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The comparative analysis of chaffinch song in geographical populations of the East Europe

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The comparative analysis of chaffinch song (
Fringilla coelebs
L.) in geographical populations of the East Europe
Song type А Song type
В
Song type
C
Astakhov
a
O.A. (dissertation) 2008.
Object
of research
Family
of
Finches
(Fringillidae
)
-
small
and
average
birds
on
the
size,
concerning
(relate)
to
group
of
Sparrow
(Passeriformes)
.
A
constitution
is
dense,
a
beak
is
strong,
conic,
rather
massive
.
They
eat
in
basically
seeds
and
fruits
of
plants,
but
almost
everyone
of
them
bring
up
(carry
)
insects
to
nestlings
.
At
the
majority
of
species,
nest
on
trees
or
bushes
in
woods
and
light
forests,
but
the
some
of
them
-
nest
on
the
ground,
in
high
mountains
or
stony
steppes
and
deserts
.
At
many
Finches
(Fringillidae)
the
song
have
in
end
a
characteristic
buzzing
sound
.
Are
very
popular
as
cellular
singing
birds
.
In
fauna
of
Russia
the
family
is
submitted
by
12
genus
and
35
species
(
Byome
,
etc
.
,
1998
;
Hrabriy
,
2006
;
Ryabitsev
,
2000
)
.
Genus
of
Finches
(Fringilla
)
-
wood
(forest)
birds
in
size
as
a
sparrow,
wings
are
black
or
dark
-
brown
with
two
white
strips
.
They
eat
usually
on
the
ground,
sing
on
trees
and
bushes
.
The
nest
-
accurate
construction
of
straw,
branches
of
trees
,
a
green
moss
-
mask
(hide)
outside
by
slices
of
lichens
or
barks
,
the
tray
(inside)
is
covered
by
feathers
and
a
wool
.
The
nest
there
is
on
trees
.
In
the
autumn
and
in
the
winter
they
meet
sometimes
as
huge
flights
.
In
Russia
there
are
2
species
-
chaffinch
(
Fringilla
coelebs
L
.
)
a
nd
brambling
(
Fringilla
montifringilla
)
(fig
.
1
)
.
At
brambling
a
head,
a
back,
wings
and
a
tail
are
black
(
male
),
and
a
throat,
above
of
tail
,
strips
on
a
wing
and
a
belly
(bottom
part)
-
white,
a
breast
-
orange
(at
female
is
less
bright)
.
The
bird
of
passage
(migrate)
.
Occupies
(inhabit)
a
southern
half
of
forest
-
tundra
and
a
wood
(forest)
zone
from
Kol
skiy
peninsula
up
to
coast
of
Ohotsk
iy
sea
.
A
song
of
brambling
-
huge
creaking
chirrup
,
having
in
end
sharp
"
chzhzhzh
",
a
call
(shout,
a
voice)
-
lingering
"
kevk
",
in
flight
-
sharp
"
chi
-
chi
"
or
"
vzhi
"
(fig
.
2
E
)
(
Byome
,
etc
.
,
1998
;
Hrabriy
,
2006
)
.
Fig. 13. Representatives of one group or genus of Finches (Fringilla):
A –
chaffinch
(
Fringilla coelebs
L.), B
–
Brambling (
Fringilla montifringilla
); 1 –
female
, 2 –
male
.
Chaffinch
(
Fringilla
coelebs
L
.
)
i
n
size
as
a
sparrow
(fig
.
1
)
.
At
male
top
of
a
head
is
grey
-
dark
blue,
a
back
is
chestnut,
a
forehead
is
black,
the
bottom
party
(side)
of
a
body
is
red
-
brown,
above
of
tail
is
greenish
.
Wings
and
a
tail
are
black
-
brown,
strips
on
each
side
a
tail
and
on
shoulders
are
white
.
Female
and
young
birds
are
brown
-
grey
.
The
bird
of
passage
(migrate)
.
Occupies
(inhabit)
wood
(forest)
and
forest
-
steppe
zones
from
the
western
borders
of
Russia
up
to
lake
Baikal
.
The
nest
builds
on
a
tree
.
In
a
laying
of
4
-
7
bluish
-
green
eggs
with
specks
and
strips
.
One
of
the
most
numerous
birds
in
woods
(forest)
and
parks
of
the
European
part
of
Russia,
also
nesting
on
Caucasus
and
in
the
south
of
Siberia
to
the
east
up
to
area
of
lake
Baikal,
it
is
usual
in
foothill
woods
(forest)
of
Caucasus
in
the
winter
.
In
some
places
it
is
possible
to
hear
some
tens
of
singing
birds
simultaneously
in
the
spring
.
A
loud
song
(a
sonorous
trill
with
the
sharp
termination
(ending)
-
a
stroke
)
-
"
fyu
-
fyu
-
fyu
-
lya
-
lya
-
lya
-
di
-
di
-
di
-
vi
-
chiu
"
(fig
.
2
)
.
Voice
(
call
,
shout)
-
sonorous
"
pink
-
pink
"
(it
is
frequently
heard
in
flight)
and
loud
ryumin
"
ryu
-
ryu
"
in
cloudy
or
cool
weather
(at
birds
from
Caucasus
and
from
Karelia
-
short
whistle)
.
At
alarm
and
occurrence
of
danger
-
a
sonorous
call
"
hyut
-
hyut
"
.
White
strips
on
wings
(a
characteristic
attribute)
are
well
visible
(
Byome
,
etc
.
,
1998
;
Hrabriy
,
2006
;
Tougarinov
,
Buturlin,
1911
)
.
Representatives
of
group
(order)
Sparrow
(Passeriformes
)
have
a
complex
(difficult)
throat
and
are
capable
to
utter
(sing)
sound
signals
-
as
simple
on
structure
(call,
shouts,
squeak
),
and
complex
(difficult)
songs
(set
of
small
sound
elements)
.
Males
s
ing
more
often
in
the
spring
-
involve
(attract)
of
females
of
their
species
in
the
borrowed
(occupied)
or
nested
sites
and
warn
others
males
,
that
the
territory
is
borrowed
(occupied)
.
Sparrow
-
youngish
(from
the
point
of
view
of
paleontology)
group
(order
),
which
is
in
process
of
active
formation
of
new
species
.
Recently
arisen
species
frequently
a
little
differ
from
each
other,
therefore
definition
of
many
sparrow
birds
in
the
nature
is
rather
difficult
.
Quite
often
close
species
of
Sparrow
differ
on
a
voice
(song)
is
better,
than
on
appearance
(
Byome
,
etc
.
,
1998
)
.
Within
the
limits
of
one
species
of
chaffinch
(
Fringilla
coelebs
L
.
)
(determined,
probably,
first
of
all
by
genetic
criterion
)
allocate
(distinguish)
13
subspecies
on
an
area
of
distribution
of
a
species
(Dementyev,
1954
)
.
The
given
subspecies
of
chaffinch
can
be
geographical
races
(the
remote
populations),
but,
nevertheless,
having
big
zones
of
contact
at
the
migrations,
overlapped
,
cooperating
(contact
)
,
dividing
(sharing)
the
general
(common),
similar,
but
also
in
many
respects
differing,
original
(distinguish)
species
specific
attributes
(signs)
.
Their
variability
can
be
shown
in
a
variation
of
character
of
the
general
(common)
color
of
feathers
,
in
its(her)
intensity,
in
a
variation
of
the
general
(common)
sizes
of
a
body,
in
the
form
and
the
size
of
a
beak
(Dementyev,
1954
)
.
Usually
subspecies
of
chaffinch
e
uropean
(
Fringilla
coelebs
L
.
)
poorly
differ
on
color
of
feathers
(distinctions
only
in
its(her)
intensity)
(Dementyev,
1954
)
and
in
song
cultures
-
in
ways
of
singing
of
a
species
specific
vocal
pattern
(a
characteristic
trill
and
a
final
stroke
in
the
end)
(
Tougarinov
,
Buturlin,
1911
)
,
while
different
species
of
chaffinch
can
have
various
color
of
covers
(for
example,
blue
chaffinch
-
Fringilla
teydea
,
on
Canary
islands)
and
the
distinguished
(different)
vocal
traditions
,
expressing
in
other
vocal
structure
of
species
specific
song
(different
ways
of
its(her)
differentiation
on
a
part,
the
phrases,
distinguished
forms
of
elements
on
sonogram
at
a
pronunciation
of
their
sound
s
)
(fig
.
2
,
3
,
4
)
(Bergmann,
1993
)
.
Fig. 4
. Blue chaffinch
(
Fringilla teydea
) on Canary islands (Tenerife, Grand Canary) (Bergmann, 1993).
Fig.
2
. Sonograms of
songs of representatives of different species
of
one group or genus of Finches
(Fringilla) and subspecies of chaffinch
(
Fringilla coelebs
L.), recorded in different points of the Earth
: A –
England, B –
Northern Africa, C, D
–
Canary
islands, E
–
Sweden (Thorpe, 1958).
Fig. 3. Subspecies of chaffinch
(
Fringilla coelebs
L.) on Canary islands (northwest of Africa) –
chaffinch
(
Fringilla coelebs tintillon
) (Bergmann, 1993).
Material and methods
of researches
It
is
known,
that
the
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
has
significant
individual
variability
(distinctions
between
individuals
and
at
singing
of
one
male
)
and
geographical
variability
(between
individuals
or
males
on
distance
-
in
the
remote
territories)
.
Vocal
variability
of
chaffinch
is
reached
(achieved)
by
"mistakes"
of
copying
at
song
training
(learning
)
in
sensitive
(perceiving)
period
of
young
males
(the
first
year
of
a
life)
by
the
senior
(older)
generation,
and
also
at
improvisations
at
singing
(change
of
phonetics
or
a
pronunciation
of
elements
of
song,
rearrangement
of
parts
of
song,
their
combination
from
different
songs
of
others
males
),
but
in
more
advanced
age
of
males
,
types
of
songs
are
stabilized
and
do
not
change
.
Females
of
chaffinch
have
species
specific
song
,
which
is
poorly
structured
,
probably,
as
a
result
of
uselessness
of
their
vocal
realization
,
and
economy
of
energy
for
other
vital
functions
(Thorpe,
1958
;
Marler,
1956
;
Nottebohm,
1967
)
.
Probably,
as
a
result
of
vocal
variability
as
in
vocal
repertoir
e
of
one
male
of
chaffinch
there
can
be
1
-
6
(
10
)
types
(different
variants
)
of
species
specific
song
(but
frequently
sing
only
2
-
3
types
of
songs),
and
in
a
population
of
chaffinch
as
a
whole,
there
can
be
on
the
average
20
different
types
of
songs
,
distinguished
(different)
in
the
form
of
elements
(syllables)
and
in
the
general
(common)
structure
(in
character
and
sequence
of
parts
or
phrases
of
songs)
on
sonograms
(a
graphic
representation
of
figure
(drawing)
of
song
structures)
(a
number
(line
,
row
)
of
males
in
a
population
can
execute
(performance)
at
singing
the
same
(similar)
song
typ
e
,
but
with
individual
specific
features
)
.
Thus
the
species
specific
song
of
chaffinch
has
to
some
extent
genetic
determination
(inherited
are
the
rhythm,
a
frequency
range,
duration
of
song),
therefore
the
differentiated
song
types
of
chaffinch
frequently
have
the
general
(common)
basis
(similarity)
at
variability
(Catchpole,
Slater,
1995
;
Kroodsma
et
al
.
,
1982
)
.
Thus,
song
training
or
learning
(copying
of
songs
from
surrounding
individuals
of
the
species
)
at
males
of
chaffinch
can
occur
far
from
a
place
of
nesting
,
where
was
born,
as
a
result
of
a
local
territorial
dispersion
(movings)
-
natal
(
postnesting
)
dispersions,
when
young
male
for
the
first
time
begins
flights
on
small
distances
within
the
limits
of
a
local
population
,
and
in
the
next
years
lives
after
the
big
migrations
,
at
which
male
could
appear
and
in
other
local
and
geographical
populations
,
remote
from
his
(its)
"native
land
"
on
100
-
1000
km
(Sokolov,
1980
;
Espmark
et
al
.
