which are included in chaffinch repertoir
e
–
six, is found in samples of Curonian spit (
area of
biological research stations "
Fringilla
") (tab. 1). The minimal size of repertoir –
1 song type. Nevertheless, quite often there were repertoir
e
s with other quantity of song types (tab. 4).
Table 4.
A percentage comparison
of chaffinch repertoir
e
s (
Fringilla coelebs L
.
) in populations of the European Russia
Repertoire size
(
quantity of song types
)
Quantity of males with the given size of repertoire
Percent from the common number of individuals in a population
1
18
22 %
2
46
56,1 %
3
8
9,76 %
4 -
6
4
4,9 %
The note: in sample of populations of the European Russia (N = 137) those were taken into account chaffinch males, from which
have been recorded on the average n>20 songs (them appeared n = 76
).
Very seldom in populations of chaffinch (
Fringilla coelebs L
.
) there were the repertoir
e
s consisting of 4
-
6 song
types (4 male
-
4,9 %). The repertoir
e
from 3 song
types –
also was rare (8 male
-
9,76 %). The often phenomenon –
2 song types in repertoir
e
(46 male
-
56,1 %). Even the repertoir
e
from one song
type met less often (18 male
–
22 %) (probably because the majority recorded males
with such repertoir
e
have not come in the account because of small song
sample, that is n <14).
It is necessary to note, that it is the generalized statistical calculations. In the nature there is all is very relative. Sometimes it is possible to record of one chaffinch male during hours, and in
its repertoir
e
only 1
-
2 song
types
are found
, though the bird can reproduce and a lot song
patterns (it is allowable, 3
-
4). But it is possible to conduct record and 10 –
15 minutes –
and to grasp thus whole repertoir
e
of an individual (4, 5, 6 types of songs? And it is possible, only and, really, –
1
-
2 types). The rare song type in repertoir
e
can appear once (time at one o'clock, at two o'clock, in day, week?). Whether it will get in sample of the researcher? (Kroodsma, 1974; Slater, 1983).
Interestingly typological comparison
of chaffinch songs (
Fringilla coelebs L
.
) in local populations. In samples on Curonian spit of Baltic sea (
N
=153)
we allocate
(
distinguish)
22 song
types which were mark
ed by latin letters (A B C D E F G H J I K L M N O P Q R S T U V). At record songs of one type came across in different points of territory (was considered, that belong to repertoir
e
s of different males
), therefore equally with the letter were designated by numbers in ascending order (for example, А1, А2, А3, etc.).
In populations of the central part of the European Russia
(
N
=65) 15 different song types
are found
,
from which 12 song types were similar in the structure to corresponding types on Curonian spit
, but frequently had the modified elements on sonogramms
, therefore were considered as dialect forms (B C D F G I J M N S U V). The
song type A was stable in the structure in different regions of Russia. It has not been found analogues six
song types (H K Q P R T), and types W
,
Y appeared new in relation to known to us song patterns on Curonian spit
. Both in northwest, and in the center of the European Russia the most widespread appeared type C. On the second place on a degree of distribution in local settlements was
the song type I. Song types R and M are more often
,
than others met at record only on Curonian spit
. But song
culture of a population
are
characterized
by different
song types, which
hav
e
the big sample
size
: A, F, S, O, T, J (tab. 5). Rare song
types are
V, P, Q, L (on Curonian spit
). In the center of the European Russia rare song types could be and what are distributed enough in northwest of Russia (G, M, S), and any song types and did not meet at all (E, H, O, R, T). It can speak about variety
and the certain originality song
cultures of different local populations.
Table 5.
A percentage comparison of chaffinch song types (
Fringilla coelebs L
.
) in different regions of the European Russia
Type of song
(
token
)
Northwest of European Russia
(
Curonian spit
)
Center of European Russia
(
Moscow
,
Zvenigorod
,
Michurinsk
)
Number of song types in sample
Percent from the common number of song type in a population
Number of song types in sample
Percent from the common number of song type in a population
A
18
9,84 %
8
8,6 %
B
5
2,73 %
8
8,6 %
C
38
20,8 %
24
25,8 %
D
11
6,01 %
3
3,2 %
Е
7
3,82 %
F
16
8,74 %
5
5,4 %
G
10
5,5 %
1
1,07 %
H
6
3,28 %
I
22
12,02 %
11
11,83 %
J
13
7,1 %
12
12,9 %
K
3
1,64 %
L
1
0,5 %
Table 5.
