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Astakhova O. A., Byome I. R. Song of chaffinch (book)

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How the evolution of chaffinch song happens? Males of chaffinch have 1-6 song type (everyone) and contain more 20 song types in population.

Astakhova O. A., Byome I. R.
Song of chaffinch (Fringilla coelebs l.):
the structure and evolution
(research works)
Content
Principles of the analysis of structure of chaffinch song (Fringilla coelebs L.). Individual variability of song3
The typological organization and microgeographical variability of chaffinch song (Fringilla coelebs L.) in the population of the Curonian spit10
Vocal variability and dialect forms of chaffinch song (Fringilla coelebs L.) in populations of the central part of the European Russia25
The typological organization of chaffinch song (Fringilla coelebs L.)
in population of Curonion spit40
Macrogeographical variability of chaffinch song (Fringilla coelebs L.)
in populations of the European part of Russia42
The analysis of phrase combinations and various ways of syllabic singing at vocal variability of song types of chaffinch (Fringilla coelebs L.) ++44
The quantitative analysis of chaffinch song types (Fringilla coelebs L.) in populations of the European part of Russia57
Vocal variability of chaffinch song (Fringilla coelebs L.) as a condition of cultural evolution in local populations72
Degree of prevalence of different song types of chaffinch (Fringilla coelebs L.)
in populations of the European Russia ++84
Results of the comparative analysis of chaffinch song (Fringilla coelebs L.) in geographical populations of the south of Ukraine (Crimea) and the European Russia93
Chaffinch (Fringilla coelebs L.) song in populations of the East Europe95
Combinative variability at different levels of the organization of a matter of the Nature: chromosomes of cells of an organism and vocal realization of birds (Aves)103
Assumptions of evolution of chaffinch song (Fringilla coelebs L.)106
Principles of the analysis of structure of chaffinch song (Fringilla coelebs L.). Individual variability of song
O. A. Astakhova, I. R. Byome
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Chaffinch (Fringilla coelebs L.) is one of the first and most widespread objects of researchers of song organizations of birds (Sick, 1939; Marler, 1956а; Thorpe, 1958).
The resounding and precise song of chaffinch frequently drew attention of bioacoustics as an accessible subject of studying of vocal laws - development of songs in ontogenesis (during life of birds), becoming of vocal repertoire, geographical variability of songs and calls (of voice) of birds (Promptov, 1930; Thielcke, 1961; Nottebohm, 1969а, 1967; Simkin, 1983; Slater et al., 1984).
In the given work, generalizing of principles of song division into structural parts are submitted, the specific features of song types at different chaffinch males is discovered.
Material and methods
During the spring period (April - May, 2005) in different points of territory Curonian spit of Baltic sea (the Kaliningrad region), where the density of a population of a chaffinch (Fringilla coelebs L.) was the high, tape recorder records of singing males (on the average n=20 songs from everyone) are made.
Record has been executed with the help of tape recorder Panasonic RQ-SX95F, of condenser microphone Philips SBC ME570. Further, sonograms of songs were analyzed with the help of computer program Avisoft-SaSLab Light. In total it has been analyzed about three thousand songs. Types of songs are marked by Latin letters.
At the quantitative analysis measured both structure of song type as a whole, and its separate phrases (in three parts of songs - started singing, a trill, a final stroke) on different frequency-and-time parameters: length (duration) of song (sec), maximal, minimal and average (median) frequency (KHz), quantity of elements (syllables), length of syllables (sec), intervals between songs (sec).
Results and discussion
Long time many fans (amateurs) of songs of birds at singing of chaffinch (Fringilla coelebs L.) defined (determined) only a long trill, frequently with the sharp termination (ending) in the end. In the middle of XX century, bioacoustics have confirmed, that the trill of song has a descending tonality (that is begins higher frequencies and passes in lower) (Witherby, 1944). In result, in a song of chaffinch (Fringilla coelebs L.) three phrases (knee) are allocated (distinguished): the first - usually increasing in a sound, the second - shorter and the constant frequency (KHz), doing (making) a song with clear step-by-step downturn of a sound, and the third - a final stroke (a sharp sound), which can be complex (Marler, 1956а; Thorpe, 1958). Also it has been marked, that the given three phrases (knee) of song can be subdivided into the parts, containing different notes (for example, the first phrase was subdivided on 1а and 1б).