,
1989
)
.
Therefore
frequently
males
of
chaffinch
in
first
year
of
their
lives
have
the
plastic,
astable
species
specific
songs
,
capable
to
change
and
form
different
song
variants
-
types
of
songs
.
(Marler,
1952
;
Slater,
1981
;
Bergmann,
1993
)
.
As
males
of
chaffinch
are
usually
numerous
in
local
populations
,
can
form
"communicative"
groups
at
antiphonal
singing
or
countersinging
("
roll
call
"
of
males
frequently
by
the
general
(common)
types
of
songs)
.
Thus
there
is
a
transfer
of
species
specific
song
at
training
(
learning
)
from
the
senior
(older)
generation
to
younger
and,
thus,
the
vocal
tradition
of
a
species
is
supported
in
aggregate
types
(variants)
of
song,
which
are
components
of
song
culture
.
Thus
young
males
of
one
year
of
birth
also
can
"train"
(learn
)
at
countersinging
each
other
by
different
types
of
species
specific
song
,
exchanging
by
them
,
probably,
altering
(change)
of
them
.
Therefore
in
each
local
population
of
chaffinch
(in
the
certain
territory
about
1
-
5
к
m
2
)
can
develop
(to
be
formed)
special
local
song
(vocal)
culture
-
distinguished
(different)
ways
of
singing
or
a
pronunciation
of
different
types
of
songs,
their
elements
.
Such
local
cultures
of
songs
at
chaffinch
can
differ
in
different
local
populations
and
consist
of
an
original
(distinguish)
set
and
a
parity
(ratio)
(
%
)
of
the
similar
or
dialect
(modified
)
types
of
species
specific
song
.
As
a
result
of
it,
probably,
different,
remote
on
the
big
distances
,
groups
of
males
at
chaffinch
(for
example,
geographical
populations
of
a
species
)
can
have
considerably
distinguished
(different)
song
(vocal)
cultures
(set
or
complex
of
types
of
songs),
but
keeping
(preserve)
general
(common)
species
specific
criteria
of
a
vocal
pattern
(Marler,
1952
;
Thielcke,
1969
;
Slater
et
al
.
,
1984
;
Simkin,
1983
,
1988
;
Bergmann,
1993
;
Mundinger,
1982
)
.
During
the
spring
-
and
-
summer
period
2005
-
2007
,
s
amples
of
songs
of
chaffinch
(
Fringilla
coelebs
L
.
)
have
been
made
in
different
geographical
populations
of
the
East
Europe
:
in
northwest
of
the
European
Russia
(
Curonian
spit
,
the
Kaliningrad
region)
(N=
158
of
males
),
in
the
center
of
the
European
Russia
(Moscow,
Zvenigorod
,
Michurinsk)
(N=
65
of
males
)
and
in
the
south
of
Ukraine
(Crimea
)
(N=
26
of
males
),
remote
from
each
other
on
1000
-
1850
km
.
And,
in
the
south
of
Ukraine
(Crimea
)
allocate
(distinguish)
a
separate
subspecies
-
chaffinch
c
rimean
(
Fringilla
coelebs
solomkoi
Menz
.
)
,
while
in
territory
of
the
European
Russia
inhabit
the
subspecies
of
chaffinch
e
uropean
(
Fringilla
coelebs
coelebs
L
.
)
.
Subspecies
of
chaffinch
can
not
differ
on
color
of
feathers
,
but
there
can
be
a
difference
in
its(her)
intensity
(Dementyev,
1954
)
.
Character
of
species
specific
song
of
chaffinch
is
capable
to
change,
but
on
the
general
(common)
species
specific
basis
(Bergmann,
1993
)
.
Tape
recorder
record
of
songs
of
chaffinch
males
(from
everyone
on
the
average
on
20
songs)
was
carried
out
(was
spent)
with
the
help
of
compact
tape
recorder
Panasonic
RQ
-
SX
95
F
and
with
an
additional
microphone
with
the
purpose
of
increase
in
quality
of
a
sound
.
The
analysis
of
sonograms
of
song
types
of
chaffinch
was
carried
out(was
spent)
with
the
help
of
computer
program
Avisoft
-
SASLab
Light
.
H
as
been
seen
about
7500
songs
,
on
different
quantitative
(time
-
and
-
frequency)
parameters
is
measured
N=
424
songs
of
different
types
.
Statistical
processing
of
data
was
carried
out
(was
spent)
with
the
help
of
programs
Microsoft
Excel,
Statistics
6
.
0
.
Different
types
of
songs
were
designated
by
latin
letters
.
At
one
type
consider
songs
of
chaffinch
,
completely
similar
by
all
three
or
two
making
parts
(
started
singing,
a
trill,
a
final
stroke
),
which
also
could
be
subdivided
into
separate
phrases
or
parts
(fluctuations,
a
knee),
consisting
of
repeating
similar
elements
(syllables)
.
At
the
analysis
of
sonograms
of
chaffinch
song
basically
applied
two
qualitative
methods
:
revealing
of
phonetic
distinctions
(frequency
of
a
sound,
his(its)
form
(figure)
on
sonograms
)
or
in
ways
of
a
pronunciation
of
syllables
of
the
phrases
,
mak
ing
songs
;
revealing
of
lexical
distinctions
(changes
of
phrases
or
parts
of
songs,
as
a
whole)
(Mundinger,
1982
)
.
Results
and discussio
n
In
different
geographical
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
(fig
.
5
)
have
been
found
the
general
(
common)
similar
types
of
species
specific
song
,
it
is
especial
in
more
geographically
close
populations
of
the
north
and
the
center
of
the
European
Russia
(distance
between
which
1000
km)
,
where
inhabit
the
general
(common)
subspecies
of
chaffinch
e
uropean
(
Fringilla
coelebs
coelebs
L
.
)
.
In
samples
on
Curonian
spit
of
Baltic
sea
(the
Kaliningrad
region)
(N=
158
of
males
)
we
allocate
(distinguish)
22
types
of
songs
:
A
B
C
D
E
F
G
H
J
I
K
L
M
N
O
P
Q
R
S
T
U
V
(where
each
song
type
is
marked
by
the
latin
letter)
(fig
.
6
)
.
At
record,
songs
of
one
type
met
in
different
points
of
territory
(was
considered,
that
belong
to
repertoir
e
s
of
different
males
),
therefore
alongside
(equally)
with
the
letter
were
designated
by
numbers
in
ascending
order
(for
example,
А
1
,
А
2
,
А
3
,
etc
.
)
.
In
populations
of
the
central
part
of
the
European
Russia
(N=
65
of
males
)
15
different
types
of
songs
are
found
(A
B
C
D
F
G
I
J
M
N
S
U
V
W
Y)
(fig
.
7
)
,
from
which
12
types
of
songs
were
similar
in
the
structure
to
analogous
types
on
Curonian
spit
,
but
frequently
had
the
modified
elements
on
sonograms
,
therefore
were
considered
as
dialect
forms
(B
C
D
F
G
I
J
M
N
S
U
V)
(Astakhov
a
,
Byome
,
2007
)
(tab
.
1
)
.
Local
variants
of
vocalizations
of
birds
can
be
considered
as
analogy
of
human
speech
and
to
name
"dialects"
(Mundinger,
1980
,
1982
)
.
It
has
not
been
found
analogues
of
six
song
types
(H
O
Е
Q
R
T),
and
song
types
W,
Y
appeared
new
in
relation
to
known
to
us
song
patterns
on
Curonian
spit
.
In
the
south
of
Ukraine
(Crimea
),
where
allocated
(distinguish)
the
separate
subspecies
-
chaffinch
c
rimean
(
Fringilla
coelebs
solomkoi
Menz
.
)
,
in
sample
of
songs
(N=
26
of
males
)
have
been
revealed
16
different
types
of
species
specific
song
:
B
BF
GC
E
O
T
V
Y
MI
(N)
MI
(B)
ME
(I)
OI
M
(O)
I
CM
CME
CB
(are
named
on
phrases
(parts)
or
as
a
whole
of
those
song
types
in
the
European
Russia
,
which
"reminded"
in
the
structure
(the
form
of
elements)
(fig
.
8
)
.
From
them
only
one
song
type
(В)
was
completely
similar
(general
(common))
with
populations
of
the
European
Russia,
and
the
majority
of
them
-
15
types
of
songs
(
93
,
75
%
)
appeared
modified
(dialect
)
on
the
basis
of
the
general(common
)
(similar)
species
specific
vocal
structure
in
relation
to
similar
(analogous)
song
types
of
individuals
of
subspecies
chaffinch
e
uropean
(
Fringilla
coelebs
coelebs
L
.
)
in
populations
of
the
European
Russia
.
Probably,
such
distinction
of
song
cultures
(
their
ways
of
singing
of
species
specific
song)
of
chaffinch
(
Fringilla
coelebs
L
.
)
i
n
geographical
populations
of
the
East
Europe
can
confirm
occurrence
of
his(its)
separate
subspecies
(tab
.
1
)
.
It is necessary to note exact distance between geographical populations
of chaffinch (
Fringilla coelebs
L.): Crimea (Simferopol, Janko
y
) –
Moscow (1500 km), Moscow –
the south of Ukraine (Melitopol) (1250 km), Crimea (Simferopol, Jankoy) –
Curonian spit
(the Kaliningrad region) (1850 km), Curonian spit
–
Moscow (
Zvenigirod
) (1000 km), Moscow –
Michurinsk (the Tambov region) (850 km), Michurinsk –
Melitopol (the Zaporozhye region) (820 km).
Fig. 5. A map of the East Europe
: red points designate places of taking of sample of chaffinch songs (
Fringilla coelebs
L.).
To have more precise representation about different song types of chaffinch
, which we discuss, it is possible to show
sonograms
of their samples (fig. 6).
Fig. 6. The most widespread song types of chaffinch (
Fringilla coelebs
L.) in a population of northwest of the European Russia
(
Curonian spit of Baltic sea): 1 –
song type A, 2 –
song type C, 3 -
song type C*, 4 –
song type F, 5 –
song type I, 6 –
song type М1.
2.
3.
4
.
5.
6.
1.
Fig. 6. The most widespread song types of chaffinch (
Fringilla coelebs
L.) in a population of northwest of the European Russia
(
Curoniab spit of Baltic sea): 7 –
song type М12, 8 –
song type O, 9 –
song type R, 10 –
song type S, 11 –
song type J.
7
.
8.
9.
10.
11.
1
.
2.
3.
4
.
5.
6.
Fig. 7. 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, 7 –
song type М
12, 8 –
song type O, 9 –
song type R, 10 –
song type S,
11 –
song type J. Some song types of chaffinch (
Fringilla coelebs
L.) in the south of Ukraine (Crimea)
(fig. 8)
:
Type of song
В6* (
Melitopol
, Zaporozhye region
)
Type of song
В6 (
Melitopol
, Zaporozhye region)
Type of song B
* (
BF
) 2 (
Melitopol
, Zaporozhye region)
Type of song
СМ 2
(
Melitopol
, Zaporozhye region
)
Type of song
CM
6 (
Simferopol
,
Crimean
region
)
Type of song
MI
(
N
) 3
(
Melitopol
, Zaporozhye region
)
Type of song
MI (
B
) 8
(
Simferopol
, Crimean
region
)
Type of song
MI
(
N
) 6
(
Jankoi
,
Crimean
region
)
Type of song
M
Е(
I
) 6
(
Simferopol
, Crimean
region
)
Type of song
M
Е(
I
) 9
(
Simferopol
, Crimean
region
)
Type of song
О (
Jankoi
,
Crimean
region
)
Type of song
О2
(
Simferopol
, Crimean
region
)
Type of song
OI
(
Simferopol
, Crimean
region
)
Type of song
OI 2
(
Simferopol
, Crimean
region)
Type of song
OI* 3 (
Simferopol
, Crimean
region)
Type of song
М(
O
)
I
(
Simferopol
, Crimean
region
)
Type of song
Е2 (
Simferopol
, Crimean
region
)
Type of song
Т2
(
Jankoi
,
Crimean
region
)
Type of song
GC
3
(Simferopol
, Crimean
region
)
Type of song
GC
* 2 (
Simferopol
, Crimean
region
)
Table 1. A degree of a generality (similarity) of different song types of chaffinch (
Fringilla coelebs
L.)
in geographical populations of east part of Europe.