A percentage comparison of chaffinch song types (
Fringilla coelebs L
.
) in different regions of the European Russia
(c
ontinuation
)
M
20
10,93 %
2
2,15 %
N
8
4,4 %
5
5,4 %
O
18
9,84 %
P
1
R
24
13,1 %
Q
1
0,55 %
S
17
9,3 %
2
2,15 %
T
14
7,7 %
U
8
4,4 %
3
3,2 %
V
2
1,1 %
2
2,15 %
W
3
3,2 %
Volume of sample individuals (
n)
158
65
Volume of sample song
types
(
n
)
183
93
Type of song
(
token
)
Northwest of European Russia
(
Curonian spit
)
Center of European Russia
(
Moscow
,
Zvenigorod
,
Michurinsk
)
Number of song types in sample
Percent from the common number of song type in a population
Number of song types in sample
Percent from the common number of song type in a population
The note: the volume of sample includes all recorded males
in populations (the bottom line); a percentage parity
(ratio) expected proceeding from the general
(common) number of song
types in samples of populations in northwest of the European Russia –
183 song types, in the central part of Russia –
93 song
types (that is repertoir
e
s of males
with small samples were taken into account also); thus it is necessary to understand, that songs of one type from everyone male
had sample on the average
n>20; the most widespread (frequently met) in populations song
types
(with sample n> 15) are allocated by a font.
To have more precise representation about song types
,
which we discuss, it is possible to result (show)
sonogramms
of their samples (fig. 1) and some statistical data of their basic time
-
and
-
frequency parameters (tab. 6).
1.
3.
4
.
7.
8.
6.
5.
2.
Fig. 1
.
Most widespread song types of chaffinch
(
Fringilla coelebs L
.
) in a population of northwest of the European Russia (
Curonian spit
): 1 –
song type A, 2 –
song type C, 3 -
song type C*, 4 –
song type F, 5 –
song type I, 6 –
song type М1, 7 –
song type М12, 8 –
song type O.
9.
10.
11.
Fig. 1
. Most widespread song types of chaffinch
(
Fringilla coelebs L
.
) in a population of northwest of the European Russia (
Curonian spit
): 9 –
song type R, 10 –
song type S, 11 –
song type J. At close (attentive) viewing of sonogramms
it is possible to allocate
(
distinguish)
a number
of features in structure of songs, characteristic only for the certain type. It is necessary to remind, that we considered songs are similar on two or all three parts as
songs of one type
(started singing or the row of whistle
elements, a trill and a final stroke)
,
which also can include one or two phrases (the elements similar under the form). The row of whistle
elements (started singing) in songs of chaffinch
(
Fringilla coelebs L.) can be as arc
-
similar
, and narrow linear forms. But on hearing all of them are perceived by higher tone, as if a number
whistle
sounds "
fuit
-
fuit
-
fuit
".
The trill with equal frequency can consist both of one phrase, and from two. Elements of a trill can be completely different form, consist of two and more subelements. But if to look narrowly, nevertheless it is possible to notice, that in many respects on the structure elements are similar –
only their primary basis in different types differently has been sung
.
The same is possible to tell about a final stroke –
usually sharp on the hearing consisting from stick
-
similar
elements on sonogramms
, has much in common at different song
types, but also is individual for everyone song pattern.
For full representation of song cultures in
different local populations
of chaffinch
(
Fringilla coelebs L
.
) it is necessary to show samples of the most widespread song
types in territory of the center of the European Russia
,
which can not coincide with a degree of their spreading in other populations
.
1.
3.
2.
4.
5.
6.
Fig. 2.
Most widespread song types of chaffinch (
Fringilla coelebs L
.
) in populations of the central part of the European Russia:
1 –
song type C, 2 –
song type C*, 3 –
song type I, 4 –
song type J, 5 –
song type A, 6 –
song type B.
