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Assumptions of evolution of chaffinch song (Fringilla coelebs L.) +

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Assumptions of evolution of chaffinch song (
Fringilla coelebs
L.) at comparison with a song of other finch (
Fringilla montifringilla
)
Modification of song type C in populations of chaffinch
(
Fringilla coelebs
L.)
Astakhova O.A. (dissertation) 2008.
Fig. 1. Representatives of one group or genus of Finches (Fringilla):
A –
chaffinch
(
Fringilla coelebs
L.), B
–
b
rambling (
Fringilla montifringilla
); 1 –
female
, 2 –
male
.
Family of Finches
(Fringillidae)
B
irds
is small
and average on size. A constitution is dense, a beak is conic, rather massive. In fauna of Russia the family is submitted by 35
species
. Brambling
(
Fringilla montifringilla
) In size as
a sparrow. The back side
is
bluish
-
black, the under of tail is
white, a throat, a craw and a breast
are
bright
-
red, abdomin (
bottom
side) is white. On a wing are white and orange strips. Female
is less brightly. The bird of passage
(migrate)
. Inhabit in
a southern half of forest
-
tundra and a wood zone
(forest)
from Kol
skiy
peninsula up to coast of Ohotsk
iy sea
. The nest arranges on a tree. In a laying of 5
-
7 greenish eggs with red
-
violet specks. A voice -
sharp "
chzhee
" and a rough song "
chzhzhzh
".
Chaffinch
(
Fringilla coelebs
L.) In size as
a sparrow. Male have
grey
-
dark blue
top of a head, a back
is
chestnut, a forehead
is
black, the bottom party
(side) of a body is red
-
brown. Wings and a tail are black
-
brown, strips on each side of a tail and on shoulders are white.
Female is
буро
-
grey. The bird of passage
(migrate)
. Occupies 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. A voice -
sonorous "
pink
-
pink
" and a loud song -
a trill "
fyu
-
fyu
-
fyu
-
lya
-
lya
-
lya
-
di
-
di
-
di
-
vi
-
chiu
". White strips on wings (a characteristic attribute) are well visible.
Both species
(
chaffinch
and brambling
) are well defined
(determined) on characteristic color of feathers
(
Hrabriy
, 2006).
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).
General characteristic of chaffinch singing (
Fringilla coelebs
L.
)
•
Species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
-
one
of
most
clear
examples
of
display
(expression)
of
geographical
and
individual
vocal
variability
,
though
at
many
other
species
of
birds
(Aves)
distinction
of
geographical
and
individual
patterns
of
species
specific
song
also
can
be
distinctly
revealed,
especially
on
sonograms
(
graphic
representations
of
sound
elements
),
even
if
vocalizations
of
some
species
of
birds
are
completely
genetically
hereditary
(Charadriiformes,Corvidae,
Anseriformes,
etc
.
)
(Miller,
1996
;
Bretagnolle,
1996
)
.
But
chaffinch
(
Fringilla
coelebs
L
.
)
belong
(relation)
to
species
of
birds
from
group
of
Sparrow
(Passeriformes
)
and
famil
y
of
Finches
(Fringillidae
)
,
which
have
social
traditions
of
song
training
(learning
),
which
are
passing
(transition)
from
father
to
son
(from
one
generation
to
next)
and
capable
to
change
(Catchpole,
Slater,
1995
)
.
•
Species
s
pecific
song
of
chaffinch
is
hereditary
fixed
in
such
features
as
certain
frequency
range
(К
Hz
),
duration
of
song
pattern
(
sec
),
intervals
(
sec
)
between
these
song
types
,
repeating
one
after
another
at
singing
of
male
,
and
also
the
general
song
structure
is
inherited
(character
of
division
into
elements,
a
rhythm
of
song
)
.
Therefore
at
exit
from
eggs
in
a
nest
and
at
activation
by
songs
of
parents
,
young
male
is
capable
to
form
at
first
(begin)
song
pattern
,
which
homogeneous
in
elements
(as
well
as
at
all
young
individuals
of
chaffinch
in
first
year
of
their
life
),
and
then,
perceiving
the
own
singing
and
singing
of
surrounding
adult
males
(not
only
of
their
parents),
complicates
the
song
in
forms
of
elements,
in
the
general(common)
structure
-
the
song
of
male
gets
the
form
of
a
stable
species
specific
vocal
pattern
(usually
consisting
of
three
parts
or
phrases
-
started
singing,
a
trill,
a
final
stroke
),
characteristic
for
all
adult
males
of
chaffinch
.
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
)
.
•
Thus
song
training
or
learning
(copying
of
songs
from
surrounding
individuals
of
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
of
lives
after
the
big
migrations
,
at
result
of
which
the
male
could
appear
and
also
in
other
local
and
geographical
populations
,
remote
from
his(its)
"native
land
"
on
100
-
1000
km
(the
Sokolov,
1980
;
Espmark
et
al
.
,
1989
)
.
Therefore
frequently
males
of
chaffinch
in
first
year
of
their
life
have
the
plastic,
astable
species
specific
songs
,
capable
to
change
and
form
different
song
variants
-
types
of
songs
.
In
repertoire
of
one
male
there
can
be
1
-
6
(
10
)
types
of
songs
,
but
frequently
sing
only
2
-
3
types
of
songs
.
And
in
a
local
population
of
chaffinch
(
Fringilla
coelebs
L
.
)
i
t
is
possible
to
meet
on
the
average
20
different
types
of
songs
.
A
number
(line
or
row
)
of
males
in
a
population
can
execute
(performance)
at
singing
the
same
(one)
song
type
,
but
with
individual
specific
features
(Marler,
1952
;
Slater,
1981
;
Bergmann,
1993
)
.
•
It
is
known,
that
the
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
has
significant
individual
variability
(distinctions
between
individuals
and
at
singing
one
male
)
and
geographica
l
(between
individuals
or
males
on
distance
-
in
the
remote
territories)
variability
.
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)
from
the
senior
(older)
generation,
and
also
at
improvisations
at
singing
(change
of
phonetics
or
a
pronunciation
of
song
elements,
rearrangement
of
song
parts,
their
combination
(mix
up)
from
different
songs
of
other
males
),
but
types
of
songs
are
stabilized
and
do
not
change
in
more
advanced
age
of
males
.
•
Appearing
as
a
result
of
migrations
in
different
populations
,
males
of
chaffinch
at
singing
"adapt"
to
each
other
,
copying
from
each
other
songs
of
different
types
,
characteristic
for
that
local
or
geographical
population
,
in
which
appeared
.