For the analysis of song structure of birds exist terms of different character: musical (the note, motif) and linguistic (a syllable, a phrase, a strophe or line). Usually both those, and others have mixing use - that it is possible to name a phrase, it name also motif or a tune (the certain part of song, that consist from similar sounds - notes or syllables, elements). The full finished song of a bird can be named a strophe (stanza) (song pattern), which is subdivided into phrases or a knee (oscillation) (Jellis, 1977).
Also there are songs-variations - different forms of one song pattern, which can be classified in variational lines (rows) (Thompson, 1970; Slater et al., 1980). On the basis of these data we have tried to analyze songs of chaffinch, which have been recorded in population of Curonian spit (N=153 males).
Fig. 1. The analysis of structure of song type С of chaffinch (Fringilla coelebs L.)Figure 1
Principles of the analysis of song structure of chaffinch (Fringilla coelebs L.) (our view with a support on references):
1) On perception (recognition) on hearing, the song was subdivided into three parts: the row (number) of whistle sounds (as if started singing), trill sounds (are as though poured each other) and a final stroke (the example of the analysis is shown on more widespread song type С) (fig. 1).
2) Within the limits of these parts of songs on sonograms it is possible to allocate (design) the phrases (the elements similar in the form), in this case a trill includes two phrases in song type C.
3) Elements (syllables) can be simple (started singing and 1-st phrase of trill in song type C) and complex (compound), consisting of two and more subelements (2-nd phrase of trill in song type C).
4) Syllables (elements) are divided (shared) by intervals, but frequently shorter, than at phrases.
5) A final stroke in many types of songs consist of elements (syllables), different in the form (it is possible, therefore it such sharp, remarkable, "bright" on hearing).
At record, songs of one type met in different points of territory (was considered, that belong to repertoires of different males), therefore alongside with the letter were marked by numbers in ascending order (for example, А1, А2, А3, etc.). At the analysis of sonograms of chaffinch songs come to light (reveal) distinctions in their time-and-frequency parameters (at songs of one type) - so-called individual variability. It is possible to show on example of song type C (fig. 2).
Individual variability - we named as variants (versions) of songs of one type, fixed (recorded) in different points of certain territory or in repertoire of one male. Songs were considered as songs of one type, that had two similar or completely three parts (started singing, a trill, a final stroke), which of them (parts) can be subdivided into phrases (the second order) (fig. 1).
In figure 2 it is necessary to note, that besides small distinction of forms of elements of different parts of variants of song type C (of started singing, trill, a final stroke), can meet and in many respects the modified variations of song - dialect forms, for example, song type С*11 (fig. 2б). This phenomenon is interesting, that dialects of songs can exist together in one local population, probably, as a result of mixture of traditions of different song cultures during migrations (Slater, Ince, 1979, 1980; Espmark et al., 1989).
At the quantitative analysis of variants of song type С (tab. 1) were found out, that the greatest distinction in their length, quantity of syllables and intervals between songs at singing. In other parameters is relative uniformity of the data is observed, but there are the values most distinguished from others.
Fig. 2. Variants of song type C of chaffinch (Fringilla coelebs L.), recorded on territories of Curonian spit in different points: a - song type C, b - song type С*11, c - song type С3, d - song type С4, e - song type С5, f - song type С6.
Variational rows (lines) have been made for the majority of song types of chaffinch, which allocated (designed) on Curonian spit of Baltic sea (in sample 22 song types), are especial for those types of songs, which had strong prevalence in a population (n>15). It is possible to present variants of song type O (fig. 3).
Despite of similarity song variants, exists improvisation at singing of different males (especially in a final stroke), forming individual variability of song type O. It is possible any of these similar song forms in further will give the new song type at the development.
Conclusion
Thus, song of chaffinch (Fringilla coelebs L.) can consist of three parts: started singing (the row of whistle sounds), trill sounds (an average part) and a final stroke. On sonograms within the limits of each part it is possible to allocate the separate phrases that consist of elements similar in the form.
Fig. 3. Variants of song type O of chaffinch (Fringilla coelebs L.), recorded in one local population: a - song type O, b - song type О1, c - song type О2, d - song type О3, e - song type О4, f - song type О5.