Song type C (fig. 2
.1) (its version type C
#
(fig. 2
.2)) and song type I (fig. 2
.3) appeared in the majority both in samples in northwest, and in the center of the European Russia (tab. 5). But song type I –
the dialect form in relation to songs of this type on Curonian spit
. The song
type J (fig. 2
.4) also is distributed enough in the center of the European Russia, but in northwest this type was average on occurrence. S
ong
t
ype A, B
(fig. 2
.5 and 2
.6) also have average prevalence too. These are the basic song
patterns
,
which frequently met in populations of all two regions of Russia where carried out research (tab. 5). It is possible to result examples of rare song types in populations of the European Russia (at record are found in 1
-
2 points) (fig. 3).
1.
2.
3.
4.
Fig. 3
.
Sonogramms of
rare
chaffinch
song types
(
Fringilla coelebs L
.
)
in European Russia
: 1 –
song type K, 2 –
song type L, 3 -
song type V, 4 –
song type Q.
Than it is possible to prove a rarity of these types of songs? (fig. 3) It is possible, such song
structure was not in the greater degree characteristic for
species
specific songs
of
chaffinch
(
Fringilla coelebs
L
.
) therefore it had no "support" for distribution to "communicative" groups of a population. But can be, it is simple song
types of migrants from other populations,
song
culture
of which includes their majority.
It is impossible to miss from attention on the so
-
called combined types of songs. In the first researches on Curonian spit these song
forms were individual
(
unique)
, and in the subsequent experiments in the center of the European Russia –
song
types
combined of modified elements
(in relation to already known) appeared in the majority and in many respects characterized song
culture of a local population (tab. 7).
Table 7.
A percentage parity (ratio) of the combined song types of chaffinch
(
Fringilla coelebs L
.
) in populations of the European Russia
Place of record
(
region of Russia
)
Song type
(
token
)
Number of song types in sample
Percent from the common number of song types in a population
Volume of sample
type songs
(
n)
Northwest of
European
Russia
(
Curonian
spit
)
Setllement Wood
(Lesnoy)
E I
1
0,55 %
183
E M
1
0,55 %
Area of
biostation
«
Fringilla
»
A C
1
0,55 %
O C6
1
0,55 %
O I 5
1
0,55 %
O C*
1
0,55 %
Total
6
3,28 %
The note: a percentage parity (ratio) expected proceeding from the common number of song types in samples of populations in northwest of the European Russia –
183 song types, in the central part of Russia –
93 song types (that is repertoir
e
s of males
with small samples were taken into account also); thus it is necessary to understand, that songs of one type from everyone male
had sample on the average n>20.
Table 7.
A percentage parity (ratio) of the combined song types of chaffinch
(
Fringilla coelebs L
.
) in populations of the European Russia
(c
ontinuation
)
Central part of European Russia
Zvenigorod
Moskow
, Michurinsk
N I E
3
3,2 %
93
Zvenigorod
S N E
4
4,3 %
Moskow
, Michurinsk
D E
2
2,15 %
Michurinsk
B F
6
6,45 %
O E
1
1,075 %
T S
2
2,15 %
C T
2
2,15 %
C M
2
2,15 %
G C
2
2,15 %
Total
24
25,8 %
Place of record
(
region of Russia
)
Song type
(
token
)
Number of song types in sample
Percent from the common number of song type in a population
Volume of sample
type songs
(
n)
The note: a percentage parity (ratio) expected proceeding from the common number of song types in samples of populations in northwest of the European Russia –
183 song types, in the central part of Russia –
93 song types (that is repertoir
e
s of male with small samples were taken into account also); thus it is necessary to understand, that songs of one type from everyone male had sample on the average n>20.
For more precise representation of these song types it is necessary to show them sonogramms
(fig. 4
).
1.
2.
3.
4.
6.
5.
7.
Fig. 4. Sonogramms of "combined" types of chaffinch songs (
Fringilla coelebs L
.
), recorded in different regions of
the European Russia:
1 -
song
type AC
, 2 –
song type E I, 3 –
song type EM, 4 –
song type ОС6 (are recorded on Curonian spit of Baltic sea
, north of Europe
); 5 -
song
type BF, 6 –
song type N I E, 7 –
song type SNE (are recorded in the center of the European Russia).
+
Table 6.
The quantitative characteristic of the most widespread song types
of chaffinch
(
Fringilla coelebs L
.