Geographi -
cal populations
Song types (sign by letter)
А
B
C
D
E
F
G
I
J
M
N
O
R
H
S
T
U
V
W
Y
BF
CM
GC
OE
Northwest of the European Russia (Curonian spit)
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
+
o
o
o
o
o
o
The center of the European Russia (Moscow,
Zvenigorod, Michurinsk)
+
+
±
+
±
±
o
×
±
±
±
±
±
±
o
o
o
±
*
±
*
±
±
+
+
+
+
+
+
The south of Ukraine (Crimea)
*
+
*
o
×
o
±
o
o
o
×
o
o
×
o
*
±
*
o
o
o
±
o
±
o
±
±
±
±
±
*
The
note
:
+
-
presence
of
song
type,
which
can
be
the
general
(common)
or
similar
to
analogous
song
type
in
other
geographical
population
;
±
-
the
modified
(
dialect
)
song
type
in
relation
to
similar
song
type
in
other
geographical
population
;
*
-
poorly
similar
(
original
)
song
type,
having
only
insignificant
similarities
in
the
form
of
elements
with
analogous
song
type
in
other
geographical
population
;
×
-
the song type is absent, but has separate similar phrases
(parts) in other allocated (different) types of songs of the given geographical population;
o
-
the
song
type
is
absent
in
the
given
geographical
population
.
The cells of the table containing two symbols
designate, that in the given geographical population the certain song type can be different character
(in relation to other geographical population), i.e. different variants
of song type (or to the full
(or partly
) it (he) at all is absent -
if sign ×
, o) are found.
Originally
sample of chaffinch songs has been taken on Curonian spit, therefore comparison of types of songs was conducted first of all wi
th a reference point on the first geographical population. The general (common), similar (or poorly modified) song types of chaffinch in all three geographical populations are marked b
y a
fat font
(letter). Original song types (not having analogues) of chaffinch (their letters) are designated by italics
.
In the south of Ukraine (Crimea) allocate (distinguish) other subspecies -
Chaffinch Crimean (
Fringilla coelebs solomkoi
Menz.), while in northwest and in the center of the European Russia the subspecies -
Chaffinch European (
Fringilla coelebs coelebs
Linn.) inhabit; species specific song of chaffinch considerably differ at different subspecies in the typological structure (song culture), in the general (common) song structu
re (in different geographical populations many the modified or "dialect" types of songs, their separate parts (phrases); also meet original
(poorly similar) song types, but there are also general
(common), similar types of songs in different geographical populations.
Similar modification of vocal cultures of chaffinch
in his(its) different subspecies
-
chaffinch
c
rimean
(
Fringilla coelebs solomkoi
Menz.) and chaffinch
e
uropean
(
Fringilla coelebs coelebs
L.), can speak about species specific song
of birds as an exponent
of general (common)
and separate (different)
of their geographical populations
, and as criteria of the species specific status
of many species
of birds
(tab. 2).
Table 2.
Degree of a generality of chaffinch songs types (
Fringilla coelebs
L.) in different geographical populations of Europe
Geographical
population
Character of variability
song types
Percentage parity (ratio) of song types in populations (%)
Northwest of the European Russia (Curonian spit)
The center of the European Russia (Moscow, Zvenigorod, Michurinsk)
The south of Ukraine
(Crimea)
Not modified
(shared)
25,9*
3,7**
29,2
4,2**
6,25
6,25*
Subdialects
2
25,9*
14,8**
29,2
25**
25
37,5*
Dialects
1
18,5*
22,2**
20,8
20,8**
37,5
31,5*
Not having similar
3
29,7*
59,3**
20,8
50**
31,25
24,75*
The note
:
dialects
1
-
different phonetic norms of song types; subdialects
2
-
small phonetic distinctions of song types; not having similar
3
-
percent of special types of songs, not having analogues and determining a degree of an originality (allogeneity or alien) of song culture of one population in relation to anothers (last line); * -
percent of the dialect (modified) or n
ot modified
(shared, common) types of songs in the given population in relation to song types in the center of the European Russia; ** -
percent of the dialect (modified) or n
ot modified
(shared, common) types of songs in the given population in relation to song types in the south of Ukraine (Crimea); without an asterisk (*) -
percent of the dialect (modified) or n
ot modified
(shared or common) types of songs in the given population in relation to song types of northwest of the European Russia; the greatest values (30 -
50 %) are allocated (distinguished) by a font.
Proceeding from table 2
, it is possible to make generalization:
1
.
Subdialects
of
song
types
of
chaffinch
(Fringilla
coelebs
L
.
)
in
the
greater
degree
are
found
in
those
geographical
populations
,
which
are
in
closer
communications
(connections)
(in
this
case,
types
of
songs
more
close
appeared
in
populations
of
northwest
and
the
center
of
the
European
Russia
,
and
also
between
populations
of
the
south
of
Ukraine
(Crimea)
and
the
center
of
the
European
Russia
)
.
2
.
Dialect
forms
of
chaffinch
songs
(
Fringilla
coelebs
L
.
)
is
more
in
a
population
in
the
south
of
Ukraine
(Crimea
)
(in
relation
to
northwest
and
the
center
of
the
European
Russia),
that
can
confirm
smaller
interrelation
of
these
populations
(on
Crimea
and
the
European
Russia)
.
3
.
Originality
(allogeneity
or
alien,
distinguish)
of
song
cultures
is
the
greatest
at
those
populations,
which
have
less
communications
(connections
)
among
themselves
(are
most
remote
from
each
other)
:
a
southern
population
of
chaffinch
(south
of
Ukraine
(Crimea))
-
in
relation
to
northwest
(Curonian
spit),
northwest
population
-
in
relation
to
central
in
the
European
Russia
,
central
population
of
chaffinch
-
in
relation
to
southern
population
(as
depends
on
a
direction
of
migratory
ways
of
chaffinch
(
Fringilla
coelebs
L
.
)
from
one
population
in
(to)
another)
.
At comparison of song
repertoir
e of chaffinch
(
Fringilla coelebs
L.) in different geographical populations
of the East Europe (at sample on average 20 songs from everyone
of male
), have come to light
(revealed)
the following results (tab. 3, fig. 9, 10): the most widespread size of repertoir
e
of chaffinch
-
2 types of songs (57,7 %), 1 and 3 types of songs
in repertoire of chaffinch
appeared almost equal in a percentage parity
(ratio) -
19,6 % and 16,6 % accordingly, the males
met seldom
with big song
repertoir
e
s -
4 types of songs
(3,1 % in a population), with 5 types of songs
(1 %), with 6 types of songs
(2,1 %). And, in different geographical populations
of chaffinch
(
Fringilla coelebs
L.) d
istinction
was observed in a percentage parity
(ratio) of the sizes of song
repertoir
e
(by tab. 3, fig. 10). In the south of Ukraine (Crimea
) was less
song repertoir
e
s of the minimal and maximal size
, but was more
enough
volumetric
(big)
repertoir
e
s from 3 and 4 types of songs
, than in other geographical populations (fig. 10). While in northern population
of chaffinch
(
Curonian spit of Baltic sea) repertoir
e
s of the maximal sizes
have been found
(though and met
seldom
) -
5, 6 and even 10 types of songs
(at singing of one male), basically reached
(achieved) by song improvisation
(change of combinations of phrases, their rearrangement, inclusion of the dialect or modified parts of songs) (fig. 10).
Table 3. A percentage parity (ratio) of the size of song
repertoir
e of
chaffinch
(
Fringilla coelebs
L.) in different geographical populations of the East Europe
Geographical population
Quantity of song
types in repertoire 1
2
3
4
5
6
Northern population (
Curonian spit, the Kaliningrad region)
n
=7
24
,1%
n
=
16
55,2%
n
=
3
10,3%
n
=
1
3,4
%
n
=2
6
,
9
%
The center or the European Russia (
Moscow, Zvenigorod, Michurinsk
)
n
=11
23
,
4
%
n
=30
63
,
8
%
n
=5
10
,
6
%
n
=1
2
,1%
Southern population
(
the south of Ukraine, Crimea
)
n
=1
4
,
8
%
n
=10
47
,
6
%
n
=8
38
,1%
n
=2
9
,
5
%
The general (common)
for
populations of a species
n
=19
19
,
6
%
n
=56
57,7
%
n
=16
16
,
6
%
n
=3
3
,1%
n
=1
1
%
n
=2
2
,1%
The note
: in the table the quantity
(amount)
of males
is specified with the certain
(determined)
size of
song repertoir
e
(n) and their percent from the general
(common) number of song
types in sample in the certain
(determined)
geographical population; repertoir
e
s of those chaffinch males were taken into account
, from which it has been recorded on average n=20 songs; on the literary data the maximal size of repertoir
e
of chaffinch
-
six types of species specific song, but also
the big
repertoire
s
size
of chaffinch
could meet (
t
o
10
-
11 types of songs)
,
achieved by song
improvisations and combinations of phrases (parts) of different song
types (
it was on Curonian spit of Baltic sea).
Fig. 9
. The diagram of a degree of distribution of the different sizes of song repertoires of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe.
Fig. 10. Schedules of different degree of distribution of the different sizes of chaffinch repertoire (
Fringilla coelebs
L.) in three geographical populations of the East Europe.
Parameters of song types
Geographical population
Average values of parameters of song types Duration of song
, sec
Min frequency
, KHz
Max
frequency
,
KHz
Average
(median)
frequency
, KHz
Degree of distribution of song type, %
Number of syllables in song
Duration of started singing, sec
Duration of trill, sec
Duration of final stroke, sec
Intervals between songs, sec Northwest of the European Russia (Curonian spit)
1,97
±
0,23
1,63
±
0,063
7,683
±
0,42
4,10
±
0,14
6,82
±
5
15,6
±
2,67
0,47
±
0,26
0,931
±
0,414
0,4
±
0,1
6,92
±
0,81
The center of the European Russia (Moscow, Zvenigorod, Michurinsk)
2,59
±
0,29
1,7
±
0,17
8,06
±
0,58
4,096
±
0,29
6,83
±
6,62
20,11
±
4,6
0,88
±
0,34
0,99
±
0,32
0,62
±
0,24
7,3
±
1,9
The south of Ukraine
(Crimea)
2,26
±
0,55
1,85
±
0,14
7,98
±
0,74
4,63
±
0,23
6,6
±
5
17,5
±
4,86
0,72
±
0,2
1,02
±
0,56
0,34
±
0,1
6,53
±
1,68
Total
(the common for song of species)
2,27
±
0,36
1,73
±
0,12
7,91
±
0,58
4,28
±
0,22
6,75
±
5,54
17,74
±
4,04
0,69
±
0,27
0,98
±
0,43
0,45
±
0,15
6,92
±
1,46
The note
: distance between geographical populations of chaffinch (
Fringilla coelebs
L.) on the average 1000 -
1500 km
; in total it is measured songs of different types (N)
: from samples in population of Curonian spit (the Kaliningrad region) -
279
, in populations of the center of the European Russia -
73
songs of different types, from samples in the south of Ukraine (Crimea) -
85
of songs; differences of parameters, a difference between which >0,5 К
Hz
in frequency and >0,3 sec
in length of song parts (
are allocated (distinguished) by a font
) were taken into account.
Table 4. Comparison of basic parameters of song types of chaffinch (
Fringilla coelebs
L.)
in different geographical populations of the East Europe.