At
countersinging
(
"roll
call"
of
males
usual
ly
by
the
general
(common)
songs
in
"communicative"
groups)
adult
individuals
of
chaffinch
form
(create)
the
final
ly
,
fixed,
stable
forms
of
species
specific
song
in
their
song
repertoir
e
s
(the
period
of
song
crystallizatio
n
)
.
At
some
species
of
birds
of
this
period
of
fastening
(stabilization)
of
species
specific
song
does
not
occur,
and
individuals
are
capable
to
alter
(change)
song
forms
by
song
improvisation
during
all
life
(Slater
et
al
.
,
1980
;
Jellis,
1977
;
Kroodsma,
1996
)
.
•
Thus,
in
different
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
mix
ing
of
set
(plural)
different
types
(variants)
of
species
specific
song
occurs
,
and
that
song
types
also
can
change
(modificate)
by
males
at
song
improvisations,
at
"mistakes"
of
copying
-
to
create
new
variants
or
types
of
songs
(distinguished
or
different
in
the
form
of
elements,
but,
nevertheless,
having
of
similar
general
(common)
features)
.
Further,
these
song
types
of
chaffinch
in
a
population
are
transferred
at
song
training
(learning)
to
younger
generation
,
which,
in
turn,
having
adopted
and
having
altered
those
types
of
songs
,
will
transfer
their
to
following
generation
,
probably,
already
in
other
populations
,
remote
on
the
big
distances
(Catchpole,
Slater,
1995
;
Kroodsma
et
al
.
,
1982
)
.
•
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,
they
hav
e
big
zones
of
contact
at
the
migrations,
they
overlap,
cooperat
e
,
they
have
(shar
e
)
the
general
(common),
similar
species
specific
attributes
(signs)
,
but
which
also
differ,
are
original
(distinguish)
in
many
respects
.
Their
variability
can
be
shown
in
a
variation
of
character
of
the
general
(common)
color
of
feather
,
in
its(her)
i
ntensity
,
in
the
general
(common)
sizes
of
a
body
,
in
the
form
and
the
size
of
a
beak
(Dementyev,
1954
,
Stepanyan,
2003
)
.
Character
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
is
capable
to
change,
but
on
the
general
(common)
species
specific
basis
(Bergmann,
1993
)
(fig
.
2
)
.
Fig. 5. А
udience model of development of song
samples (patterns) in the ontogenesis (during life) of
singing birds (
Slater, 1983d
).
Song
training
(or learning)
of chaffinch
(
Fringilla coelebs
L.)
Fig. 6. Sonograms of
song
at
typical
wild chaffinch
(
Fringilla coelebs
L.) and song
of chaffinch,
brought up i
n isolation
(Slater, 1989).
Fig. 8. Sonograms of
two different types of songs from repertoir
e of
chaffinch male
(
Fringilla coelebs
L.); also division of song on
(into)
phrases, syllables and elements is shown
(
Slater, Ince, 1979
) . Fig. 7. Songs of young males
of chaffinch
(
Fringilla coelebs
L.), brought up in isolation
(
the letters are specified sonograms of
songs from different individuals). The frequency rank of species specific song
is kept
(preserve)
, but song structure is indistinct, poorly divided
(s
eparate)
into departments
(section, phrases), the elements of songs are badly
(weak)
differentiated
, a final stroke is absent or is distinguished hardly up (
Thorpe, 1958
).
Principles of the analysis of
structure chaffinch
song
(
Fringilla coelebs L
.)
(
our view with a support on literary references
) (Astakhova, Byome, 2007)
•
on
perception
on
hearing
the
song
was
subdivided
into
three
parts
:
the
row
(number)
of
whistle
sounds
(as
if
started
singing)
,
sounds
of
trill
(are
as
though
pored
each
other)
and
a
final
stroke
or
terminal
flourish
(the
example
of
the
analysis
is
shown
on
more
widespread
song
type
C
)
:
fuit
-
fuit
-
fuit
-
fuit
til
-
til
-
til
-
tel
-
tel
chi
-
kuik
•
Within
the
limits
of
these
parts
of
songs
on
sonogramms
it
is
possible
to
allocate
(design,
mark)
the
phrases
(the
elements
similar
under
the
form),
in
this
case
a
trill
includes
two
phrases
;
•
Elements
(syllables)
can
be
simple
(the
row
of
whistle
sounds
and
1
-
st
phrase
of
trill
in
type
C)
and
complex
(difficult),
consisting
of
two
and
more
subelements
(
2
-
nd
phrase
of
trill
in
type
C)
;
•
Syllables
(elements)
are
divided
by
intervals
,
but
frequently
shorter,
than
at
phrases
;
•
A
final
stroke
(flourish)
in
many
types
of
songs
will
consist
of
elements
(syllables),
different
under
the
form
(it
is
possible,
therefore
it
such
sharp,
remarkable,
"bright"
on
hearing)
.
Song
str
u
cture of chaffinch
(
Fringilla coelebs
L.)
Fig. 9. Songs of chaffinch
(
Fringilla coelebs
L.): (а) the typical stanzas
(strophe or line),
which have been recorded near the
Cambridge (England) in territory of nesting of chaffinch,
where experiment of song
training
or learning
was carried out
(was spent)
(Thorpe, 1958); (b) the diagrams of two typical stanzas
(strophes) of
chaffinch
-
lines represent the basic sections (departments), and circles -
elements of a final phrase (stroke) (
Marler, 1956
).
Fig. 10. Structure of chaffinch song
(
Fringilla coelebs gengleri
), living
(inhabit)
in England
(
Thorpe, 1958
). Fig. 11. Time of record and the analysis of song structure of chaffinch
(
Fringilla coelebs
L.) in Germany (
Bergmann, Helb, 1982
). Different song types of chaffinch (
Fringilla coelebs
L.) in a population
Fig. 12. Songs
-
stanzas
(strophes, lines) of chaffinch
(
Fringilla coelebs
L.). (а) and (d) from same
(one)
male
, (b) and (c) from another (second) male
. (а) and (b) -
stanzas (lines) (songs) of one type, (с) and (d) -
different types. (а), (b) and (e) have a particle is "
kit
"
(additional element
) at the end of a stanza
(strophe, line)
; (f) and (g) have variants of "
kit
"
(
additional element)
. The songs a
re recorded in different places: (а
-
d) a southwest of Germany, (е) Austria, (f, g) Dolomite (
Thielcke, 1969
).