) (the example of population in Curonian spit
)
Parameters
of songs
Song type (
token
)
_
x ±
õ
А
S
M
F
C
I
O
R
Duration of song
, sec
1,96
±
0,25
2,29
±
0,25
1,875
±
0,16
2,24
±
0,3
1,96
±
0,11
1,86
±
0,22
1,56
±
0,15
2,05
±
0,086
1,97
±
0,23
Min frequency
, KHz
1,6
±
0,16
1,605
±
0,05
1,718
±
0,12
1,57
±
0,038
1,59
±
0,12
1,56
±
0,065
1,64
±
0,17
1,72
±
0,13
1,63
±
0,063
Max
frequency
, KHz
7,63
±
0,19
7,51
±
0,15
7,6
±
1,25
7,18
±
0,3
8,36
±
0,9
7,8
±
0,47
7,15
±
0,28
7,44
±
0,7
7,683
±
0,42
Median (average
) frequency
, KHz
4,22
±
0,2
3,84
±
0,06
4,06
±
0,38
4,01
±
0,11
4,2
±
0,12
4,1
±
0,22
4,3
±
0,09
4,05
±
0,21
4,1
±
0,14
Number of syllables
14,9
±
1,86
19,45
±
1,48
17,7
±
1,7
15,6
±
1,7
16,03
±
1,4
11,9
±
1,35
12,05
±
1,61
17,5
±
0,95
15,6
±
2,7
Duration
of syllables
,
s
ес:
0,04
±
0,007
0,12
±
0
0,046
±
0,017
0,11
±
0,02
0,14
±
0,03
0,14
±
0,011
0,06
±
0,004
0,039
±
0,007
0,087
±
0,045
started singing
trill
0,11
±
0,014
0,06
±
0,004
0,08
±
0,006
0,096
±
0,04
0,066
±
0,01
0,083
±
0,004
0,072
±
0,004
0,16
±
0,02
0,09
±
0,032
1
phrase
2
phrase
0,1
±
0,001
0,11
±
0,017
0,06
±
0,005
0,09
±
0,03
final stroke
0,12
±
0,036
0,12
±
0,007
0,041
±
0,016
0,13
±
0,02
0,072
±
0,007
0,15
±
0,043
0,15
±
0,035
0,19
±
0,03
0,12
±
0,05
The note: average value and a standard deviation of parameters of song types are specified; the songs of one type
,
which has been recorded in different points of territory, but not having any differences in the structure
,
were taken into account; in the last column
-
the generalized values on all measured song
types (are allocated or signed by font).
Table 6.
The quantitative characteristic of the most widespread song types
of chaffinch
(
Fringilla coelebs L
.
) (by the example of population Curonian spit
) (c
ontinuation
)
Intervals between syllables
, sec
0,047
±
0,009
0,083
±
0,0003
0,048
±
0,016
0,067
±
0,005
0,061
±
0,02
0,063
±
0,008
0,03
±
0,005
0,057
±
0,017
start singing
trill
0,054
±
0,017
0,026
±
0,002
0,0405
±
0,01
0,052
±
0,01
0,026
±
0,009
0,0485
±
0,005
0,044
±
0,007
0,03
±
0,006
0,04
±
0,011
1
phrase
2
phrase
0,035
±
0,1
0,042
±
0,015
0,03
±
0,007
0,03
±
0,021
final stroke
0,05
±
0,014
0,033
±
0,007
0,055
±
0,009
0,059
±
0,014
0,035
±
0,005
0,069
±
0,0038
0,08
±
0,011
0,05
±
0,041
0,054
±
0,016
Intervals between phrases
,
s
ес:
0,05
±
0,012
0,054
±
0
0,047
±
0,002
0,057
±
0,03
0,05
±
0,006
0,059
±
0,02
0,052
±
0,009
0,033
±
0,004
0,05
±
0,008
start singing
and trill
trill
and
final stroke
0,074
±
0,015
0,066
±
0,016
0,04
±
0,009
0,06
±
0,007
0,047
±
0,022
0,056
±
0,012
0,048
±
0,012
0,056
±
0,016
0,056
±
0,011
Intervals between songs
, s
ес
6,54
±
2,8
7,21
±
3,11
6,6
±
0,37
5,2
±
3,56
7,13
±
2,5
7,58
±
2,18
7,45
±
4,17
7,66
±
0,016
6,92
±
0,81
Number of measured song (
n)
15
24
11
34
57
33
68
37
34,88
±
19,56
Parameters
of songs
Song type (
token
)
_
x ±
õ
А
S
M
F
C
I
O
R
The note: average value and a standard deviation of parameters of song types are specified; the songs of one type
,
which has been recorded in different points of territory, but not having any differences in the structure
,
were taken into account; in the last column
-
the generalized values on all measured song
types (are allocated or signed by a font).