Individual variability of songs of one type exists, but is frequently insignificant. Though in one territory it is possible to meet dialect forms of one song type (enough modified in the form of elements, but having the general (common) similarity in structure).
The table. The basic time-and-frequency characteristics of song type C of chaffinch (Fringilla coelebs L.)
Type of song (token)Sample size (songs)The name of record placeThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables
in song typeLength of syllables of start singing, secLength of syllables trill, sec*Length of syllable of final stroke, sec*Intervals
between songs,
sec1
phrase2
phraseС15Trial
point2,025
±0,111,447
±0,1668,01
±0,374,134
±0,21516,7
±1,160,13
±0,010,084
±0,0280,13
±0,0070,085
±0,00511,61
±5,06C*1122
point2,55
±0,0821,3
±0,377,751
±04,478
±019
±1,40,175
±0,0070,034
±0,010,084
±0,0010,915
±0,00222,9С3208
point2,074
±0,1181,627
±0,1319,698
±0,414,0996
±0,17317,25
±0,850,1203
±0,0050,068
±0,0030,1155
±0,0080,071
±0,0056,423
±1,9С4219
point2,01
±0,211,75
±0,159,25
±0,64,056
±0,2216,45
±1,370,122
±0,010,066
±0,0050,128
±0,010,074
±0,0087,06
±0,01С5611
point1,75
±0,151,7
±0,137,8
±0,54,3
±0,0713,7
±1,030,133
±0,0050,067
±0,0040,103
±0,0030,068
±0,0055,4
±0,85С6313
point1,96
±0,121,55
±07,522
±0,364,36
±0,215
±10,135
±0,010,0632
±0,0050,0913
±0,050,07
±0,0064,44
±0,91
The note: specified are average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type, which were reproduced by chaffinch males in the given points of record; the strongest differences of song types counted a difference of parameters > 0,5 КHz in frequency и > 0,02 sec in length (are allocated by a font); * - the trill of song type C consist of two phrases; the values of parameters, most distinguished from others, are separately allocated (designed).
Gratitude
Authors express deep gratitude to employees of a biological research station "Fishing" of the Zoology Institute of the Russian Academy of Science on Curonian spit for the help and support in work, gratitude to prof. G. N. Simkin for valuable advice at the analysis of a material. Work is executed at financial support of the Russian Federal Property Fund, the grant 04-04-49276.
The literature
Promptov A.N. Geographical variability of chaffinch song in connection with the general questions of seasonal flights of birds // Zoology Journal. 1930. V. 10, № 3. P. 17-40.
Simkin G.N. The typological organization and populational phylogeny of birds song // The bulletin of Moscow society of nature researchers. Section of Biology. V .88, №1. 1983. P. 15-27.
Espmark Y.O., Lampe H.M., Bjerke T.K. Song conformity and continuity in song dialects of redwings Turdus iliacus and some ecological correlates // Ornis Scand. 1989. № 20. P. 1-12.
Jellis R. Bird sounds and their meaning. Cambridge, 1977. 256 p. Marler P. The voice of the chaffinch and its function as a language // Ibis. 1956b. № 98. P. 231-261. Nottebohm F. The "critical period" for song leaning in birds // Ibis. 1969a. № 111. P. 386-387.
Nottebohm F. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs., 1967. - P. 265-280.
Sick H. Ueber die Dialektbildung beirn Regenruf des Buchfinken // J. Ornit. 1939. № 87. P. 568-592.
Slater P.J.B., Ince S.A. Cultural evolution in chaffinch song // Behaviour. 1979. № 71. P. 146-166.
Slater P.J., Ince S.A., Colgan P.W. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. 1980. № 75. P. 207-218. Slater P.J.B., Clement F.A., Goodfellow D.J. Local and regional variations in chaffinch song and the question of dialects // Behaviour. 1984. № 88. P. 76-97.
Thielcke G. Stammesgeschichte und geographische Variation des Gesanges unserer Baumläufer // Verh Ornithol. Ges Bayern. 1961. № 14. P. 39-74.
Thorpe W.H. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. 1958. № 100. P. 535-570.