Fig. 13. Song types of chaffinch
(
Fringilla coelebs
L.) of the center and the south of the European Russia and the Ukraine
,
which has been recorded in different
ecological places (biotops
) (
Simkin, 1983
): 1 a -
a fir forest; 1 b
-
d -
a birch wood (forest); e
-
a pine wood (forest); 1 f
-
a oak wood
(forest)
; 1
g
-
a taiga (a pine wood). Southern chaffinch
: 2 a -
oak groves (Uzhgorod), 2 b -
oak groves (Melitopol), 2 c -
Kerch, 2 d
-
Krasnodar, 2 e
, f
-
Chernovtsy (birch
-
fir forest).
1
2
1
.
2.
3.
4
.
5.
6.
Fig. 14. Most widespread song types of chaffinch (Fringilla coelebs L
.)
in populations of the central part of the European Russia (Astakhova, Byome, 2007):
1 –
song type C, 2 –
song type C
#
, 3 –
song type I, 4 –
song type J, 5 –
song type A, 6 –
song type B (different types of songs we have marked by latin letter). Variability of calls or voices (simple sounds) of chaffinch
(
Fringilla coelebs
L.)
Fig. 15. Geographical variability of a call (voice)
"
hyut
"
of chaffinch
(
Fringilla coelebs
L.) in different local populations of Europe (
Thielcke, 1969
).
Fig. 17. Different types of calls (voices) and their variability in local populations of chaffinch
(
Fringilla coelebs
L.) in researched territory in Germany; there are combinations
(mixing)
of calls
(voices) "
hyut
-
dschad
" (
Bergmann, 1993
). Fig. 16. Different types and variability of calls (voices) of chaffinch
(
Fringilla coelebs
L.) on sonograms
: d -
the West of France (1976); i -
Denmark (1980); k -
the West of Germany (1979); m -
island of Crete (1983), n -
island of Tenerife, Canary islands (
Fringilla c. tintillon
) (1989), o -
island of Las
Palmas
, Canary islands (
Fringilla c. palmae
) (1991), p -
calls (voices) "
hyut
" and
"
wrud
" (Germany), r -
rain call (
ryumin
) of chaffinch
in cloudy
or cool
weather (Germany) (1983) (
Bergmann, 1993
). Process of evolution of chaffinch song (
Fringilla coelebs
L.)
Fig. 18. Evolution of song types of chaffinch
(
Fringilla coelebs
L.) in a population
. Numbers
-
song types of chaffinch
; arrows
(pointers
) -
transfer (transmission) of song types to the following generation
(young individuals) by song
training or learning (copying
); the numbers,
which have been signed
round
by three forms of figures
(a circle, a square, a triangle) -
initial
song types
(at the senior or older individuals) in a population; figures in green triangles
-
song types of chaffinch, taught
(learnt) by posterity without mistakes and improvisations
, are kept in a population without changes
; the numbers,
which have been signed
round by dark blue circle
-
unlearned (were not coped)
types of songs
by the following generation in a population
of chaffinch
; numb
e
r
s in a red square
-
song types of chaffinch, taught
(learnt) by posterity with mistakes
(or with improvisation
at singing); numbers
in a red circle
-
the
modified song types
being a new forms of song in a population (
a cultural mutation
) (Catchpole, 1995
).
The answer by
the same (one) song
typ
e
16,8%
9,5%
Song type F
Song type BF
The answer by a song, similar on sounding (motifs)
The vocal adaptation of chaffinch males (
Fringilla coelebs
L.) in a population
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)
song
types
). Thus
,
there is a transfer of species specific songs at training (
learning
) from the senior
(older)
generation to young
individuals (next generation
)
and, thus, the vocal tradition of a species
is supported
at
aggregate (set, complex) of song
types
(variants),
as
components
of song culture
. Thus
,
young males of one year of birth
also can "train"
each other at countersinging in
different types of species specific song
, exchanging by them, probably, altering (change) of them (Nottebohm, 1967; Jellis, 1977). Scripts (scheme) of intraspecific
song
interactions of chaffinch males (
Yablonovska
-
Grishenko, 2006
)
The answer by universal song type (most widespread) 43,2%
30,5%
Song type J
Song type
В Song type
М
Opposition of very unsimilar (different) or more complex (difficult) song type
Therefore in each local population of chaffinch
(in the certain territory is abou
t
1
-
5 km
2
) can develop (to be formed) special local song
(vocal) culture
-
distinguished (different, special) ways of singing or a pronunciation of different song types (each of
them
), their elements
. Such local cultures of song
at chaffinch
can differ in different local populations
and
consist
of an original
(distinguished, special)
set (complex) 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 chaffinch males
(for example, geographical populations
of a
species
) can have considerably distinguished (different) cultures
of song
(vocal) (
set 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).
Countersinging males of chaffinch (
Fringilla coelebs
L.)
(Astakhov
a
, Byome
, 2007)
Fig. Countersinging males of chaffinch (
Fringilla coelebs
L.) by different, but by the general (common) (similar) types of songs -
three cases: A, B, C (though there is a lot of cases of countersinging by the same song types). As males of chaffinch are usually numerous in local populations, than they can form "communicative" groups at antiphon singing or countersinging (call of males frequently by the general (common) types of songs). Thus there is a transfer of species specific song at training (song learning) from the senior (older) generation to younger and, thus, the vocal tradition of the species in aggregate of song types (variants), song components is supported (kept, memory). Thus young males (of the same year of life) also can "train" at countersinging each other in different types of species specific song, exchanging by them, probably, altering (changing) them (Nottebohm, 1967; Jellis, 1977).
+
•
In
many
literary
references
in
bioacoustics
of
birds,
the
view
(themes)
is
described
about
formation
of
original
(special)
vocal
traditions
of
birds
in
their
different
local
and
geographical
populations
-
that
is,
there
is
a
distinction
between
sets
(complex)
of
different
types
of
species
specific
song
in
the
geographical
populations
,
remote
from
each
other
.
Even
if
types
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
can
differ
inside
a
geographical
population
in
aggregate
(set,
complex
)
they
are
capable
to
reflect
in
the
structure
some
features
of
a
climate,
geology
(mountains
or
plains)
,
character
of
vegetation
of
district
-
and,
thus,
to
form
special
local
culture
of
vocal
(
song
)
(ways
of
singing
of
species
specific
song),
distinguished
(different)
from
vocal
culture
of
chaffinch
in
other
geographical
population
(Thielcke,
1969
;
Slater,
Ince,
1979
;
Slater
et
al
.
,
1984
;
Simkin,
1983
;
Mundinger,
1982
)
.
•
Thus,
in
different
geographical
populations
of
chaffinch
(
Fringilla
coelebs
L
.