From table 6 it is visible, that values of parameters of song types are various, but nevertheless are in the certain range that can confirm the precise organization and in many respects a genetic basis species specific song of chaffinch
(
Fringilla coelebs L
.
). Despite of the big similarity of values of parameters of
song types, nevertheless it is interesting to find out, on what from them variability was the greatest. For this purpose it is necessary to calculate factor of a variation (V, %) each parameter. The factor of a variation represents a parity
(ratio) of a standard deviation
and average value, shows a degree of a variation of an attribute. It is possible to make the diagram on character of variability of parameters of different types of songs (fig. 5
).
Fig. 5.
Degree of a variation of values of parameters of different song types of chaffinch
(
Fringilla coelebs L.)
on Curonian spit of Baltic Sea (north of Europe)
: 1 –
length (duration) of song, 2 –
the minimal frequency, 3 –
the maximal frequency, 4 -
average (median) frequency, 5 –
number of syllables, 6 –
length (duration) of syllables in started singing
, 7 -
length of syllables of a trill (1 phrase), 8 –
length (duration) of syllables of a trill (2 phrase), 9 –
length of syllables of a stroke, 10 –
intervals between s
yllables (started singing), 11 –
intervals between syllables of a trill (1 phrase), 12 –
intervals between syllables of a trill (2 phrase
), 13 -
intervals between syllables of a stroke, 14 –
intervals between phrases (
started singing
and a trill), 15 -
intervals between phrases (a trill and a stroke), 16 –
intervals between songs.
The most varying attributes (33 –
50 %) different song types appeared (
on Curonian spit, north of Europe)
: length (duration) of syllables in started singing
(6), trill (7, 8) and a final stroke (9) (and, started singing and a final stroke were more variability
). Significant stability (small variability) (3 –
5 %) has come to light at minimal (2), average (median) (4) and the maximal (3) frequencies. Other parameters of different song types had an average variation (10 –
30 %)
,
which can enter into a category of homogeneous values, but among these data there are also those, which variability was the greatest –
intervals between syllables in started singing
(
the row of whistle
elements) (10), intervals between syllables of a trill (11) and intervals between elements of a stroke (13).
Thus, songs of chaffinch
(
Fringilla coelebs L
.
) different types can be stable in frequency, but are more individual in values of length
(duration)
of syllables (elements) and intervals between them.
During the quantitative analysis of song types comparison of their basic time
-
and
-
frequency parameters has been lead (has been carried out) in view of individual (within the limits of one population) and geographical (in populations in different territories) variability (tab. 8).
Fig. 5*. A degree of a variation of values of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in the center of the European Russia: 1 -
length of songs (sec), 2 -
minimal frequency (
К
Hz), 3 -
maximal frequency (
К
Hz), 4 -
average (median) frequency (
К
Hz), 5 -
number of syllables in a song, 6 -
length of started singing syllables (sec), 7 -
length of syllables of a trill (1 phr
ase) (sec), 8 -
length of syllables of a trill (2 phrase) (sec), 9 -
length of syllables of final stroke (sec), 10 -
intervals between syllables
of started singing (sec), 11 -
intervals between syllables of a trill (1 phrase) (sec), 12 -
intervals between syllables of a trill (2 phra
se) (sec), 13 -
intervals between syllables of final stroke (sec), 14 -
intervals between phrases (started singing and a trill) (sec), 15 -
inte
rvals between phrases (a trill and final stroke) (sec), 16 -
intervals between songs (sec).
The factor of a variation (V, %) (fig. 5
, 5*
) shows a degree of a variation (variability) of this or that attribute
(sign),
proceeding from his(its) average arithmetic value (x) and a standard deviation (õ) (tab. 3), which shows as far as on the average each value of an attribute (sign) of the general (common) sample "deviates" from average value of this attribute. The more value of factor of a variation (V, %) is the more variability of values of parameter. Such attitude (relation) can be expressed by the general (common) formula (
Nimenya
, 2003):
õ
V
= –––
×
100%
x
Conclusions on a variation of time
-
and
-
frequency parameters of different song types of chaffinch (
Fringilla coelebs
L.) from
two geographical populations of the East Europe
а)
Similarities
on
a
variation
of
values
of
quantitative
parameters
of
chaffinch
songs
(
Fringilla
coelebs
L
.