At generalization
of quantitative measurements of parameters of different song types
at chaffinch,
has come to light, that in the south of Ukraine (Crimea
) (N=172
of
songs), more southern in relation to populations of the European Russia, average (median) frequency (К
Hz
) of
song types represents median=4,63
±
0,23 К
Hz
, that is much higher
, than at song
types of chaffinch
in northwest
(
Curonian spit
, the Kaliningrad region) (N=279 of songs) median=4,10
±
0,4 К
Hz
and in the center of the European
Russia
(Moscow, Zvenigorod
, Michurinsk) (N=73
of
songs)) median=4,096
±
0,29 К
Hz
, that it is possible to explain
by warmer
climate
in the south
and raised
(increased) song
activity
of chaffinch
(
Fringilla
coelebs
L.) as a result of surplus
(supply)
of a forage
(food)
and a normal metabolism in an organism, not requiring
in economy of forces (energy
) at song
realizations
(tab. 4). Interesting
there
were
similarities
of
average
prevalence
of
one
song
type
in
each
of
geographical
populations
of
the
East
Europe
(in
their
different
vocal
culture
s,
each
of
which
represents
the
certain
set
and
a
parity
(ratio)
of
song
types
in
a
local
population)
:
in
northwest
(
Curonian
spit
,
the
Kaliningrad
region)
6
,
82
±
5
%
,
in
the
center
of
the
European
Russia
(Moscow,
Michurinsk)
6
,
83
±
6
,
62
%
and
in
the
south
of
Ukraine
(Crimea
)
6
,
6
±
5
%
(tab
.
4
)
.
But
distinguished
(widely
varying
in
values)
time
-
and
-
frequency
parameters
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
in
three
geographical
populations
of
the
East
Europe
appeared
:
length
of
songs
(
sec
),
number
of
syllables
(elements)
in
songs
,
intervals
between
songs
(
sec
)
.
At
individual
variability
(tab
.
5
)
of
songs
of
one
type
at
chaffinch
is
observed
also
similar
distinction
of
these
time
-
and
-
frequency
parameters
.
Probably
,
individual
level
of
song
organizations
of
chaffinch
is
reflected
(repeat
)
on
population
level
of
species
specific
song
(tab
.
4
)
.
Fig. 1
1
. A degree of similarity of different chaffinch song types (
Fringilla coelebs
L.) under the basic forms of elements (syllables) in populations of European Russia
Thus, chaffinch
(
Fringilla coelebs coelebs
L.) has more similarities
(shared
)
of song types
in geographical populations of the European Russia
(fig. 11). On dendrogram
(Statistics 6.0)
groups of close
song
types
are allocated (distinguished) as
"relationship
(relatives
)
", as the most similar
among themselves
and, probably, they mak
e
(form) certain vocal lines of development
(at transition
of one song
type into another
song
type from one generation to next
generation, keeping (preserving
) structure of each type by vocal tradition
, but also altering
(change
)
it(him)) (fig. 11). The given types of songs can represent the result
of differentiation (modification
) of one initial ("
ancestor
") song
type
(in figure 11 are vocal lines
of chaffinch
(
Fringilla coelebs coelebs
L.) -
types of songs is close on similarity -
U I A
, V M F D
, T S E B
, W N H G R
, J O C
).
Fig. 12
. A percentage parity (ratio) of different chaffinch song types (
Fringilla coelebs
L.) in populations of northwest, the center of the European Russia and in the south of Ukraine (Crimea):
without an asterisk (*) -
types of songs from samples of northwest of the European Russia (Curonian spit, the Kaliningrad region); * -
types of songs from populations of the center of the European Russia (Zvenigorod, Moscow, Michurinsk); ** -
types of songs from populations in the south of Ukraine (Crimea) ; the types of songs designated by three letters (MI (B), MI (N), ME (A), M (O) I –
were considered as consisting of phrases known song types in populations of the European Russia) as concern to samples in the south of Ukraine (Crimea).
However, it is possible to compare more or less the general (common) (similar) types of songs at chaffinch
(
Fringilla coelebs
L.) from all three researched geographical populations of the East Europe
,
even if they represent different subspecies of chaffinch
(fig. 12). It speaks not only about difference of song
cultures
at
chaffinch
, but also about their many similarities
, about the uniform initial beginning
in development of these vocal (
song
) cultures
,
which represent
parts of one uniform species specific vocal pattern
. In figure 12
it is possible to note a different percentage parity
(ratio) of similar types of songs in different populations
-
in one population of chaffinch
the song
type
can be strongly widespread
(distributed
), and in other population
the
same song
type
(or similar to it(him)) can be rare or average on prevalence
.
At
comparison
of
song
types
of
chaffinch
in
the
south
of
Ukraine
(Crimea),
where
allocate
(distinguish)
subspecies
is
chaffinch
c
rimean
(
Fringilla
coelebs
solomkoi
Menz
.
),
in
relation
to
song
types
in
populations
of
the
European
Russia,
where
the
subspecies
is
chaffinch
e
uropean
inhabit
(
Fringilla
coelebs
coelebs
L
.
),
has
come
to
light
(reveal)
their
full
dialect
(
modification
of
all
song
types
)
.
Song
dialect
of
birds
is
a
variant
of
the
traditional
form
of
song
,
divided
(shared)
by
members
of
a
local
population
of
birds
and
forming
borders
of
a
dialect
(separating
from
other
variant
of
song
pattern)
,
within
the
limits
of
which
there
is
a
traditional
training
(learning
)
of
song
components
,
characteristic
for
this
population
.
But
thus
big
song
repertoir
e
s
of
some
species
of
sparrow
interfere
to
objective
definition
of
dialect
borders
(Kreutzer,
1974
;
Kroodsma,
1974
)
.
The
phenomenon
of
vocal
variability
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
in
geographical
populations
of
the
East
Europe
can
be
tracked
(revealed)
on
many
examples
of
different
types
of
songs
.
Transition
of
one
song
type
in
(to)
other
song
type
in
vocal
culture
of
chaffinch
population
meets
enough
frequently
.
S
imilar
cases
are
e
specially
interesting
between
different
vocal
lines
at
chaffinch,
which
are
made
similar
,
but
in
a
different
degree
remote
«in
relationship
(as
relatives)»
from
initial
"
ancestor
"
song
pattern
,
which
once
at
differentiation
has
created
these
lines
of
development
of
vocalizations
(fig
.
11
,
16
,
18
)
.
Regional
variability
of
songs
at
birds
can
be
determined
in
qualitative
aspect
(the
form
of
a
syllable,
also
syntax
or
sequence
of
phrases
or
song
parts
,
elements)
and
in
quantitative
(time
-
and
-
frequency)
parameters
.
The
concept
of
a
dialect
should
be
closely
(cautiously
)
and
is
carefully
argued
at
establishment
of
regional
variability
of
signal
and
song
patterns
.
The process of a modification of song type Y
is seen (show) in figure 13
. Between populations of chaffinch
in Michurinsk
(the Tambov region) and Simferopol
(the Crimean region)
distance of 1070 km
. It is interesting, that the structure of song
type Y
(fig. 13) has remained similar as a whole
, but nevertheless elements of started singing
(
the row of whistle
elements) and a final stroke
have considerably changed
. Elements of a trill
of this song type
have changed not so strongly
in comparison with elements of started singing and a final stroke. Probably, here stability of elements of a trill
of one
type of song
is expressed at their individual and microgeographical variability
, revealed as a result of
the
quantitative analysis
. Together with it, elements of started singing
and a final stroke
of one type
were less stable
in the quantitative (time
-
and
-
frequency) parameters
, frequently differed
(had a wide range of a variation at similarity of elements), therefore, probably, they and are subjected in the greater degree
to
modifications (than a trill
) at formation of dialect forms
of songs of one type
. Fig. 13. Geographical variants of song type Y of chaffinch
(
Fringilla coelebs
L.):
1 –
song type Y 2 (record in Simferopol, the Crimean region –
the south of Ukraine), 2 –
song type Y 2 (record in Michurinsk, the Tambov region –
the center of the European Russia).
One more remarkable example shows, how is unpredictable
, but also naturally the typological organization of chaffinch song is constructed
within the limits of all species specific area (fig. 14). The
song type Е
has been found only in northern population of chaffinch
(
Curonian spit of
Baltic sea) (fig. 14.3), but in samples of closer population of the center of the European Russia
(Moscow, Zvenigorod
, Michurinsk) has not been found out
. While in the southern population
of chaffinch (Crimea)
,
most remote on 1850 km
,
the
song type Е meets
, but in the dialect (modified) kind
(fig. 14.1). And the reduced form of song
type Е
(fig. 14.2) is more similar
to earlier found song
type Е in the north
.
Fig. 14. The dialect (modified) forms of song type Е of chaffinch
(
Fringilla coelebs
L.):
1 –
song type Е (Simferopol, the Crimean region), 2 –
song type Е* (the reduced form) (record in Simferopol, the Crimean region), 3 –
song type Е 2 (
Curonian spit
, the Kaliningrad region).
How there was the process of a modification of song type Е
(fig. 14)? Probably, in the south of Ukraine (Crimea
) other final stroke
simply
"
increased (
add)
"
at
song
improvisation
and was fixed in such kind
in a population
? Or these two forms
of song
type Е
in parallel developed
in local
song
cultures
(local populations) from initial ("
ancestor
") song
type
,
which once
was in a population of chaffinch
, which had
not yet so widely open borders of a species specific area
? But it only assumptions.
The song type V
(fig. 15) is found as in northwest (
Curonian spit
), in the center of the European Russia (Moscow, Michurinsk), and in the south of Ukraine (Crimea). But all these geographical variants of song
type V are dialect
(are modified
), as seen
on sonograms
. A percentage parity
(ratio) of song
type V (among all types of songs) in each of different geographical populations also miscellaneous
(different)
: in the north (
Curonian spit
) -
1,1 %, in the central part (Moscow, Michurinsk) -
2,15 %, in the south (Crimea) -
3,27 %, but everywhere the song type V appeared rare
(on occurrence in samples) (fig. 12). Fig. 15. Geographical variants of song type V of chaffinch
(
Fringilla coelebs
L.):
1 –
song type V (record in Zvenigorod
, the Moscow region), 2 –
song type V (Simferopol, the Crimean region), 3 –
song type V 1 (record on Curonian spit
, the Kaliningrad region).
On sonograms
the song
type Т
is not changed almost
(fig. 16) in the south of Ukraine
(
in
Janko
y
and Melitopol
) (fig. 16.3, 16.4)
,
though these local populations divide 150 km
. In comparison with variants of song
type Т
in populations of the
European Russia
(fig. 16.1, 16.2) is changes in the form of syllables of a trill
(in the south are more simplified) and in a final
stroke
,
which also appeared more simple
in the south. The
song type ТS
(fig. 16.2) -
in the central part of the European
Russia
so is named, as is dialect
in relation to song
type Т on Curonian spit of Baltic sea
and consist of a trill
,
similar to a trill
of song
type S
(fig. 16.5).
Fig. 16. The dialect (modified) forms of song type Т of chaffinch (
Fringilla coelebs
L.) (last sonogram only for comparison):
1 –
song type Т 4 (record on Curonian spit
, the Kaliningrad region), 2 –
song type TS (record in Michurinsk, the Tambov region –
the center of the European Russia), 3 –
song type Т (record in Melitopol, the Zaporozhye region –
the south of Ukraine), 4 –
song type Т 2 (record
to Jankoi, the Crimean region), 5 –
song type S 11 (
Curonian spit
, northwest of the European Russia).
The song type O
(fig. 17) -
one of the shortest song types
of chaffinch
(
Fringilla coelebs
L.), has not been found
in populations of the center of the European Russia
, but in the north
w
est of the European Russia
(
Curonian spit of Baltic sea) was one of the widespread types of songs
(9,84 %). And this song
type
represented the modified (dialect) form
in
populations of the south of Ukraine
(Crimea)
and was not
numerous
(3,27 %). Fig. 17. Geographical variants of song type O of chaffinch
(
Fringilla coelebs
L.):
1 –
song type O (record to Jankoy, the Crimean region), 2 –
song type О*2 (the reduced form of type songs OI) (Simferopol, the Crimean region), 3 –
song type O 14 (record on Curonian spit
, the Kaliningrad region).