)
from
one
generation
to
next
generation
can
develop
(to
be
formed)
distinguished
(different)
traditions
of
vocal
(
song
)
,
probably,
giving
rise
(
beginning)
for
new
forms
(norms)
of
species
s
pecific
song
or
beginning
for
distinguished
(special)
vocalizations
of
new
species
of
birds
.
•
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)
.
Conclusions from researches of chaffinch song (
Fringilla coelebs
L.) (Astakhov
a
, Byome
, 2007)
•
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
.
•
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
)
.
•
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
"
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)
.
•
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
.
*
)
.
•
«
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
.
•
Whether
the
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
is
criterion
of
process
of
speciation
(separation)
of
species
and
a
is
characteristic
attribute
of
subspecies
and
other
formed
groups
of
birds?
Probably,
as
well
as
any
species
specific
attribute
(sign)
,
a
song
of
chaffinch
it
is
capable
to
be
steady
at
the
certain
norm
,
but
within
the
limits
of
any
norm
can
and
vary
at
individual
and
geographical
variability
of
individuals
in
populations,
at
their
special
song
cultures
in
aggregate
(which
represent
the
original
or
distinguish
set
and
a
parity
(ratio)
(
%
)
types
or
variants
of
species
specific
song)
.
Thus
it
is
necessary
to
take
into
account,
that
these
distinguished
(different)
song
cultures
of
chaffinch
in
different
populations
will
consist
of
constantly
mixing
up
types
of
songs
of
these
different
populations
,
where
they
change
or
remain
constant
,
being
transferred
to
the
following
generation
(fig
.
*)
.
Fig. *
. C
haracter of evolution as "
net
" (from N.
N.
Iordansk
iy
, 2001).
Any evolutionary trunk is formed by filetical (development or evolutionary
)
lines
(AE, AH, AG, AF, BF)
,
which branch out or diverge and again merge (converge); sections of filetical (development)
lines are divided by “area
of time
”
(T1, T2) -
biological species (
also as species
-
specific vocalizations); parts of filetical (
evolutionary
)
lines MN and PQ
represen
ted in the paleontologic
al
annals -
paleontologic
al
species
.
•
The
unity
of
the
general
(common)
and
differing
in
species
specific
song
of
chaffinch
is
same
as
and
at
any
attribute
(sign)
of
a
species
.
But
formation
of
new
song
norms
(tradition
)
within
the
limits
of
which
there
will
be
the
new
other
variation
,
which
is
distinct
from
character
of
variability
of
former
norm
of
species
specific
song
,
can
be
shown
more
on
level
of
population
and
a
speci
es
of
birds
(at
occurrence
of
separate
different
close
species
of
birds)
.
•
But
when
species
specific
populations
are
uniform
enough
and
form
a
single
whole
-
difficultly
to
speak
about
formation
of
separate
independent
song
norms
.
Most
likely,
on
all
a
species
specific
area
of
distribution
of
chaffinch
(
Fringilla
coelebs
L
.
)
there
can
be
smooth
(gradual)
transitions
from
one
vocal
norm
in
another
or
also
combination
(mix
up)
of
these
song
norms
as
a
result
of
migrations
of
birds
,
and
then
there
can
be
a
formation
of
one
or
several
combined
song
cultures
(including
cultures
of
set
or
many
),
as
occurs
in
species
specific
populations
of
chaffinch
(
Fringilla
coelebs
L
.
),
without
dependence
from
to
what
subspecies
it(he)
concerns
.
•
It
is
possible,
that
more
frequently
evolution
occurs
in
time
,
instea
d
of
in
space
-
and
the
species
specific
song
of
chaffinch
here
can
be
an
example
.
During
time
,
variants
or
types
of
song
at
chaffinch
can
form
the
certain
variation
numbers
(lines
),
philetical
(evolutionary
),
frequently
the
crossed
,
vocal
lines
,
representing
more
and
more
new
distinguished
(different
)
,
but
also
similar
vocal
forms
,
also
including
and
former,
initial
types
of
songs
(fig
.
*)
.
•
At
the
quantitative
analysis
have
been
revealed
less
stable
features
of
species
specific
song
of
chaffinch
(
Fringilla
coelebs
L
.
)
which,
probably,
and
in
the
greater
degree
will
change
at
formation
of
new
vocal
norms
(basically
it
concerns
fine
or
small
details
of
song
-
length
and
intervals
of
elements
songs,
song
parts
and
phrases,
probably,
their
quantity),
rather
than
those
song
parameters
and
their
correlations
,
which
appeared
more
stable
(a
frequency
range,
character
of
singing
of
song
pattern
and
intervals
between
them,
some
form
of
a
pronunciation
of
elements),
that
also
is
capable
to
change
.
Fig. **. A degree of similarity of different chaffinch song types (
Fringilla coelebs
L.) under the basic forms of elements (syllables) in populations of European Russia (Astakhova, Byome, 2007)
Lines of vocal development (vocal lines
) -
a number (line) of song types, similar on the structure, "related" vocal forms by origin, probably, generated (formed) at change and transition of one song type in other song type, can represent result of differentiation (modification) of one initial ("
ancestor
") song
type (on fig
. **
are vocal lines of chaffinch -
types of songs are close on similarity (more similar) -
U I A
, V M F D
, T S E B
, W N H G R
, J O C
).
At results (at support) of dendrogram of similarities of song
types
of
chaffinch
e
uropean (
Fringilla coelebs coelebs
L.) (fig. **), it is possible to make conclusion,
what types of songs
more
differ from the others
, and what song
types
of chaffinch
"
are more related
" (
are close)
with each other
in the origin
,
and
which of song types
create (f
orm
)
separate evolutionary trunks
(
fig
. *) of the
vocal lines
,
crossed and differentiated again and again. Also it is possible to make attempt to reveal the most ancient vocal forms of song
at
chaffinch
e
uropean (
Fringilla coelebs coelebs
L.), i
t is possible to tell, song
types -
"ancestors
" of separate evolutionary vocal trunks (fig. *) of species specific song of chaffinch
, existing now
. Probably, direct
(similar) descendants
of ancient forms of chaffinch song are types of songs
J, O, C, V, A, R (
fig
. ***). Probably, these
song types represent the most ancient structure
of
chaffinch song
, kept (preserve
) or poorly changed till now
, existing equally
with other song
types
, probably, already with their descendants
, which are
changed enough
and representing already other,
younger
, vocal lines
or a separate large evolutionary trunk
of chaffinch song, developing in parallel
with ancient vocal forms
, their vocal lines
. 1.
2.
3.
4
.
5.
6.