)
in
two
geographical
populations
:
•
p
oorly
(
weak)
varied
(factor
of
variability
(V)
3
-
9
%
)
in
all
two
geographical
populations
of
chaffinch
are
minimal,
maximal
and
average
(median)
frequency
(
KHz
)
–
it
is
probable,
the
frequency
range
of
species
specific
song
of
chaffinch
is
genetically
determined
;
•
t
he
average
variation
(factor
of
variability
(V)
10
-
30
%
)
in
all
two
geographical
populations
of
chaffinch
was
observed
at
such
parameters
as
length
of
songs
(
sec
),
the
number
of
syllables
in
a
song,
intervals
between
songs
(
sec
)
,
i
ntervals
between
syllables
of
a
trill
(
2
phrase)
(
sec
),
intervals
between
phrases
(
started
singing
and
a
trill)
(
sec
),
intervals
between
a
trill
and
a
final
stroke
(
sec
)
–
is
probable,
it
also
genetically
determined,
more
or
less
stable
attributes
(signs)
of
species
specific
song
of
chaffinch
,
but
capable
in
the
greater
degree
to
change,
vary,
than
frequency
of
songs
(К
Hz
)
at
individuals
of
chaffinch
at
any
geographical
population
;
•
t
he
strong
variation
(factor
of
variability
(V)
33
-
50
%
)
in
all
two
geographical
populations
of
chaffinch
was
at
parameters
of
song
types
:
length
of
syllables
in
started
singing
(
sec
),
length
of
syllables
of
a
trill
(
1
phrase)
(
sec
),
length
of
syllables
of
a
final
stroke
(
sec
)
–
these
parameters
of
songs
are
capable
to
change
strongly
in
a
population
of
chaffinch
at
song
training
(learning)
and,
probably,
at
genetic
variability
as
well
to
create
different
song
forms
(types
of
species
specific
song
,
distinguished
on
structure
and
the
form
of
elements)
.
Table 8.
Results of comparison of parameters of different song types of chaffinch
(
Fringilla coelebs L
.
) in populations of the European Russia at individual and geographical variability
+
Parameters
of songs
(
quantity
of distinctions
)
Song type (
token
)
А
G
C
E
I
B
Ind
.
1
variab
.
Geog. 2
variab.
Ind.
1
variab
.
Geog 2
variab.
Ind.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind
.
1
vari
.
Geog 2
variab.
Duration of song
, sec
number
21
7
8
8
14
19
4
3
20
7
3
6
% *
20,2
17,5
13,3
15,4
18,9
16
14,3
15
15,87
12,73
18,75
14,3
Min
frequency
, KHz
number
3
%
2,5
Max
frequency
, KHz
number
1
2
5
4
9
8
4
1
11
2
6
%
0,9
5
8,3
7,7
12,2
6,72
14,3
5
8,7
3,6
14,3
b
)
D
istinctions
on
a
variation
of
values
of
quantitative
parameters
of
chaffinch
songs
(
Fringilla
coelebs
L
.
)
in
two
geographical
populations
:
•
t
he
length
of
syllables
of
a
trill
(
2
phrase)
(
sec
)
is
average
varied
(factor
of
variability
(V)
10
-
30
%
)
in
populations
of
the
center
Europe
an
Russia
,
but
in
northern
population
is
strongly
varied
(factor
of
variability
(V)
33
-
50
%
)
-
probably,
chaffinch
was
a
nesting
species
on
Curonian
spit
(north),
had
the
big
population
density
and
many
the
differentiated
types
of
species
specific
song
(
22
song
types
in
sample),
therefore
elements
of
a
trill
could
be
various
under
the
form
and
quantitative
parameters
;
while
in
populations
of
the
center
Europe
an
Russia
can
be
many
migrants,
flying
groups
of
chaffinch,
which
adapt
in
many
respects
to
local
song
cultures
of
nesting
chaffinch
and
have
poorly
differentiated
under
the
form
and
time
-
and
-
frequency
parameters
elements
(syllables)
of
a
trill
(especially
2
-
nd
phrases
at
songs
with
a
complex
(difficult)
trill
on
structure)
;
•
i
ntervals
between
syllables
in
started
singing
(
sec
),
intervals
between
syllables
of
a
trill
(
1
phrase)
(
sec
),
intervals
between
syllables
of
a
final
stroke
(
sec
)
is
average
vary
(factor
of
variability
(V)
10
-
30
%
)
in
northern
population
,
but
is
strongly
vary
(factor
of
variability
(V)
33
-
50
%
)
in
the
center
Europe
an
Russia
it
is
possible,
nesting
chaffinch
better
kept
of
song
tradition
in
the
north
(
Curonian
spit
)
and
the
given
attributes
(signs)
of
species
specific
song
were
more
stable
at
variability,
than
among
the
majority
of
flying
chaffinch
(migrants)
of
the
center
Europe
an
Russia
.