Thompson W.L. Song variation in a population of indigo buntings // Auk. 1970. № 87. P. 58-71. Witherby H.F., Jourdain F.C.R., Ticehurst N.F., Tuccker B.W. The Handbook of British Birds. London, 1944. 156 p.
Principles of the analysis of structure of chaffinch song (Fringilla coelebs L.). Individual variability of song
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology,
Faculty of biology, Moscow State University of M. V. Lomonosov
e-mail: beme@nm.ru
chaffinch@bk.ru
Summary
Song of birds is subtle and flexible species specific feature, like as the morphology signs. Species specific song of chaffinch (Fringilla coelebs L.) can be judged as precise character in the image on sonograms. The song has the certain structure and shares (divides) on the parts, elements, which consists of them. Individual variability of songs of one type will consist in unique singing each of chaffinch males, who form a local population.
Key words: chaffinch song, species specific features, individual variation of song, parts of song, polymorphism of song, development of song.
The typological organization and microgeographical variability of chaffinch song (Fringilla coelebs L.) in the population of the Curonian spit
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology,
Faculty of biology, Moscow State University of M. V. Lomonosov
e-mail: beme@nm.ru
chaffinch@bk.ru
Singing for the majority of sparrow birds - are the integral feature of their ability to live determining and building preparation and process of reproductive cycle (Lukanus, 1907; Darling, 1938 from Simkin, 1972). Some researchers believe, that the song of birds serves (means) not only for attraction of female, but carries out also functions for scaring away contender (other male), and representation (advertising) of the territory of nest, relative definition of habitat borders (Catchpole, Slater, 1995). Thus spring singing represents the multifunctional phenomenon (Simkin, 1972) and can carry various value. At detailed studying of song repertoire of many sparrow species of bird it was revealed, that the individual has not only one, and also some number types of songs. Variants or types of species song are individually various and distributed between individuals of a population (Slater et al., 1980; McGregor, Krebs, 1982). In many works proves to be true distinction of species specific song of birds at individuals of different local and geographical populations (Marler, Tamura, 1962; Thielcke, 1969; Baptista, King, 1980; Krebs, Kroodsma, 1980), also individual variability of song types at different males is shown, and these birds of one species inhabit in one local population in the certain territory (Hultsch, Todt, 1981; Thompson, 1970; Slater et al., 1980; Whitney, Miller, 1987).
Chaffinch (Fringilla coelebs L.) is classical object of studying of becoming of vocal repertoire (Thorpe, 1958; Marler, 1956; Nottebohm, 1967) and geographical variability species specific song in a population (Promptov, 1930; Sick, 1939; Thielcke, 1961; Simkin, 1983; Slater et al., 1984).
The purpose of the given work will consist in revealing the typological organization and microgeographical variability of song types of chaffinch in population of Curonian spit of Baltic Sea (Kaliningrad region). Material and methods
The basic part of work was conducted in area of biological research station "Fringilla" on Curonian spit of Baltic Sea (Kaliningrad region), where the density of a population of chaffinch (Fringilla coelebs L.) was the greatest. Also were made tape records of chaffinch songs and in other areas of Curonian spit - settlement Sea (or Morskoy), settlement Fishing (or Ribachiy) and settlement Wood (or Lesnoy), which are removed from each other on the average on 10 km (fig. 1). To territorial distribution of chaffinch individuals was applied approach of search of birds groups (Simkin, 1983), and points of record are marked on map.
Thus, during the spring period (April - May, 2005) in territory of a pine forest with an alder and a birch (h =10-15 м) have been made tape records of actively singing chaffinch males (N=153). For record of songs tape recorder Panasonic RQ-SX95F and microphone Philips SBC ME570 have been used. In different points of territory we tried to fix complete repertoire of chaffinch male (from everyone were recorded on the average 20 songs). In the general (common) sample about three and a half of thousand songs of different types has been considered.
Further, sonograms of songs were analyzed with the help of program Avisoft-SaSLab Light. In this article are submitted results of measurements only on nine basic time-and-frequency parameters for each song type with the purpose of their comparison: duration of song types (sec) and number of elements in song, duration of elements in started singing, trills and a final stroke (sec), maximal, minimal and average (median) frequency (KHz) of song types, intervals between songs (sec). Traditionally the song of chaffinch (Fringilla coelebs L.) shares (divided) on three parts (on perception on hearing): sounds of whistle (started singing), trill sounds (an average part) and a final stroke (a sharp sound) (Marler, 1956; Thorpe, 1958). On sonograms within the limits of each part of song it is possible to allocate (distinguish) separate phrases (a parts, fluctuation), which consist of elements (syllables) similar in the form (fig. 2).