Fig. ***. Ancient forms of song at
chaffinch e
uropean (
Fringilla coelebs
L.),
being founders of the separate evolutionary vocal lines
,
existing equally
and in parallel
in relation to evolutionary
more
young song
types (it is possible, their close "descendants") and
in relation to their vocal lines (Astakhov
a
, Byome
, 2007): 1 -
song type J, 2 -
song type O, 3 -
song type C, 4 -
song type V, 5 -
song type A, 6 -
song type R (different stages of evolution of vocal lines
are represent -
earlier vocal forms of chaffinch song are the first
(1
-
3)
, and further
(4
-
6)
-
later vocal forms and their evolutionary lines).
Fig. 20
. Representatives of one group or genus of Finches (Fringilla)
:
A –
chaffinch
(
Fringilla coelebs
L.), B
–
Brambling (
Fringilla montifringilla
); 1 –
female
, 2 –
male
.
Fig. 22
. Blue chaffinch
(
Fringilla teydea
) on Canary islands (Tenerife, Grand Canary) (Bergmann, 1993).
Fig.
19
. 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. 21
. Subspecies of chaffinch
(
Fringilla coelebs
L.) on Canary islands (northwest of Africa) –
chaffinch
(
Fringilla coelebs tintillon
) (Bergmann, 1993).
Assumptions of evolution of chaffinch song (
Fringilla coelebs
L.) a
t comparison with a song of brambling
(
Fringilla montifringilla
)
Fig. 23. Sonograms of
songs and calls (voices)
of different species of
chaffinch
(Fringilla): a, b -
Fringilla coelebs tintillon
(island of Tenerife
, Canary islands), c
, е -
alarm (danger) calls (voices), d -
Fringilla teydea
(
blue chaffinch
), Canary islands (
Bergmann, 1993
).
Discussion of laws of evolution of chaffinch song (Fringilla coelebs
L.)
(Astakhova, Byome, 2007)
At comparison of sonograms of
songs
of different species
of birds in genus of
Finches
(Fringilla) and some of subspecies of chaffinch
e
uropean
(
Fringilla coelebs
L.) there is a question
,
how
singing (song) of birds can to develop, to change
(modification)
, proceeding
at beginning
(differentiation) from the general
(common) ancestor
song
forms to
songs
, which we can observe now at different philetical
(evolutionary) lines
. Whether
it is
influence
of ecological conditions
on evolution of specific songs
of birds
or it is only result of their genetic and "social" (
song learning
) variability
? To answer this question, precise
and, at the same time
the thin analysis of songs
(their types) is necessary at close (closely related) species
and at subspecies of birds
(in this case, at chaffinch
)
.
It is interesting to compare two close species
of the birds
, belong to genus of Finches (Fringilla) -
chaffinch
(
Fringilla coelebs
L.) and brambling
(
Fringilla montifringilla
). Brambling
(fig. 20.2) has not sonorous
,
but
has "buzzing"
species specific song
(fig. 19
E
, 23*.1), as
against
chaffinch
. Probably, such song of brambling
was generated
(
formed)
initially
,
there where brambling
arose (origin) as a
species
-
in habitats with low and rather thin (sparse) vegetation (forest
-
tundra
), therefore a species specific song of brambling
is narrower in frequency range (К
Hz
) (that is its(her) minimal and maximal frequencies have smaller limits
(difference) in the values), than a species specific song of chaffinch
, frequently having wide frequency range (К
Hz
) (fig. 23*.2). Therefore the song of brambling on hearing is perceived
by
"
creaking
"
sound
or as "
gnash
", almost as "
ryumin
" (
ryu
-
ryu
) of chaffinch
-
his(its) call (voice) in cloudy or cool weather.
Probably, such narrow frequency range of song at brambling
has appeared
,
because of the greater ease of passage of a sound among poor vegetation of forest
-
tundra
–
“
it is not necessary to try
”
to sing sonorous
and more complex
(difficult). And, probably, the "modest" song of brambling
also was f
ormed
at economy of expenses of power
(energy
),
because
the forest
-
tundra as northern zone of the Earth
is
colder on mid
-
annual temperature
and is poorer in food resources
-
probably, therefore song realization of brambling
appeared not such bright
, as at chaffinch
. In southern forest
-
tundra
prevail bushes
and low trees
, which far remote
from each other. After occurrence of precisely established
species specific song
of
brambling
on area
, which, probably, he(it) inhabited long time
in the north, at his(its) following distribution to other areas
of continent, the song has remained
"buzzing
"
,
though also has variants or types.
Fig. 23*. Species specific songs
of brambling
(1) (
Fringilla montifringilla
) and chaffinch
(2) (
Fringilla coelebs
L.) in England
(
Thorpe, 1958b
). Both species
of birds belong
to genus of Finches
(Fringilla).
At chaffinch
in process of “
cultural (
song
) evolution
”
the wide frequency range
of species specific song
has developed
, it is probable, because it(he) widely occupied a wood (forest) zone
, forest
-
steppe zone
, where among dense vegetation
of leaf
-
bearing forest
and coniferous woods
(forest)
passage of a sound is more complicated
. Therefore, probably, chaffinch was capable to form loud (sonorous) and complex
(difficult) songs
-
for their best passableness
through vegetation, moreover -
the reserve of
forage (food
) and good temperature conditions
of an occupied natural zone allowed
to chaffinch
to do
(make) it
and promoted
it(him) song realization
. On the physical data
,
in air space
the most good
(optimum) passes
of a sound
by
frequency 3,5 К
Hz
. A
verage frequency of species specific
song a
t chaffinch
is represent
about 4 К
Hz
(though at many other species
of sparrow birds the song has similar frequency). Probably, such adaptation of song
of birds (
chaffinch
, in particular) in a frequency
range
is intended for the best passage
of a sound in
a wood
(forest)
zone
,
where chaffinch
also
inhabit
. Thus chaffinch
is a "social" species
of birds, and chaffinch
males create (
form
) microgroups
, settlements in local populations
,
where antiphonal sing with
each other
("
roll call
" by songs), thus training (learning) of the following (young) generation
by species specific song
, by "
local culture" of song
(by ways of singing, variants or types of species specific song). Probably, therefore, for original (special) "dialogue" or "communication
" for song of chaffinch
it is necessary to be well passable (loudness) in sound energy (
Db
) among wood and bushes
vegetation
. In connection with good audibility
of sonorou
s chaffinch song
, his(its) individuals cooperate
and are interconnected
by a uniform species specific song
(different types, variants, but similar in the structure) in different local populations
during all area of distribution of a species
(continent of Eurasia, England, northern Africa, on Canary islands), that, thus, support
(keeping) a species of chaffinch
in integrity (uniform) at formatio
n
of some his(its) subspecies or geographical races
(the geographical populations
,
located from each other at a great distance) (fig. 19). Individuals of chaffinch of different subspecies
(geographical populations) can differ
by
the intensity of feather color
(according to atlas of determi
nation
of birds) (Dementyev, 1954) and differ i
n typological structure
of species
specific song
(
in “song culture
”
,
in ways of singing of species specific song), in its(her) character (features) of structure, in a frequency range
(Marler, 1952; Thielcke, 1969; Slater et al., 1979; Ince et al., 1980; Catchpole, Slater, 1995; Simkin, 1983; Bergmann, 1993).