Table 8.
Results of comparison of parameters of different song types of chaffinch
(
Fringilla coelebs L
.
) in populations of the European Russia at individual and geographical variability
Median (average
) frequency
, KHz
number
1
1
3
2
4
2
% *
0,9
2,5
5
1,7
3,17
3,6
Number of syllables
number
19
5
8
8
13
18
3
2
18
7
3
6
%
18,27
12,5
13,3
15,4
17,6
15,1
10,7
10
14,3
12,73
18,75
14,3
Parameters
of songs
(
quantity
of distinctions
)
Song type (
token
)
А
G
C
E
I
B
Ind
.
1
variab
.
Geog. 2
variab.
Ind
.
1
variab
.
Geog 2
variab.
Ind.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind.
1
var
.
Geog 2
variab.
Duration
of syllables
,
s
ес:
started singing
number
1
2
2
5
13
2
1
8
4
1
2
%
0,9
3,3
3,8
6,8
10,9
7,14
5
6,35
7,27
6,25
4,8
trill
1
phrase
number
5
4
4
2
2
6
3
2
1
%
4,8
10
6,7
3,85
2,7
5,04
10,7
10
1,8
2
phrase
number
7
6
8
12
5
%
11,7
11,5
10,8
10,08
11,9
final stroke
number
15
4
5
2
1
4
3
15
5
3
%
14,4
10
8,3
3,8
1,35
3,36
15
11,9
9,09
7,14
The note: % * -
percent from the common number of distinctions on all parameters of the given song type at individual (geographi
cal) variability; 1. I
ndividual variability –
variants (version
s
) of songs of one type
,
which has been recorded in different points of certain territory or in repertoir
e
of one male
; 2. G
eographical variability –
variants (version
s
) of songs of one type
,
which has been recorded in different territories; differences of parameters, a difference between which
> 0,5 К
Hz
in frequency and > 0,02 sec in length were taken into account; e
mpty cells of the table –
the data of this parameter were within the limits of an insignificant difference; values of parameters with a plenty of distinctions are allocated
(signed) by a font.
Intervals between syllables
, sec
start singing
number
3
4
7
3
2
8
5
2
% *
2,9
7,7
5,88
10,7
10
6,35
9,09
4,8
trill
1
phrase
number
10
5
4
1
2
1
3
%
9,6
12,5
6,7
1,9
7,14
5
5,45
2
phrase
number
4
5
1
2
%
7,7
4,2
6,25
4,8
final stroke
number
1
2
6
1
7
2
2
3
%
0,9
5
5,04
5
5,6
3,6
12,5
7,14
Parameters
of songs
(
quantity
of distinctions
)
Song type (
token
)
А
G
C
E
I
B
Ind
.
1
var
.
Geog. 2
variab.
Ind.
1
variab
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Ind.
1
var
.
Geog 2
variab.
Ind
.
1
var
.
Geog 2
variab.