It is necessary to note distance between geographical populations
of chaffinch
(
Fringilla coelebs
L.): Crimea (Simferopol, Janko
y
) –
Moscow (1500 km), Moscow –
the south of Ukraine (Melitopol) (1250 km), Crimea (Simferopol, Jankoy) –
Curonian spit (the Kaliningrad region) (1850 km), Curonian spit
–
Moscow (
Zvenigorod
) (1000 km), Moscow –
Michurinsk (the Tambov region) (850 km), Michurinsk –
Melitopol (the Zaporozhye region) (820 km).
Fig. 18. Geographical variants of song type C of chaffinch
(
Fringilla coelebs
L.) at a combination to individual vocal variability (last sonogram
only for comparison):
1 –
song type CM 3 (record in Melitopol, the Zaporozhye region –
the south of Ukraine), 2 –
song type CM 5 (record in Simferopo
l, the Crimean region), 3 –
song type CM 2 (Michurinsk, the Tambov region –
the center of the European Russia), 4 –
song type С
0
4 (the reduced form) (record in Zvenigorod
, the Moscow region), 5 –
song type С*11 (
Curonian spit
, the Kaliningrad region), 6 –
song type C 3 (Curonian spit, the Kaliningrad region –
northwest of the European Russia), 7 –
song type C* (Moscow), 8 –
song type M 12 (
Curonian spit
, the Kaliningrad region).
Transition
of
one
song
type
in
(to)
other
song
type
in
vocal
culture
of
chaffinch
population
meets
enough
frequently
.
Similar
cases
between
different
vocal
lines
of
chaffinch,
which
are
made
similar
,
but
in
a
different
degree
remote
‘’
in
relationship
(as
relatives)
’’
from
initial
"
ancestor
"
song
pattern
,
which
once
at
differentiation
has
created
these
lines
of
development
of
vocalizations
(fig
.
11
,
16
,
18
)
are
especially
interesting
.
The
phenomenon
of
existence
in
one
local
population
of
chaffinch
two
and
more
dialect
forms
of
one
song
type
is
interesting
,
and
they
have
,
probably,
initially
arisen
(were
differentiated
from
uniform
ancestor
type
)
in
different
local
song
cultures
(
in
different
populations),
but
at
migration
and
mixing
of
vocal
traditions
at
chaffinch
,
appeared
together
(can
be,
there,
where
earlier
were
executed
(
sing
)
and
now
execut
e
(performance)
at
singing
of
males
th
is
ancestor
song
type
,
from
which
there
were
formed
data
,
already
independent
,
song
patterns
)
(fig
.
14
,
17
,
18
)
.
At
the
quantitative
analysis
of
geographical
variants
of
song
type
C
(fig
.
18
,
tab
.
5
),
has
come
to
light
(reveal)
,
that
the
greatest
quantity
(amount)
of
distinctions
contains
in
such
time
-
and
-
frequency
parameters
,
as
:
length
(duration)
of
songs
(
sec
),
the
maximal
frequency
of
songs
(К
Hz
),
length
of
syllables
(elements)
of
started
singing
(
sec
)
and
intervals
between
songs
(
sec
)
.
But,
nevertheless,
length
of
songs
(
sec
)
and
intervals
between
songs
(
sec
)
form
the
certain
group
of
close
values
,
varying
on
average
degree
in
comparison
with
other
parameters
of
songs,
for
example,
elements
of
started
singing
(
sec
)
and
the
maximal
frequency
of
songs
(К
Hz
),
capable
to
vary
more
widely
(tab
.
5
)
.
Table 5 on the following page
Table 5. The basic time
-
and
-
frequency parameters of geographical variants of song type C of chaffinch (
Fringilla coelebs
L.), which belong to one vocal line
of development Song type (design by latter)
Number of sons
(
n
)
Name of place
of record
Length of song, sec
Min frequency, К
Hz
Max frequency, К
Hz
Median
(average) frequency, К
Hz
Number of syllables in songs
Length of syllables in started
singing, sec
Length of syllables in trill
, sec
Length of syllables in final stroke,
sec
Intervals between songs, sec
1
phrase
2
phrase
С
M
3
(
fig
. 18.
1
)
33
Melitopol
(
Zaporozhye region
)
2,14
±
0,18
1,92
±
0,21
7,15
±
0,5
4,41
±
0,19
16,8
±
1,33
0,145
±
0,016
0,05
±
0,01
0,097
±
0,015
7,68
±
2,87
CM 5
(
fig
. 18.2)
16
Simferopol
(
Crimean region
)
1,9
±
0,26
2
±
0,18
8
±
1,04
4,44
±
0,15
14,5
±
2,1
0,11
±
0,02
0,055
±
0,03
0,11
±
0,039
6,06
±
2,49
С
M
2
(fig
. 18.
3
)
5
Michurinsk
(
Tambov region)
2,48
±
0,027
1,72
±
0,21
7,4
±
0,12
4,2
±
0,15
17
±
0
0,24
±
0,01
0,034
±
0,004
0,11
±
0,005
0,082
±
0,007
6,58
±
1,46
C
0
4
(
fig
. 18.
4
)
10
Zvenigorod
(
Moscow region
)
2,39
±
0,12
1,41
±
0,1
7,75
±
0,24
4,48
±
0,21
12,3
±
0,48
0,21
±
0,03
0,12
±
0,01
0,1
±
0,085
6,54
С*11 (
fig
. 18.5)
2
Curonian spit (
Kaliningrad region
)
2,55
±
0,082
1,3
±
0,37
7,751
±
0
4,478
±
0
19
±
1,4
0,175
±
0,007
0,034
±
0,01
0,084
±
0,001
0,0915
±
0,002
22,9
С 3 (
fig
. 18.6)
20
Curonian spit (
Kaliningrad region
)
2,074
±
0,118
1,627
±
0,131
9,698
±
0,41
4,0996
±
0,173
17,25
±
0,85
0,1203
±
0,005
0,068
±
0,003
0,1155
±
0,008
0,071
±
0,005
6,423
±
1,9
С
#
(
fig
. 18.
7
)
3
Moscow
2,73
±
0,3
1,78
±
0,1
8,096
±
0,6
4,54
±
0,43
20
±
1,73
0,22
±
0,006
0,076
±
0,003
0,12
±
0
0,081
±
0,019
8,26
The note:
average value and a standard deviation of parameters of song types are specified from statistical calculations for all songs
o
f one type, which were performance (executed) at singing by males of chaffinch in the given points of record; the strongest differences counted a difference of parameters
> 0,5 К
Hz
in frequency and > 0,02 sec
in length (are allocated
(distinguish)
by a font).
In table 5
it is necessary to note, t
hat not all song types
, which more similar on structure
(the form of elements, on ways of division into phrases or parts) can have similar time
-
and
-
frequency parameters
. On the contrary, more similar songs
in the given example
could have the big distinctions in quantitative parameters
, even if they were in the same (in one) local or geographical population
-
for example, song types С*11
(fig. 18.5) and
C
3
(fig. 18.6), recorded
on Curonian spit of Baltic sea
, or song
types C
M 3
(fig. 18.1) and C
M 5
(fig. 18.2), recorded
in Simferopol
. These examples of "related
(as relatives)
" song
types
show distinctions
as in length (
sec
), frequency (К
Hz
) of songs
, and in length of elements (
sec
)
,
parts of songs
(
in started singing
, trills, a final stroke), that speaks about modification
(
dialect
) of given song
types
,
even if in many respects their general vocal structure is kept
(preserved)
.
Why
in
some
vocal
cultures
the
initial
song
type
can
change
more
,
than
in
other
populations
–
so,
that
through
any
time
to
make
(to
form)
completely
different
dialect
forms
,
but
of
one
type
of
song,
than
that
initial
ancestor
song
type
in
other
population
?
Probably,
the
degree
of
combination
(
mixing
)
of
one
population
here
influences
,
and
a
share
(
%
)
of
birds
-
migrants
in
population
,
therefore
,
as
a
result,
different
song
cultures
of
chaffinch
can
strongly
mix
up
,
and
initial
song
types
can
change
to
new
types
,
than
in
other
populations
,
which
are
less
mixed
by
representatives
of
different
local
and
geographical
populations
,
where
more
or
less
supported
stability
of
vocal
culture
,
ways
of
execution
(performance)
at
singing
of
species
specific
song,
and
where
there
is
a
smaller
modification
of
song
types
,
which
for
many
song
types,
already
changed
and
differentiated
,
in
other
populations
can
be
initial,
ancestor
and
are
capable
to
exist
equally
(together)
with
them
(Espmark
et
al
.;
Nelson
et
al
.
,
1995
;
Kroodsma,
1996
)
.
Results of the quantitative analysis
of different songs types in three geographical populations
of chaffinch
(
Fringilla
coelebs
L.) in the East Europe
in the greater degree are submitted as schedules
(diagrams)
, but on the basis of tables
.
The combination of individual and geographical variability
of different song
types
at
chaffinch
(
Fringilla coelebs
L.) in populations of the East Europe
represents special
(distinguished, different)
character
for everyone
separate song
type
(tab. 6, fig. 19).
Table 6. A degree of distinction of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe at individual and geographical variability
Character of variability
Song types (designed by latter)
А
В
С
E
G
I
ME(A)
GC
BF
MI(N)
MI(B)
CM
OI
Individual
5,48
5,7
6,57
5,6
7,5
6,3
6,67
5
5
7,67
Microgeographical
5,71
6
6,26
6,67
5,78
7,86
4,4
1
6,67
Geographical
6,25
7,39
6,79
8,33
10,5
6
The note
: individual variability
-
variants (version) of songs of one type, recorded in different points of the certain one territory
or in repertoir of one males; geographical variability
-
variants (version) of songs of one type, recorded in different territories (remote on 1000
-
1800 km
); microgeographical variability
of songs of one type was defined (determined) in the territories remote on the average on 100 km
; a degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs song types (average value from different populations is shown); the more these values -
the more this or that variability of song types; the greatest difference of value
s (> 1,5) different kinds of variability of the certain song types is allocated (distinguished) by a fat font.
Thus, different types of songs
of chaffinch (
Fringilla coelebs
L.) are capable to vary
differently, with specific features
(to change in the parameters) on the big and smaller distances
between populations. Some song types
of chaffinch can have the big individual variability
(within the limits of one population) of time
-
and
-
frequency parameters, but smaller their microgeographical variability
(on distance of populations on the average 100 km). Other song types
of chaffinch are capable to have smaller individual variability
(quantity of distinctions) of quantitative parameters, but big their geographical variability
(on distance of populations on 1000
-
1800 km), that is (the last fact) observed at songs of chaffinch in the greater degree
.
Fig. 19.
A degree of distinction of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe at individual and geographical variability:
a degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs of song types (average value from different populations is shown); than the more these values is the more this or that variability of types of songs.