Here also it is possible to note ideas of G.N.Simkin
(Simkin, 1983) about "adaptation" of chaffinch song to geobotanical conditions of a habitat
of a population of birds and to change
s
of their song in structure
(frequency, length, number of syllables or elements, number of phrases or parts of songs) according to
(at influence)
the certain ecological factors
. Variants of song
types
of chaffinch
(
Fringilla coelebs
L.) in different "biotops"
(
in territories with different ecological conditions
)
or song versions
are
named
by G.N.Simkin
as "biomorphs
"
(as biotypes)
, so
-
called results of "ecological adaptation
“ (fig. 13)
.
But nevertheless the idea of "
biomorph
i
s
m
"
(as
ecological
vocal polymorphism
) in a song of chaffinch
can have
and precise frameworks (limits
), because
it
is feebly
(
weak
) marked
(expressed)
in song
types at dwelling
(inhabit)
of chaffinch
in similar biotops
(different ecological places)
(
leaf
-
bearing and coniferous
vegetation -
wood and bushes
), despite of the big variety and variability of
song types
of chaffinch
in these territories
of a wood
(forest)
and forest
-
steppe. A species specific area
of chaffinch
(
Fringilla coelebs
L.) b
asically it is located in zones
of wood
(forest)
and forest
-
steppe
,
where biotops
(different ecological places)
in many factors are ecologically similar
, but a variety of song
types thus remains
. Therefore to allocate
(distinguish)
here "ecological phenotypes
" of song it would be not absolutely correct
. Here there can be simply a variety of song types
of chaffinch
, formed in result of "mistakes" of copying at song
training
(learning
)
by young birds and at improvisation of songs
in their structure at singing. But it is impossible to assert
(approve) precisely
,
wh
ich
of
song
types of chaffinch
"is adapted" in the structure to birch or oak woods
(forest
)
and w
hich
of
these song
types
were generated
(created
)
in fur
-
tree or pine vegetation
. Though some
time was proved, that the structure of song
depends
on a degree of passage
of a sound
through an environment
-
certainly, with increase
of density
of vegetation (environment) can increase
are
force of a sound
(loudness of singing (
Db
)), complexity of song
(longer and various on structure)
, can
widen (broaden) the
frequency range of song (К
Hz
) (Handford, 1988; Lougheed et al. 1989; Handford and Lougheed, 1991, Wiley, 1991). Probably, also species specific song
of chaffinch (its(her) variants, types) is developed
in this direction
(scheme)
in
process of song (culture) evolution
during
time
-
in more dense woods (forest
) (taiga, for example) a song
could have the variants, which are similar
in the greatest sound capacity
(loudness) (
Db
), in the greatest complexity
(the greater number of syllables or elements, number of phrases or parts of songs), i
n wider frequency range (К
Hz
), than types of species specific song
,
sung
(performance)
in woods (forest
) (
their biotops or different ecological places
) with smaller density of vegetation
,
where
these song types
can have
smaller (weak) loudness (
Db
), thin and frequently short (
sec
), simple structure
, narrower frequency range (К
Hz
). Thus the form (figure) of elements
(syllables, phrases or parts) of songs
can not to change
under action (
at influence) of ecological factors
, because in the greater degree
qualitative aspect of songs
(a variety of forms of elements
in the songs, observable on sonograms
) are result of "social" interaction
of birds
, their copying
from each other song types
(their elements),
result of song
improvisations
or also
it displays
(expresses)
genetic variability
(at species
of birds without social traditions
of song training
or learning
). And, mental
essence
(psychical sign)
of birds
(though and in a rudiment) is so plastic
, that its(her) genetic inclinations
can to change
already
in the first generation
, a
t adapt
ion
to an environment
. In our researches
(2005
-
2007
) the same
(one)
song
type
of chaffinch
(
Fringilla coelebs
L.) has been found
in the center of the European Russia
, but
in different ecological conditions
it(he) was performance at singing
with different intensity
(sound force (
Db
)), with
different complexity
: for example, in an old dense wood
(forest)
chaffinch
male sang (performance) song
type
C
is loudly enough (
Db
), the song included two trill
phrases (part
s
), she(it) was wide in frequency range (К
Hz
), but in seldom growing birches
(about 5 km from the first territory) the chaffinch male
sang
(performance) song type C
is more "simplified
": the second phrase (part) of a trill
was absent
, the song had smaller sound capacity (
Db
), had thin structure on sonogram
, though the frequency range (К
Hz
) could remain former
or hardly smaller
(figures in following page 34
-
35)
.
Also, for example
, on Curnian spit of Baltic sea
(the Kaliningrad region) the chaffinch song
appeared more sonorous
(more loudly or more powerfully in sound energy (
Db
)), than
frequently in the center of the European part of Russia
(
Zvenigorod
, Moscow, Michurinsk). Probably, such distinction of loudness (
Db
) and clearness at
singing
of chaffinch,
because
that on
Curonian spit
the noise background
was higher or as a result of affinity of Baltic sea
(his(its) noisy waves
at coast), or in result of higher density
of a
population
of chaffinch on Curonian spit
. Probably, such phenomenon
also is possible to count
(
consider)
as adaptation of singing (songs) of chaffinch
to an environment
. As the fact
, that in a city zone
(settlement)
,
where noise background is higher
,
because of an abundance of the vehicles
, of cooperating social groups of people
, different species
of sparrow birds sang
(performance)
much more loudly (
more sonorous
) (
Db
), than in quieter and not noisy zone of woods
(forest
) (Rheindt, 2003).