Types of songs have been designated (marked) by Latin letters. We considered songs of one type, if they had two or all three similar parts (phrase). For example, to song type A are belong songs А1 and А2 (fig. 2.1), and belong to type Е - songs Е1 and Е2 (fig. 2.2), which similar on two phrases (song parts). But the majority song types were similar on all three phrases (parts) (fig. 2.3). At record, songs of one type met in different points of territory (was considered as belong to repertoires of different males), therefore alongside with the Latin letter were designated (marked) by numbers in ascending order (for example, А1, А2, А3, etc.).
Fig. 1. Map of Curonian spit of Baltic Sea (Kaliningrad region).
Results and discussion
In a researched population of chaffinch we revealed 22 types of songs, 4 from which appeared rare (are found in 1-2 points of record) and 5 types of songs were combined (have consisted of phrases or parts of different types of songs). The average size of chaffinch repertoire on Curonian spit has been represented by 2-3 types of songs (max - 6 song types, min - 1 song type). Example of repertoire and sequence of execution (performance) of song types we give as an example of songs of one of chaffinch male, recorded in a point №14+:
R R E2 R E2 E2 E2 E2 R R R R R R R R R R R R R R R R R R E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 E2 R R E2 R R R R R R E2 E2 E2
In his repertoire are marked only 2 types of songs (R - 29 (56,9 %), Е2 - 22 (43,14 %), n = 51), which are executed (performance) almost with identical or similar frequency (fig. 3).
Parity (ratio) of song types in repertoires of different males of chaffinch (Fringilla coelebs L.) are reflected in table 1.
At the analysis and comparison of chaffinch songs on Curonian spit have been found out individual and microgeographical variability of the some song types.
Table 1. Parity (ratio) of song types in repertoires of chaffinch (Fringilla coelebs L.) in area of biological research station "Fringilla"
Number of maleQuantity of song types
in repertoireParity (ratio) of song types12С - 35% G - 65%22C*2 - 51% H1 - 49%32M - 43,5% N - 56,5%41N1 - 100%52Е1 - 21% О - 79%61С3 - 100%71С4 - 100%81О1 - 100%96О2 - 18% О3 - 13% Р - 5%
С5 - 16% I 2 - 45% А4 - 3%105А5 - 6% А6 - 6% О3* - 35%
О4 - 12% I 3 - 41%112Е2 - 56,9% R - 43,14%122Т - 58% S2 - 42%
The note: the characteristic of chaffinch repertoire was taken into account at sample volume of songs from one male on the average n> 20; songs of one type in different points of record were marked by numbers in ascending order.
Fig. 3. Song types of chaffinch, which are being in repertoire of one male (record in area of biological research station "Fringilla"): a - song type R, b - song type Е2.
Individual variability of song types
At the analysis of songs of one type within the limits of one micropopulation (some groups) of chaffinch on Curonian spit some distinctions in basic parameters of these song types are allocated (distinguished) (sometimes in their qualitative or phonetic features) - so-called individual variability (variants or a versions of songs of one type, recorded in different, close on distance, points in the certain territory (1 км2) - was considered tape records different males, or in repertoire of one male). Individual variability can be tracked at comparison of time-and-frequency parameters of the most widespread types of songs (A and С) in a population of chaffinch on Curonian spit (fig. 8). Probably, at other types of songs (them more than 20), found by us in a population, individual variability has other character. For songs of both types (A and С) the greatest distinctions are found out in duration songs (sec), quantity of syllables (though song type C is more stable in this parameter) and intervals between songs (sec) (tab. 2, 3). The minimal and maximal frequency (KHz) of songs varies insignificantly. Some distinctions in duration of syllables (elements) (sec) of separate parts in chaffinch songs (started singing, trills, and a final stroke) are observed.