For discovering (revealing), as far as differs from each other and varies each separate of quantitative (time
-
and
-
frequency) parameter at different song types of chaffinch
(
Fringilla coelebs
L.) at individual and geographical variability, we have made some of tables (tab. 7, 8, 9). Table 7. A general degree of distinction of the certain parameters of different song types of chaffinch (
Fringilla
coelebs
L.) in geographical populations of the East Europe at individual and geographical variability
Parameters of songs
(degree of distinctions)
Character of variability of song types
Individual variability
Microgeographical variability
Geographical variability
Length of songs, sec
0,94
0,9
2
0,91
Min
frequency
, К
Hz
0,16
0,08
0,19
Max frequency
, К
Hz
0,58
0,6
0,56
Median (average) frequency
, К
Hz
0,22
0,08
0,51
Number of syllables in songs
0,96
0,87
0,99
Length of syllables, sec
:
started singing
0,41
0,58
0,51
trill
1
phrase
0,5
0,17
0,49
2
phrase
0,44
0,43
0,66
final stroke
0,64
0,45
0,66
Intervals between syllables, sec
:
started singing
0,66
0,56
0,28
trill
1
phrase
0,33
0,19
0,3
2
phrase
0,31
0,16
0,51
final stroke
0,52
0,5
0,43
Intervals between phrases, sec
:
started singing and trill
0,43
0,23
0,46
trill and final stroke
0,48
0,37
0,45
Intervals between songs, sec
0,92
0,84
0,78
The note
: individual variability
-
variants (versions) of songs of one type, recorded in different points of the certain one territory or in repertoire of one male
; geographical variability
-
variants (versions) of songs of one type, recorded in different territories (remote on 1000
-
1800 km); microgeographical variability
of songs of one type was defined (determined) in the territories, remote on the average on 100 km; a degree of distinction of parameters of songs is defined (
det
ermined) as a parity (ratio): quantity(amount) of distinctions of parameters / quantity (amount) compared
pairs of song
types (average value from different populations is shown); than the more these values -
the more this or that variability of types of songs; the greatest difference of values (>0,3) of different kinds of variability of the certain
pa
rameter of song types is allocated (distinguish) by italics
; the fat font designates the greatest values (>0,5) in the certain kind of variability.
In
table
7
the
more
differing
time
-
and
-
frequency
parameters
of
songs
of
one
type
at
all
kinds
of
vocal
variability
of
chaffinch
(
Fringilla
coelebs
L
.
)
are
appeared
:
length
of
songs
(
sec
),
maximal
frequency
(К
Hz
),
number
of
elements
(syllables)
in
songs
,
intervals
between
songs
(
sec
),
and
also
length
(
sec
)
of
started
singing
,
a
final
stroke
and
length
(
sec
)
of
their
elements
(syllables
)
.
Thus
different
song
types
were
analyzed
,
within
the
limits
of
which
songs
of
one
type
were
allocated
(distinguished)
in
repertoir
e
s
of
several
males
of
chaffinch
in
a
population
.
Probably,
individual
variability
(within
the
limits
of
one
local
population)
of
these
parameters
of
songs
of
one
type
is
reflected
(repeat
)
on
population
level
of
chaffinch
,
in
their
geographical
variability
(in
different
local
populations),
that
is
expressed
in
similarity
of
song
differences
of
these
levels
(fig
.
20
)
.
Also
it
is
necessary
to
note,
that
in
some
time
-
and
-
frequency
parameters
of
songs
of
one
type
at
chaffinch,
individual
variability
can
exceed
geographical
variability
(that
is
differences
of
these
parameters
at
songs
of
one
type
can
be
big
between
individuals
of
chaffinch
in
one
local
population
in
territory
1
км
2
,
than
on
distance
of
100
-
1000
km)
:
length
of
syllables
of
the
first
phrase
of
a
trill
(
sec
),
intervals
between
elements
of
started
singing
and
a
final
stroke
(
sec
),
intervals
between
songs
(
sec
)
(tab
.
7
,
fig
.
20
)
.
It
is
possible
to
explain
it
that
at
an
individual
level
is
more
probability
of
display
of
occasions
(chances
)
and
a
randomness
in
values
of
parameters
of
songs,
than
on
larger
population
level
,
where
the
law
of
variability
of
song
parameters
already
is
more
shown
(revealed)
.
Figure 20 on
the following page
Fig. 20.
A degree of distinction of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe at individual and geographical variability:
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 phrase) (sec), 8 -
length of syllable
s 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 phrase) (sec), 13 -
intervals between syllables of final stroke (sec), 14 -
intervals between phrases (started singing and a trill) (sec), 15 -
intervals between phrases (a trill and final stroke) (sec), 16 -
intervals between songs (sec). A degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs of song types (average value from different populations is shown); the more these values -
the more this or that variability of song types.
Now it is interesting to consider, how parameters of chaffinch song
(
Fringilla coelebs
L.)
differ
in different geographical populations of the East Europe
at the account
(including)
of the same characters of variability
of song
types -
individual, microgeographical and geographical
(tab. 8).
In
table
8
we
notice
the
same,
that
was
and
in
table
7
-
an
individual
level
(smaller)
is
reflected
(repeat
)
on
larger
population
level
at
similarity
in
differences
of
songs
of
one
type
in
such
parameters
,
as
length
of
songs
(
sec
),
number
of
syllables
in
songs,
maximal
frequency
(К
Hz
),
intervals
between
songs
(
sec
),
length
of
syllables
(elements)
(
sec
)
and
intervals
between
them
(
sec
)
in
started
singing
and
a
final
stroke
.
But
here
come
to
light
(reveal)
features
of
vocal
variability
inside
each
of
geographical
populations
of
chaffinch
,
including
distinctions
of
character
of
song
variability
between
different
subspecies
-
chaffinch
e
uropean
(
Fringilla
coelebs
coelebs
L
.
)
and
chaffinch
c
rimean
(
Fringilla
coelebs
solomkoi
Menz
.
)
(tab
.
8
)
.
For
example,
in
the
south
of
Ukraine
(Crimea
)
individual
variability
of
length
(
sec
)
of
syllables
(elements)
and
intervals
between
them
(
sec
)
at
the
second
phrase
of
a
trill
is
less
,
than
in
the
European
Russia
.
Probably,
it
is
connected
,
that
in
the
south
of
Ukraine
(Crimea
)
song
types
of
chaffinch
are
poorly
differentiated
,
were
very
similar
with
each
other
(fig
.
8
,
18
),
therefore
parameters
of
a
trill
of
these
song
types
also
had
small
variability
.
In
general,
weak
(small)
separation
(
differentiation
)
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
o
n
separate
(different)
types
in
the
south
of
Ukraine
(Crimea
)
(fig
.
8
,
18
)
can
speak
about
"young
",
yet
not
stabilized
vocal
(
song
)
culture
of
chaffinch
,
about
his(its)
"recent",
in
the
evolutionary
plan,
settling
(inhabit)
in
Crimea,
Caucasus
.
It
is
surprising,
that
in
the
south
of
Ukraine
(Crimea
)
individual
variability
of
parameters
of
started
singing
and
a
final
stroke
is
somewhat
more
,
than
in
the
European
Russia
(tab
.
8
)
.
Probably,
the
big
variability
of
quantitative
parameters
of
started
singing
and
a
final
stroke
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
also
can
speak
about
enough
astable
(
not
established
to
the
full
)
in
the
character
of
song
culture
of
chaffinch
in
the
south
(Crimea
)
.
Also
in
table
8
it
is
possible
to
note,
that
individual
variability
of
average
(median)
frequency
(К
Hz
)
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
is
more
in
the
center
of
the
European
Russia
in
comparison
with
other
geographical
populations
.
Probably,
it
also
testifies
to
instability
(
about
structural
instability
)
of
song
(vocal)
culture
of
chaffinch
in
territory
of
East
European
plain
(fig
.
7
)
and,
probably,
it
occurs
as
a
result
of
significant
mixing
of
nesting
and
flying
(migratory)
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
a
t
his(its)
gradual
settling
(inhabit
)
nested
territory
in
the
East
Europe
.
Though,
most
likely,
in
comparison
with
difficult
and
precisely
differentiated
song
types
,
complicated
song
culture
of
chaffinch
in
the
Western
Europe
(fig
.
6
)
(
primary
,
his(its)
ancient
species
specific
area
),
both
in
the
south
of
Ukraine
(Crimea
),
and
in
the
East
Europe
is
observed
smaller
differentiation
and
clearness
of
song
types
of
chaffinch
(fig
.
7
,
8
),
their
instability
,
that
speaks
about
chaffinch
as
"recently
"
occupied
(inhabit)
species
in
the
East
Europe
,
even
if
as
one
of
numerous
.
Table 8 on the following page
Table 8. A degree of distinction of the certain parameters of song types of chaffinch (
Fringilla coelebs
L.) in different geographical populations of the East Europe at individual and geographical variability
Parameters of songs
(degree of distinctions)
Character of variability of song types in geographical populations
Northern population
(
Curonian spit
)
The center of the European Russia (Moscow, Zvenigorod, Michurinsk)
The south of Ukraine (Crimea)
Individual variability
Microgeogra -
phical variability
Individual variability
Geographical variability
Individual variability
Microgeograph.
variability
Geographical variability
Length of songs, sec
0,97
0,96
1
0,93
0,86
0,87
0,89
Min
frequency
, К
Hz
0,16
0,16
0,21
0,21
0,1
0
0,17
Max frequency
, К
Hz
0,43
0,15
0,67
0,55
0,63
0,75
0,56
Median (average) frequency
, К
Hz
0,16
0,44
0,4
0,35
0,1
0
0,67
Number of syllables in songs
0,89
0,88
1
0,97
1
0,87
1
Length of syllables, sec
:
started singing
0,26
0,49
0,4
0,58
0,57
0,67
0,44
trill
1
phrase
0,57
0,33
0,1
0,31
0,83
0
0,67
2
phrase
0,65
0,66
0,67
0,65
0
0,2
0,67
final stroke
0,56
0,4
0,5
0,31
0,87
0,5
1
Intervals between syllables, sec
:
started singing
0,61
0,44
0,71
0,4
0,67
0,67
0,17
trill
1
phrase
0,47
0,38
0,2
0,27
0,33
0
0,33
2
phrase
0,33
0,31
0,6
0,35
0
0
0,67
final stroke
0,23
0,33
0,83
0,43
0,5
0,67
0,44
Intervals between phrases, sec
:
started singing and trill
0,45
0,46
0,33
0,36
0,5
0
0,56
trill and final stroke
0,35
0,36
0,6
0,33
0,5
0,37
0,56
Intervals between songs, sec
0,76
0,67
1
0,67
1
1
0,89
The note
: individual variability
-
variants (versions) of songs of one type, recorded in different points of the certain one territory or in repertoire of one male
; geographical variability
-
variants (versions) of songs of one type, recorded in different territories (remote on 1000
-
1800 km); microgeographical variability
of songs of one type was defined (determined) in the territories, remote on the average on 100 km; a degree of distinction of parameters of songs is defined (
det
ermined) as a parity (ratio): quantity(amount) of distinctions of parameters / quantity (amount) compared
pairs of song
types (average value from different populations is shown); than the more these values -
the more this or that variability of types of songs; the greatest difference of values (>0,3) of different kinds of variability of the certain
pa
rameter of song types is allocated (distinguish) by italics
; the fat font designates the greatest values (>0,5) in the certain kind of variability.
Fig. 21.
A parity (ratio) of different kinds of vocal variability of chaffinch (
Fringilla coelebs
L.) for the certain parameters of song types in three geographical populations of the East Europe:
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 phrase) (sec), 8 -
length of syllables of a trill (2 phrase) (sec), 9 -
length of syllables of final str
oke (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 phrase) (sec), 13 -
intervals between syllables of final stroke (sec), 14 -
intervals between phrases (started singing and a trill) (sec), 15 -
intervals between phrases (a trill and final stroke) (sec), 16 -
intervals between songs (sec). Individual variability
-
variants (versions) of songs of one type, recorded in different points of the certain one territory or in repertoire of one male; geographical variability
-
variants (versions) of songs of one type, recorded in different territories (remote on 1000
-
1800 km); microgeographical variability
of songs of one type was defined (determined) in the territories, remote on the average on 100 km; equal
-
signs between kinds of variability of songs are shown: equally (=), it is more (>), it is less (<).
Character
of
variability
of
each
separate
quantitative
parameter
of
different
song
types
of
chaffinch
is
submitted
in
the
diagram
(fig
.
21
)
.
Some
parameters
of
different
types
of
songs
of
chaffinch
(
Fringilla
coelebs
L
.
)
can
be
characterized
by
the
greater
individual
variability
(within
the
limits
of
one
local
population)
and
smaller
geographical
(microgeographical)
variability
(between
populations
on
distance
of
100
-
1000
km)
.