Intervals between phrases
,
s
ес:
start singing
and trill
number
6
3
3
2
1
11
6
1
%
5,8
7,5
5
7,14
5
8,7
10,9
2,4
trill
and
final stroke
number
6
2
3
2
8
5
2
1
4
6
2
%
5,8
5
5
3,8
10,8
4,2
7,14
5
3,17
10,9
12,5
Intervals between songs
, s
ес
number
9
5
8
9
14
11
3
2
20
5
1
3
% *
8,7
12,5
13,3
17,3
18,9
9,24
10,7
10
15,87
9,09
6,25
7,14
Quantity of compared pairs of song (
n)
21
8
8
8
15
19
5
3
20
7
3
7
Quantity of parameters
distinctions
104
40
60
52
74
119
28
20
126
55
16
42
The note: % * -
percent from the common number of distinctions on all parameters of the given song type at individual (geographical) variability; differences of parameters, a difference between which
> 0,5 К
Hz
in frequency and > 0,02 sec
in length were taken into account; songs of one type from one male
had sample
> 20; empty cells of the table –
the data of this parameter were within the limits of an insignificant difference.
In table 8 distinctions of parameters were considered within the limits of each song type (the font allocated
(
distinguished)
the greatest by quantity of distinction at songs of one type). At comparison of values of different song types by the common there were distinctions for such parameters as length (duration) songs
(sec)
, number of syllables and intervals between songs
(sec)
. More individual (not for all song types) distinctions in the maximal frequency (types of songs G, C, Е, В), length
(duration)
of syllables in started singing
(song
type С), trills (types of songs G, C, B), a final stroke (song
types A, G, E, I), intervals between syllables and phrases (song
types A, Е, I, C) were characteristic.
Values of parameters
,
which entered into limits of an insignificant difference (had small distinction) –
minimal and average (median) frequency
(KHz)
, intervals between syllables (elements) of phrases
(sec)
were characteristic for songs of one type, but the marked similarities came to light at different types (having numbers song variants), measured in parallel, independently from each other. If to pay attention to distinctions of values at individual and geographical variability (tab. 8) it is possible to note their different parity (ratio) even within the limits of one song type. So, sometimes individual variability of parameters of songs of one type (which, apparently, should be the least) had a lot of distinctions, than values of the same parameters of songs at geographical variability which could and not have at all distinctions (measurement of song types from the different local populations removed from each other frequently on the big distances) –
for example, at types of songs A and G –
length (duration) of a stroke
(sec) , the maximal frequency (KHz) –
at types of songs Е, I, G, intervals between a trill and a stroke (sec) –
at song type C. N
evertheless
,
song
types
in most cases distinctions or coincided at individual and geographical variability, or prevailed at geographical variability.
Thus, the majority of distinctions has been found in values of length (duration) of songs, numbers of syllables and intervals between songs of one type (it has come to light for all song types
,
which have been analy
z
ed), the least distinctions –
minimal, average (median) frequency
(KHz)
, intervals between syllables and phrases
(sec)
. Other parameters in the distinctions were individual for each type. Geographical and individual variability had no precise parity (ratio) in distinctions (songs of one type in one territory could differ more, than songs of the same type in different local populations), but presence of distinctions on the certain parameters frequently coincided.
Conclusions
•
As a result of statistical calculations it has been found, that the average size of chaffinch
repertoir
e
(
Fringilla coelebs L
.
) in different areas of research made approximately 2 song
types.
In different populations song culture of chaffinch can be characterized by different song
types and their different percentage parity
(ratio) –
for example, those song types
,
which were in the majority in one population, could make small percent in another population
–
and, some song types were as dialect forms (modified in the form of elements on sonogramms
). •
The “
c
ombined" types of songs
(consisting of phrases of different song types) could be as individual in one population (were considered as «a cultural mutation»), and more widespread in other local populations, describing song
culture
in the greater degree
.
•
Chaffinc
h
songs (
Fringilla coelebs L
.
)
of
different types
could be stable in frequency
(KHz)
, but more individual (to differ) in values of length
(duration)
of syllables (elements)
(sec)
and intervals between them
(sec)
. •
Songs of one type
less differed on values minimal, average (median) frequencies
(KHz)
, intervals between syllables and phrases
(sec)
. The greatest quantity of distinctions has fallen to such parameters of songs of one type
as length (duration) of songs
(sec)
, number of syllables, intervals between songs
(sec)
. Geographical and individual variability could coincide on presence of distinctions in the same parameters of songs of one type, but had no conformity in a percentage parity (ratio).
+
•
The quantitative analysis of songs –
one of complex (difficult), relative, but accessible and
precise ways of the description individual and population
structures of song organization of different
bird
species
.
Автор
Olesya Astakhova
Документ
Категория
Зоология
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Теги
song, chaffinch, analysis, quantitative
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