Fig. 2. Principles of allocation (distinguish) of songs of one type in a population of chaffinch (Fringilla coelebs L.) on Curonian spit: 1а - song type А5, 1б - song type А6, 2а - song type Е1, 2б - song type Е2, 3а - song type I2, 3б - song type I6.
Table 2. The basic time-and-frequency parameters of song type A
Type of song (token)Number of songs (n)The name of a place of recordThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables in song typeLength of syllables in started singing, secLength of syllables in trill, secLength of syllables in final stroke, secIntervals between songs, secА11Trial point2,01141,7227,4073,962160,0563
±0,0060,136
±0,0240,1746
±0,0903А343
point2,36
±0,0261,5
±0,17,6
±0,464,2
±0,118,3
±1,150,04
±0,010,12
±0,010,145
±0,0111,039
±5,68А4*111
point1,86051,557,9244,306130,0425
±0,0090,099
±0,0070,136
±0,094А5512
point1,9712
±0,131,45
±0,157,85
±0,364,5
±0,2215
±0,70,0425
±0,0060,1
±0,020,092
±0,026,74
±2,27А6313
point2,0511
±0,161,5
±0,17,5
±0,64,306
±0,315
±10,042
±0,0060,11
±0,0060,14
±0,016,04
±1,06А7*113
point1,87211,7228,0964,478140,0435
±0,010,11
±0,0034,8704
The note: average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced of males chaffinch in the given points of record are specified; the strongest differences counted a difference of parameters >0,5 КHz in frequency and >0,02 sec in length (are allocated or distinguish by a font).
Table 3. The basic time-and-frequency parameters of song type C
Type
of song (token)Number of songs (n)Place of recordThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables in song typeLength of syllables in started singing, secLength of syllables in trill, sec *Length of syllables in final stroke, secIntervals between songs, sec1
phrase2
phraseС15Trial point2,025
±0,111,447
±0,1668,01
±0,374,134
±0,21516,7
±1,160,13
±0,010,084
±0,0280,13
±0,0070,085
±0,00511,61
±5,06С*2222
point1,917
±0,111,48
±0,147,88
±0,484,164
±0,21317,06
±1,160,199
±0,010,05
±0,0040,094
±0,0090,063
±0,0097,85
±5,06С3208
point2,074
±0,1181,627
±0,1319,698
±0,414,0996
±0,17317,25
±0,850,1203
±0,0050,068
±0,0030,1155
±0,0080,071
±0,0056,423
±1,9С4219
point2,01
±0,211,75
±0,159,25
±0,64,056
±0,2216,45
±1,370,122
±0,010,066
±0,0050,128
±0,010,074
±0,0087,06
±0,01С5611
point1,75
±0,151,7
±0,137,8
±0,54,3
±0,0713,7
±1,030,133
±0,0050,067
±0,0040,103
±0,0030,068
±0,0055,4
±0,85С6313
point1,96
±0,121,55
±07,522
±0,364,36
±0,215
±10,135
±0,010,0632
±0,0050,0913
±0,050,07
±0,0064,44
±0,91
The note: average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced of males chaffinch in the given points of record are specified; the strongest differences counted a difference of parameters >0,5 КHz in frequency and >0,02 sec in length (are allocated or distinguish by a font); * - the trill of song type C will consist of two phrases
Fig. 8. Individual variability of song types A (1) and C (2) in a population of chaffinch (Fringilla coelebs L.) on Curonian spit: (1) a - song type А1 (trial point), b - song type А3 (record in 3-d point), c - type А4 (11-th point), d - song type А5 (12-th point), e - song type А6 (13-th point), f - song type А7 (13-th point); (2) a - song type C (trial point), b - song type С*2 (record in 2-nd point), c - song type С3 (8-th point), d - song type С4 (9-th point), e - song type С5 (11-th point), f - song type С6 (record in 13-th point).+
Microgeographical variability
At the quantitative analysis of songs of one type in different areas of Curonian spit (fig. 1) is found out small microgeographical variability (in variants or the versions of songs of one type which has been recorded on different sites of Curonian spit - distance between them on the average on 10 km).
For example, variants of song type G (fig. 4) differed on length (sec), number of syllables (elements) and on intervals between songs (sec). Some difference of values and in the maximal frequency (KHz), length of syllables (elements) in started singing (sec) and a final stroke of songs of this type (tab. 4) is found.