Certainly, all this proves immediate reaction of birds to an environment
and the adaptation to its(her) qualities
, but only plasticity of mental essence(psychical sign) of birds
(its(her) rudiments
) allows
these "adaptations" to change quickly and sharply so
, that they (
psychical adaptations
) can be imperceptible, insignificant, astable, short in the existence
. Other talk, if the question is about long (during many years) adaptations of any population of birds
in the certain
(
definite
)
habitat
with the certain
(
definite
)
ecological conditions
and formation
thus more or less stable "
song
cultures
"
(the certain set or complex of variants or types of species specific song
), but if also this population
could have small (weak) communications (connections
) with other populations
of a species
-
here is
already possible to speak about occurrence of "adaptations to enviroment
", including song aspect at birds
. Though, certainly, the majority of populations
of birds represent "open system
", interconnected and constantly mixed with each other
, especially during migrations
, therefore "song
culture
"
(as well as many others populational
attributes
of birds) will be constantly updated, change
d
, but, nevertheless, there is populational
variability as a whole
, its(her) fitness
(
adaption
)
and formation
of
separate populational
(and then, may
be, and
species
specific
) attributes
(signs
)
in the certain
(
definite
)
territory
during certain
(
definite
)
time
. There is an interesting fact
,
which shows, that songs of nesting birds
(
residents
-
come back from places of winterings on the certain or definite place of dwelling (inhabit) from year to year enough long time
) differ
by the greater clearness (accuracy) in structure of song
elements
(syllables, phrases), than it happens in songs of birds
-
migrants
of the same (one) species
,
which can be characterized by the greater uncertainty
(unclear
)
in song structure
, in the form of elements
, phrases (parts) of song
types. Probably, the reason
(cause
)
of it is
,
that migrants still
should occupy any territory of a species specific area
,
where they will be compelled to ad
a
pt
to local "
song cultur
e"
of birds
(of r
esidents
), nesting there
(or are trained or learned
by given "
song culture
"
-
copying of local ways of singing of species specific song
, its(her) types
, variants). Probably, therefore less precise songs of birds
-
migrants
thus are adapted more and faster to change in the structure
(to be the greatest plastic
at training or copying of songs) and as a whole to be better adapted to local "
song culture
"
(Nelson et al., 1995). The figures in following page 34
-
35
Some conclusions from
researches
(Astakhova, Byome
, 2007)
The song type C on Curonian spit of Baltic sea
(where
chaffinch
(
Fringilla coelebs
L.) is a usual nesting
species
) -
elements of songs
are more precise
, well differentiated and definite (determined
).
Song type C in the Center of the European Russia (Moscow
) (where flying or migratory populations
of chaffinch
(
Fringilla coelebs
L.) are strongly mixed with his(its) nesting populations
) -
elements of songs
are not precise, poorly (weak) differentiated, uncertain (undefinite, unclear
).
Song type С* (the reduced form) of chaffinch in the Center of the European Russia
(
Zvenigogod
,
the Moscow region) -
record of song in an old fur
-
tree wood (forest
), his(its) density is significant
(
the song has more powerful sound energy
).
Song type С* (the reduced form) of chaffinch
in the Center of the European Russia
(
Michurinsk
, the Tambov region) -
record of song in young birchs on edge of road
, trees are rare
, are planted (put) in one row (line
), density
of planting (spreading) is low
, the trees border on a field
(
the song has smaller
or weak
sound energy, thin structure
).
The song type I
on Curonian spit of Baltic sea
(where
chaffinch
(
Fringilla coelebs
L.) is a usual nesting
species
) -
elements of songs
are more precise
, well differentiated and definite (determined
).
Song type I
in the Center of the European Russia (
Zvenigogod
,
the Moscow region) (where flying or migratory populations
of chaffinch
(
Fringilla coelebs
L.) are strongly mixed with his(its) nesting populations
) -
elements of songs
are not precise, poorly (weak) differentiated, uncertain (undefinite, unclear
).
Geographical populations of chaffinch
(
Fringilla coelebs
L.) in northwest (
Curonian spit
, the Kaliningrad region) and in the center of the European Russia (Moscow, Zvenigorod
, Michurinsk) are on distance of 1000 km
. Between populations of Moscow, Zvenigorod
(the Moscow region) and Michurinsk (the Tambov region) distance of 850 km
.
In a case of chaffinch song it also was well appreciable. On Curonian spit of Baltic sea
(the Kaliningrad region), w
here chaffinch
is a nesting species
, we have noted the big clearness, a subtlety, definiteness of song structures at
singing
and of separate song elements
, than at chaffinch
in the central part of the European Russia
,
where it(he), apparently, in many populations is "flying"
(passed)
or the migrant
. In these "passed"
local populations of chaffinch
(migrants
) many
song types were indistinctly
(
not thin, unclearly, not definite
) are determined in forms of the elements
(syllables, phrases) (as it was seen
on sonograms
), and also in populations was revealed (
observed
) the greater number of the "combined" (mixed) song
types
(consisting of phrases or parts of other types of songs)
.
It can speak about formation
of cumulative (complex) "
song
cultures
"
of chaffinch
in
this territory
, probably, sets of song
types
of several local populations of chaffinch
, capable to adapt
to a
row (
line) of local "
song
cultures
" of
chaffinch
on spreading
on the big territory of a species specific area
, to coexist with a wide spectrum of local dialect forms of species specific song
(
local ways of singing
song
types and their variants
) (
figures on previous page)
.