Probably,
these
parameters
of
songs
are
more
instable
(not
stable)
in
the
values
in
populations
of
a
species
,
are
subject
to
the
greater
song
improvisation
,
change,
transformation,
though
in
different
geographical
populations
such
song
parameters
of
a
species
can
have
less
distinctions
.
Thus,
most
individually
in
changeable
parameters
of
chaffinch
song
(but
capable
to
be
geographically
similar)
are
(fig
.
21
)
:
maximal
frequency
(
К
Hz),
intervals
between
syllables
of
started
singing
(sec),
intervals
between
syllables
of
a
final
stroke
(sec),
length
of
syllables
of
a
final
stroke
(sec),
intervals
between
songs
(sec)
.
More
distinguished
at
geographical
(microgeographical)
variability
(on
distance
of
100
-
1000
km),
but
more
stable
(with
smaller
distinctions)
in
the
values
of
song
parameters
of
chaffinch
in
a
local
population
at
individual
variability
can
be
(fig
.
21
)
:
average
(median)
frequency
(
К
Hz),
length
of
syllables
of
started
singing
(sec),
length
of
syllables
of
a
trill
(
2
phrase)
(sec),
intervals
between
syllables
of
a
trill
(sec)
.
Probably,
on
these
song
parameters
of
chaffinch
the
vocal
tradition
in
the
certain
populations
of
a
species
is
established
,
and
transferred
the
subsequent
generation
at
song
learning
as
a
feature
of
a
local
dialect
.
Equal
in
individual
and
geographical
(microgeographical)
variability
of
parameters
of
chaffinch
songs
(fig
.
21
)
are
:
length
songs
(sec),
minimal
frequency
(
К
Hz),
intervals
between
phrases
(started
singing
and
a
trill)
(sec),
intervals
between
a
trill
and
a
final
stroke
(sec)
.
Probably,
these
song
parameters
of
chaffinch
can
vary
(individually
and
geographically
to
change),
but
as
a
whole
remain
stable
for
populations
of
a
species
and
define
(determine)
species
specific
song
tradition
(
a
species
specific
song
pattern
of
chaffinch
)
.
Other
song
parameters
of
chaffinch
can
have
both
individual,
and
geographical
variability
in
different
parities
(ratio)
(fig
.
21
)
.
The degree of individual and geographical variability of different parameters of species specific song
of chaffinch
can be tracked (revealed)
separately in each of geographical populations of the East Europe (fig. 22, 23), that can be generalization of the previous diagram.
Fig. 22.
Individual variability (within the limits of one certain territory) parameters of different types of songs of chaffinch (
Fringilla coelebs
L.) in populations of the East Europe:
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 phrase) (sec), 8 -
length of syllables of a trill (2 phrase) (sec), 9 -
l
ength 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 phrase) (sec), 13 -
intervals between syllables of final stroke
(sec), 14 -
intervals between phrases (started singing and a trill) (sec), 15 -
intervals between phrases (a trill and final str
oke) (sec), 16 -
intervals between songs (sec). A degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs of song types (average value from
different populations is shown); than the more these values -
the more this or that variability of song types.
Individual
variability
of
song
types
of
chaffinch
(
Fringilla
coelebs
L
.
)
shows
a
degree
(quantity)
of
distinctions
of
parameters
in
their
values
at
songs
of
one
type
within
the
limits
of
one
territory
of
the
area
1
km
2
.
On
the
schedule
average
values
of
a
degree
of
distinction
of
different
song
types
of
chaffinch
in
each
geographical
population
are
resulted
.
Differences
of
parameters,
a
difference
between
which
>
0
,
5
К
Hz
in
frequency
and
>
0
,
02
sec
in
length
were
taken
into
account
(were
considered)
.
Songs
of
one
type
from
one
male
had
sample
on
the
average
n>
20
(fig
.
22
)
.
Distinctions
of
song
parameters
at
males
of
chaffinch
in
each
population
can
be
different
by
quantity
(amount)
-
greater,
smaller,
or
equal
in
this
or
that
geographical
population
.
Probably,
individual
variability
of
parameters
of
song
types
shows
a
degree
of
combination
(mixing)
of
chaffinch
individuals
from
different
local
populations
-
as
far
as
many
migrants
in
a
population
,
and
also
can
speak
about
a
degree
of
definiteness
(stability)
of
those
or
other
parameters
of
songs
in
any
geographical
population
of
chaffinch
.
Than
the
more
individual
variability
of
parameters
of
chaffinch
song
(
Fringilla
coelebs
L
.
)
–
the
more
can
be
combination
(mixing
)
of
his
(its)
population
,
and
more
not
stable
character
of
parameters
(fig
.
22
)
.
Fig. 23.
Geographical variability (on distance on the average 1000
-
1800 km) and microgeographical variability (on distance on the average 100 km) parameters of different song types of chaffinch (
Fringilla coelebs
L.) in populations of the East Europe:
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 betwee
n syllables of started singing (sec), 11 -
intervals between syllables of a trill (1 phrase) (sec), 12 -
intervals between syllabl
es of a trill (2 phrase) (sec), 13 -
intervals between syllables of final stroke (sec), 14 -
intervals between phrases (started singin
g and a trill) (sec), 15 -
intervals between phrases (a trill and final stroke) (sec), 16 -
intervals between songs (sec). A degre
e of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs of song types (average value from different populations is shown); the more these values -
the more this or that variability of song types.
Geographical
(microgeographical)
variability
of
different
types
of
songs
of
chaffinch
(
Fringilla
coelebs
L
.
)
in
the
certain
geographical
populations
shows
a
degree
(quantity)
of
distinctions
of
different
parameters
of
song
types
of
chaffinch
individuals
at
a
great
distance
(from
100
km
up
to
1800
km)
(fig
.
23
)
.
Geographical
(microgeographical)
variability
of
parameters
of
chaffinch
songs
shows
a
degree
of
distinction
of
vocal
cultures
in
different
geographical
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
.
Also
microgeographical
variability
of
songs
(on
distance
on
the
average
100
km)
can
show
a
degree
of
change
of
song
types
of
chaffinch
in
space
within
the
limits
of
this
or
that
geographical
population
(fig
.
23
)
.
In
cases,
when
microgeographical
variability
of
song
parameters
of
chaffinch
is
more,
than
their
geographical
variability,
it
is
possible
to
speak
about
the
phenomena
of
distant
migrations
of
birds
,
their
gradual
settling
(inhabit)
of
nested
territory
and
thus
mixing
of
different
song
forms
(from
distinguished
or
different
cultural
vocal
traditions
of
a
species)
of
individuals
of
chaffinch
from
different
local
populations
(fig
.
23
)
.
Thus, it is possible to make the general (common) result on geographical and individual variability of quantitative parameters of species specific song of chaffinch
(
Fringilla coelebs
L.) (tab. 9, fig. 24).
The table 9. A general degree of distinction of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe at individual and geographical variability
Character of variability
of song types
Geographical populations of chaffinch
Northern population (
Curonian spit
)
The center of the European Russia (Moscow, Michurinsk)
The south of Ukraine (Crimea)
Individual variability
5,76
6,24
6,56
Microgeographical variability
6,4
4,02
Geographical variability
7,5
8,12
The note
: individual variability
-
variants (versions) of songs of one type, recorded in different points of the certain one territory or in repertoire of one male; geographical variability
-
variants (versions) of songs of one type, recorded in different territories (remote on 1000
-
1800 km); microgeographical variability
of songs of one type was defined (determined) in the territories, remote on the average on 100 km; a degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinc
tio
ns of parameters / quantity(amount) compared pairs of song types; than the more these values -
the more this or that variability of song types
; the fat font designates the greatest values in the certain kind of variability
.
As a result, songs of one type
at chaffinch (
Fringilla coelebs
L.) could be more similar
on the greater distance
(up to 100 km) at microgeographical variability
, than songs of the same type
, but taking place (recorded) in one local population
(in territory 1 к
m
2
) or in repertoire of one male at individual variability (
tab
. 9, fig
. 24
)
. Probably, such combination (mixing) of different song forms of one type
occurs as a result of migrations of birds
or at local territorial dispersions
(moving) of individuals, including at distribution in the postnestling period
(postnatal dispersion).
Fig. 24.
Degree of variability of song types of chaffinch (
Fringilla coelebs
L.) at different distances of individuals in geographical populations of the East Europe
: a degree of distinction of parameters of songs defined (determined) as a parity (ratio): quantity (amount) of distinctions of parameters / quantity (amount) compared pairs of song types (average value from different populations is shown); than the more these values is the more this or that variability of song types.
And, at last, comparison of a degree of a variation (variability) of time
-
and
-
frequency parameters
of species specific song of chaffinch
in different geographical populations of the East Europe
(fig. 25).
Fig. 25
.
A degree of a variation of values of parameters of different song types of chaffinch (
Fringilla coelebs
L.) in geographical populations of the East Europe:
1 –
length of song (
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 syllables in started singing
(
sec
), 7 -
length of syllables of a trill (1 phrase) (
sec
), 8 –
length of syllables of a trill (2 phrase) (
sec
), 9 –
length of syllables of a 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 phrase) (
sec
), 13 -
intervals between syllables of a final stroke (
sec
), 14 –
intervals between phrases (
started singing
and a trill) (
sec
), 15 -
intervals between phrases (a trill and a final stroke) (
sec
), 16 –
intervals between songs (
sec
).
The factor of a variation (V, %)
(fig. 25
) 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
) , 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.) inside of each from
three geographical populations of the East Europe
а)
Similarities
on
a
variation
of
values
of
quantitative
parameters
of
chaffinch
songs
(
Fringilla
coelebs
L
.
)
in
three
geographical
populations
(fig
.
25
)
:
p
oorly
varied
(factor
of
variability
(V)
3
-
9
of
%
)
in
all
three
geographical
populations
of
chaffinch
are
minimal,
maximal
and
average
(median)
frequency
(
KHz
)
(fig
.
25
)
–
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
of
%
)
in
all
three
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
)
(fig
.
25
)
–
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
of
%
)
in
all
three
geographical
populations
of
chaffinch
was
at
parameters
of
song
types
:
length
of
syllables
in
stareted
singing
(
sec
),
length
of
syllables
of
a
trill
(
1
phrase)
(
sec
),
length
of
syllables
of
a
final
stroke
(
sec
)
(fig
.
25
)
–
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)
.
b
)
D
istinctions
on
a
variation
of
values
of
quantitative
parameters
of
chaffinch
songs
(
Fringilla
coelebs
L
.
)
in
three
geographical
populations
(fig
.
25
)
:
t
he
length
of
syllables
of
a
trill
(
2
phrase)
(
sec
)
is
average
varied
(factor
of
variability
(V)
10
-
30
of
%
)
in
populations
of
the
center
and
the
south
of
the
East
Europe
,
but
in
northern
population
is
strongly
varied
(factor
of
variability
(V)
33
-
50
of
%
)
(fig
.
25
)
-
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)
(fig
.
6
)
,
therefore
elements
of
a
trill
could
be
various
under
the
form
and
quantitative
parameters
;
while
in
populations
of
the
center
and
the
south
of
the
East
Europe
(fig
.
7
,
8
)
can
be
many
migrants,
flying
groups
of
chaffinch,
which
adapt
in
many
respects
to
local
song
cultures
of
nesting
chaffinch
(
Nelson
et
al
.
,
1995
)
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)
(fig
.
25
)
;
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
of
%
)
in
northern
population
,
but
is
strongly
vary
(factor
of
variability
(V)
33
-
50
of
%
)
in
the
center
and
in
the
south
of
the
East
Europe
(fig
.
25
)
-
it
is
possible,
nesting
chaffinch
better
kept
of
song
tradition
in
the
north
(
Curonian
spit
)
(fig
.
6
)
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
and
the
south
of
the
East
Europe
(fig
.
7
,
8
)
;
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
average
vary
(factor
of
variability
(V)
10
-
30
of
%
)
in
northern
and
central
populations
of
the
European
Russia
,
but
is
strongly
vary
(factor
of
variability
(V)
33
-
50
of
%
)
in
the
south
of
Ukraine
(Crimea)
(fig
.
25
)
-
probably,
because
the
first
phrase
of
a
trill
(or
last
phrase
of
started
singing)
of
chaffinch
songs
in
many
song
types
in
the
south
could
be
in
form
of
stick
-
similar
horizontal
element
(as
"
ryu
-
ryu
(ryumin)
"
of
chaffinch
in
cloudy
or
cool
weather,
or
as
"
whistle
")
(fig
.
8
)
,
and
the
second
phrase
of
a
trill
was
equally
with
less
stable
first
phrase
of
a
trill
in
the
European
Russia
(fig
.
6
,
7
,
8
)
.
Сonclusion
Stability
of
song
culture
(set
or
complex
of
song
types)
in
a
population
of
chaffinch
(
Fringilla
coelebs
L
.
)
during
time
and
on
space
,
probably,
is
a
result
of
precise
or
certain
(determined)
"social"
traditions
of
song
learning
:
training
of
species
specific
song
by
individuals
of
chaffinch
in
the
first
years
of
a
life
at
singing
of
grown
-
ups
(older)
males
,
and
antiphonal
singing
(
"roll
call"
by
the
general
(common)
songs),
the
song
adaptation
(change,
transition
of
plastic
song
in
(to)
the
stable
form)
to
each
other
by
a
generality
of
song
types
,
by
their
similarity
in
"communicative"
groups
at
young
males
of
chaffinch
–
as
between
males,
having
of
the
same
age
,
at
individuals
after
the
first
year
of
a
life
,
and
with
grown
-
ups
(older)
males
.
Also
stability
of
species
specific
song
is
shown
as
result
of
selective
individual
training
(copying
)
of
species
specific
song
by
individuals
of
chaffinch
in
sensitive
period
at
young
age
(
the
first
year
of
a
life
),
and
also
in
this
case
the
significant
hereditary
predefiniteness
(determinancy)
of
parameters
of
species
specific
song
of
chaffinch
especially
influences
(a
frequency
range
and
a
rhythm,
general
structure
of
songs
)
by
genetic
inclinations
of
display
(expression)
of
mentality
(psychica
)
(its(her)
rudiments
at
birds),
which
is
capable
to
change
owing
to
action
of
"social"
traditions
of
song
learning
("
mistakes"
of
copying
)
and
the
individual
improvisations
in
the
species
specific
song
(
in
depending
on
"creative"
potential
)
of
individuals
of
chaffinch
at
singing
(at
realization
of
types
of
species
specific
song)
.
Song
types
(
in
th
eir
set
or
complex
)
in
one
geographical
population
of
chaffinch
(
Fringilla
coelebs
L
.
)
differ
from
set
(or
complex
)
of
different
song
types
in
another
remote
on
the
big
distances
(
1000
-
1800
km)
geographical
population
on
the
following
quantitative
(time
-
and
-
frequency)
parameters
:
length
(duration)
of
songs
(
sec
),
number
of
syllables
in
songs,
intervals
between
songs
(sec)
.
On
the
same
parameters
the
song
types
differ
among
themselves
and
at
individual
variability
(at
individuals
of
chaffinch
in
one
local
population
or
in
repertoir
e
of
one
male
)
.
So
,
it
is
possible,
that
population
level
of
the
song
organization
of
chaffinch
(
Fringilla
coelebs
L
.
)
is
reflection
(display
or
repeat
)
of
individual
level
of
his(its)
song
organization
,
because
similarity
of
distinctions
of
chaffinch
song
types
comes
to
light
(reveal)
in
different
biological
scales
.
At
different
biological
subspecies
of
chaffinch
(
Fringilla
coelebs
L
.
)
(the
geographical
populations
,
remote
from
each
other
on
the
big
distances)
"
song
cultures
"
(set
of
song
types,
their
certain
percentage
parity
(ratio))
considerably
differed
(in
the
general
(common)
structure
of
songs,
under
the
form
of
elements
on
sonograms
)
.
In
the
given
geographical
populations
of
two
subspecies
of
chaffinch
-
chaffinch
e
uropean
(
Fringilla
coelebs
coelebs
L
.
)
and
chaffinch
c
rimean
(
Fringilla
coelebs
solomkoi
Menz
.
),
w
ere
observed
(revealed)
the
smaller
quantity
(amount)
of
the
general
(common)
(similar)
types
of
songs
(even
"dialect"
or
modified)
and
a
lot
of
original
(dissimilar,
not
the
general
(common))
types
of
songs
,
which
had
no
analogues
in
the
geographical
population
,
which
are
remote
on
big
(
1500
-
1800
km)
distance
from
the
first
.
A
lot
of
"dialect"
(modified
)
and
"
subdialect"
(a
little
modified
)
in
the
structure
(the
form
of
elements)
of
general
(common)
(similar)
types
of
songs
were
appeared
in
those
local
and
geographical
(more
removed)
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
,
which
were
in
big
(closer)
communications
(connections)
among
themselves
(
by
migrations
-
fig
.
26
,
at
having
of
smaller
distances
of
an
arrangement
to
each
other)
.
In
migratory
("flying")
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
in
wide
territories
(East
European
plain)
the
effect
"cumulative"
song
cultures
was
observed
(revealed),
which
could
include
a
plenty
of
the
"combined"
types
of
songs
(consisting
of
phrases
or
parts
of
different
other
types
of
songs)
and
was
indistinctly
,
is
not
thin
(not
determined
)
expressed
in
song
structure
and
forms
of
elements
of
song
types,
that,
probably,
was
the
adaptation
to
the
greater
plasticity
(changeability)
of
types
of
songs
at
their
adaptation
to
local
song
culture
in
process
of
song
training
or
learning
(
copying
of
species
specific
song
of
chaffinch
,
local
ways
of
its(her)
singing
,
phonetic
variants
of
different
song
types
)
.
While
at
nesting
populations
of
chaffinch
(residents)
the
types
of
songs
were
characterized
by
clearness,
a
subtlety
(definiteness)
in
structure,
the
form
of
elements
(fig
.
26
)
.
Fig. 26
. The basic flying ways (migration) of birds (from S.
P.
Nayumov, 1965):
chaffinch (
Fringilla coelebs
L.), apparently, is the migrant of west
-
african
lines (on the plan
is a continuous line)
.
In
northern
population
of
chaffinch
(
Fringilla
coelebs
L
.
)
(
Curonian
spit
,
the
coast
of
Baltic
sea)
the
species
specific
song
was
characterized
by
smaller
length
(duration)
(
sec
)
and
lower
frequency
range
(К
Hz
),
than
in
a
southern
population
of
chaffinch
(Crimea
)
-
it
is
probable,
the
reason
for
this
is
the
cold
climate
of
the
north
,
capable
to
lower
a
metabolism
of
an
organism
and
compelling
to
save
(to
economy
)
of
forces
(energy)
at
realization
(execution
or
performance,
singing)
of
songs
.
A
lot
of
subdialects
(small
phonetic
changes)
of
general
(common)
song
types
of
chaffinch
is
found
in
those
populations
,
which
were
more
contacted
to
each
other
(
by
migrations
,
smaller
distances
among
themselves
(more
closely))
and
they
formed
one
(uniform)
subspecies
of
chaffinch
(
Fringilla
coelebs
coelebs
L
.
)
.
While
in
less
contacted
populations
of
chaffinch
(the
geographical
populations
,
removed
on
the
big
distances
-
1500
-
1800
km,
can
concern
or
relate
to
different
subspecies
of
chaffinch
)
w
ere
observed
(revealed)
a
lot
of
original
types
of
songs
(
few
similar
,
dialect,
not
having
analogues
)
,
determining
a
degree
of
“
a
lien"
of
song
cultures
(their
distinction)
at
given
populations
of
chaffinch
.
To
individual
and
geographical
variability
in
the
greater
degree
are
subjected
started
singing
(an
initial
part)
and
a
final
stroke
of
species
specific
song
of
chaffinch
,
than
a
trill
(an
average,
basic
part)
,
which
is
more
stable
in
quantitative
and
qualitative
parameters
.
The
big
variability
of
started
singing
and
a
final
stroke
of
chaffinch
song
(
Fringilla
coelebs
L
.
)
i
t
is
possible
to
explain
,
that
these
parts
of
species
specific
song
are
poorly
and
indistinctly
audible
at
singing
of
males
in
populations
(an
illegibility
of
structure
of
a
sonorous,
sharp
final
stroke
,
probably,
because
of
his(its)
brevity
),
therefore
at
song
training
(learning
)
and
at
singing
of
chaffinch
the
started
singing
(more
silent
)
and
a
final
stroke
(shorter
)
more
exposed
and
give
chance
to
the
greater
song
improvisations
or
to
mistakes
of
copying
as
in
the
form
of
elements
(
qualitative
aspect
),
and
time
-
and
-
frequency
parameters
.
While
the
trill
is
more
stable
in
quantitative
and
qualitative
parameters
at
individuals
of
chaffinch
in
different
geographical
populations
,
because
in
the
greater
degree
it
is
genetically
determined
(in
beat
,
the
general
(common)
song
structure
,
a
frequency
range
)
and
more
audibl
e
at
execution
(performance)
of
species
specific
song
by
males
of
chaffinch
,
as
the
trill
is
an
average
,
basic
part
of
songs
.
Singing
of
chaffinch
(
Fringilla
coelebs
L
.
)
–
the
feature
(sign)
is
parallelly
in
the
development
to
others
attributes
(signs)
of
this
species
(morphological,
genetic,
physiological,
ecological,
geographical)
though,
certainly,
in
many
respects
together
with
them
forms
a
single
whole
,
but,
nevertheless,
equally
with
these
attributes
(signs
),
a
song
of
chaffinch
also
develop
on
the
special
laws
,
is
differentiated
(changes
on
the
basis
of
initial
"
ancestors
"
song
forms
and
"relatives"
song
lines
during
time
and
on
space),
creates
separate
steady
song
norms
(set
of
the
most
widespread
song
types),
consisting
of
vocal
lines
of
similar
,
but
also
distinguished
(different
)
song
variants
of
one
type
,
forming
certain
typological
vocal
polymorphism
,
from
which
proceed
many
other
polymorphic
branches
(or
big
trunks)
of
more
distant
or
more
close
«on
relationship»
lines
of
song
culture
(
a
variety
of
song
types
,
a
manner
or
ways
of
execution
(performance)
at
singing
their
structure,
separate
elements,
phrases
)
(fig
.
27
)
.
«
Song
cultures
»
(
set
of
song
types,
their
certain
percentage
parity
(ratio
))
in
local
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
gradually
(smoothly)
pass
(transition
)
to
each
other
on
all
a
species
specific
area
:
the
"intermediate"
condition
of
song
culture
of
chaffinch
in
a
separate
local
population
can
confirm
interrelation
and
a
continuity
of
song
types
of
chaffinch
,
their
smooth
(gradual)
transition
in
the
variability
from
one
type
(
song
forms,
a
pattern)
in
(to)
another
song
type
(the
form,
a
pattern)
on
all
area
(territory
)
of
distribution
of
a
species
.
Fig. 27
. C
haracter of evolution as "
net
" (from N.
N.
Iordansk
iy
, 2001).
Any evolutionary trunk is formed by philetical (development or evolutionary
)
lines
(AE, AH, AG, AF, BF)
,
which branch out or diverge and again merge (converge); sections of philetical (development)
lines divided by «planes (flats) of time» (T1, T2) -
biological species (species specific vocalizations in particular); pieces of philetical (
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)
lines MN and PQ, submitted in the paleontologic annals, -
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535
-
570
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The song of chaffinch
–
is especially important
!
She(it) is same, as well as everything, that exists.
We sing, because we exist.
Автор
Olesya Astakhova
Документ
Категория
Зоология
Просмотров
54
Размер файла
226 093 Кб
Теги
europe, song, populations, chaffinch, geographical, comparative, analysis, east
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