If to speak about habitats of chaffinch
in other natural zones of the Earth
(rather than only in wood
(fores
)
and forest
-
steppe zones
)
,
where chaffinch is common
, we admit
(allow),
that at autumn migration
,
chaffinch can be on the south
(a place of winterings
) -
area of the Mediterranean sea
, the north of Africa
, Caucasus (Asia
), where there can be a zone of subtropics
, semideserts
, mountains
, and
, certainly, in this case is possible to pay attention
to the adaptation of chaffinch song
to surrounding conditions of the nature
(more likely, to character of vegetation
, a geology
, a climate of a place
of chaffinch dwelling
, where inhabit
). If the population of chaffinch inhabit
in these natural zones
enough
long time
, despite of constant mixing of residents (nesting) and migrants
of chaffinch
, independent of mixing of individuals
there is a gradual adaptation of the given population of birds to local ecological conditions
, and in song
aspect
-
also
(besides, in quantitative parameters of song
-
length, a frequency range, a degree of differentiation of
song structures or its(her) complexity), but thus at song
training (learning
) occurs also qualitative transformation
of species specific song
(the form of its(her) elements
, their variety
) in the given population
of birds, by creation
certain
(determined) "
song
cultures
"
(ways of singing species specific song, the certain set (complex) of its(her) types
(variants), distinguished or different
(but, may be, in something and similar
) from "
song
cultures
"
of other local or geographical populations of this species
(in this case, of chaffinch
). Therefore
,
now
we
can observe
(reveal)
so wide variety
of species specific song
s
of sparrow birds
, and as the private
(individual) phenomenon
-
so mysterious differentiation (division) of initial (
ancestor
) form
s of
species specific song
of
chaffinch
into
different song variants
(
song
forms
) at different subspecies
(geographical populations) of chaffinch
, their establishment
(at forming)
in separate song norms
,
including the certain (definite) rows
or lines
of development, modification
, characteristic for the certain
(definite)
habitat
and local ecological conditions
. Fig. 24
. Song
types of chaffinch
(
Fringilla coelebs
L.),
making (
forming) different vocal lines, but passing
(transition)
each other:
1 –
song type С*11 (record on Curonian spit
, the Kaliningrad region), 2 –
song type CM 5 (Simferopol, the Crimean region), 3 –
song type CM 3 (record in Melitopol, the Zaporozhye region), 4 –
song type CM 4 (Jankoy, the north of Crimea), 5 –
song type OI*3 (Simferopol, the south of Crimea), 6 –
song type OI 2 (record in Simferopol, the Crimean region), 7 –
song type M(O)I (Simferopol, the south of Ukraine), 8 –
song type O (
Curonian spit
, northwest of the European Russia). Comparison of song types of chaffinch (
Fringilla coelebs solomkoi
Menz.) in the south of Ukraine (Crimea) and chaffinch (
Fringilla coelebs coelebs
L.) in northwest of the European Russia (
Curonian spit of Baltic sea
) (
Astakhov
a
, Byome
, 2007
)
Simferopol, Jankoy (Crimea) and Melitopol (the south of Ukraine, the Zaporozhye region) are remote from each other on the average on 100 km.
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. Comparison of basic parameters of song types of chaffinch (
Fringilla coelebs
L.)
in different geographical populations of the East Europe
(
Astakhova, Byome, 2007
)
.
At results
of quantity analysis
, i
n 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
(the table is lower) (Astakhova, Byome, 2007)
.
Probably, "displacement upwards" of frequency range (К
Hz
) of species specific song
of
chaffinch
(
Fringilla coelebs
L.) in the south
also is feature of southern song
cultures
(singing of species specific song) of chaffinch
and can give rise
to (beginning)
of
the new vocal form
,
already adapted to local ecological conditions in the south
, probably, initial for species specific songs
of many
future new species
and wider groups of birds
. Comparison of some song types of chaffinch (
Fringilla coelebs
L.) in a population of the south of Ukraine with corresponding dialect (modified) song
types in a population of northwest of Caucasus.
While in the south (Caucasus, Crimea
) could meet those general (common) (similar) song types
of chaffinch
(
Fringilla coelebs
L.)
,
which were absent in the north or in the center of the European Russia
.
Song type ME (I) (
Teberda
, Karachaevo
-
C
he
rcassia, 1985) Song type ME (I) 9 (Simferopol, the Crimean region, 2007)
Song type GC
(
Teberda
, Karachaevo
-
C
he
rcassia, 1985) Song type GC3
(Simferopol, the Crimean region, 2007)
Simferopol –
Pitsunda
(560 km)
Simferopol –
Teberda
(670 km)
Song type MI (B) (
Pitsunda
, Abkhazia, 1985)
Song type В* (BF) (
Bordgomi
, Georgia, 1985)
Song type CM
(
Teberda
, Karachaevo
-
C
he
rcassia, 1985) Type of song
MI (
B
) 8 (
Simferopol
, Crimean
region
, 2007
)
Type of song B
* (
BF
) (
Melitopol
, Zaporozhye region
, 2007
)
Type of song
СМ 2 (
Melitopol
, Zaporozhye region
, 2007
)
Simferopol –
Pitsunda
(560 km)
Simferopol –
Teberda
(670 km)
Melitopol -
Teberda
(630 km)
Melitopol -
Bordgomi
(830 km)
Apparently from represented
sonograms
, songs of chaffinch (
Fringilla coelebs
L.) in different geographical populations
of the south of Ukraine (Crimea) and of northwest Caucasus can represent dialect forms of one song type. Though the songs of chaffinch
on Caucasus
have been recorded a
long time
ago
-
in 1980th years
(Sultanov, 1985). Nevertheless, it is possible to distinguish and now
similarity of some song types of chaffinch
, that speaks about stability of song
cultures
of chaffinch
in u
n
it
ed
or connected (cooperated
)
local populations
during
time and in space
, despite of individual and geographical variability of their
song
types
. At
similarity
of structure of some dialect song
types
(see sonograms
), geographical populations
of the south of Ukraine and the western Caucasus can be connected as a result of constant migrations
of individuals
,
that leads to mixing
and, somewhat, to joint development of song cultures of chaffinch
in local populations
. Probably, the distance
(on the average 600 km) allows to support
these populational
communications (connections
) between regions of the south of Eurasia
.
Thus, the song (singing) of birds
develop (evolution
) on the same, similar
biological laws
, as also many other displays (expressions) of life of an organism
. But it is necessary to emphasize
special (particular) plasticity
in functioning and the adaptation
of mental essence
(psychical sign) (though only and its rudiments at birds
) -
it is capable plural (
changeable
) and quickly to react
(by perception)
on irritation
(on surrounding)
,
also
can quickly
turn psychical action
(by transition)
to
new way
s
(special manners or patterns of acts)
and can quickly "
forget"
, change former
(
previous) course
. But in larger scales of evolution
(
populations
, species
) in displays
(expressions)
of psychical essence
(songs of birds)
,
nevertheless
,
can be
revealed
(
observed
) also ecological differentiation
and an establishment
(at becoming, forming) of "adapted" more stable forms (norms) of song
, that are similar
, for example, as development of
morphological attributes
(signs)
of birds
. Thus, song (singing) of birds
includes
"
social" character of evolution (
development
)
, and that
is unique
and it seems "parallel
" in the existence to other morphological attributes
(signs)
of an organism
, -
as if song (singing) of birds
represents
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-
such, as display
(
expressing) of mental (psychical) sign
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(energy)
of an organism
, as essence of higher level of the organization
, but existing on the same laws
, as also others properties
, more material for the person (human
), "
more biological
" and appreciable attributes
or signs for an organism
(color of covers, the form of a body and his(its) parts, character of a skeleton at animals, structure and features of internal bodies of an organism, differentiation of his(its) cells and tissues, features of a nourishment (food), reproduction of descendant individuals, a way or pattern of life as a whole). L
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chaffinch
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in
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Автор
Olesya Astakhova
Документ
Категория
Зоология
Просмотров
138
Размер файла
148 860 Кб
Теги
fringilla, coelebs, song, evolution, chaffinch, assumptions
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