Fig. 4. Sonograms of variants of song type G, recorded in different areas of Curonian spit: a - song type G1 (record in a settlement Fishing or Morskoy), b - song type G2 (record in a settlement Wood or Lesnoy), c - song type G3 (record in 33-d point, "Fringilla").
Song type M of chaffinch was very interesting for the qualitative analysis (fig. 5). A variety of this song type is observed within the limits of one area (individual variability), and on different sites of Curonian spit (microgeographical variability). It is possible to consider as a combination of individual and microgeographical variability.
On hearing three parts of songs of this type are perceived differently (differ from each other): first two phrases М1, М2, М4 (fig. 5а, 5e, 5f) sound as started singing (a number or row sounds of whistle), while in song types М5, М11, М16 (fig. 5c, 5b, 5d) these elements are in the middle of song and are heard as a trill. It is interesting to note, in song type М16 (fig. 5d) the started singing ("sounds of whistle") is duplicated by two identical phrases, and last element of these phrases does not sound as thin sound of whistle, and more likely - as sharp "gnash". Nevertheless, this element ("gnash") enters into the first part of song, but first phrase at the majority of song types is submitted by sounds of higher tone (started singing). Trill elements of songs of this type differ on the form and somewhat on motif (a phrase which is taking place before a final stroke): at some song types the accent at singing is done (made) on the top bend of elements of a trill (М2, М5, М11), at others - on the bottom bend (М1, М4, М16). Fig. 5. Sonograms of variants of song type M, which have been recorded in different areas of Curonian spit: a - song type М1 (record in 11-th point, "Fringilla"), b - song type М12 (record in 78-th point, "Fringilla"), c - song type М5 (record in a settlement Sea or Morskoy), d - song type М16 (record in a settlement Fishing or Ribachiy), e - song type М2 (record in a settlement Wood or Lesnoy), f - type songs М4 (record in a settlement Wood or Lesnoy).
Table 4. The basic time-and-frequency parameters of song type G
Type
of song (token)Number of songs (n)Place of recordThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables in song typeLength of syllables in started singing, secLength of syllables in trill, sec *Length of syllables in final stroke, secIntervals between songs, sec1
phrase2
phraseG18Settlement Fishing2,72
±0,181,47
±0,098,65
±0,673,89
±0,12518,2
±0,970,132
±0,130,076
±0,0040,165
±0,0080,11
±0,014,94
±1,15G228Settlement
Wood2,3
±0,211,54
±0,147,9
±0,493,76
±0,17 16,2
±1,80,106
±0,0150,064
±0,0050,156
±0,0120,13
±0,0259,43
±8,14G328Station
"Fringilla"2,46
±0,021,58
±0,0778,23
±0,63,72
±0,094 17
±00,12
±0,0140,067
±0,0020,15
±0,0040,12
±0,0068,64
±0,8
The note: average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced of males chaffinch in the given points of record are specified; the strongest differences counted a difference of parameters >0,5 КHz in frequency and >0,02 sec in length (are allocated or distinguish by a font); * - the trill of song type G will consist of two phrases.
Thus all these "dialect" forms of song type M are found in repertoires of chaffinch male at one settlement (on one site), and in micropopulations of different areas of Curonian spit (in settlements Wood, Sea, Fishing, remote on 10 km from each other). Geographical variability of song of different species of sparrow birds (Passeriformes) and forming song "dialects" (so-called "local songs") were discussed in many works (Baker, 1975; Lemon, 1975; McGregor, 1980; Mundinger, 1982). Thus, microgeographical variability of time-and-frequency parameters of songs of one type in a researched population of chaffinch appeared insignificant. Probably, in conditions of high density of chaffinch population on Curonian spit, males formed songs without obvious deviations in basic parameters in the "communicative" groups.
The mixed (combined) song type
At the analysis of chaffinch songs on Curonian spit have been marked cases of formation new song forms as a result of a combination of phrases of different types of songs (fig. 6). Earlier at chaffinch (Fringilla coelebs L.) similar cases have been described only for the individuals, which have been brought up in experimental conditions, when caught young male in sensitive period of song learning (early period of life), and bird hear played by tape daily different songs of two others wild males. In the result, after some time the caught bird sang a song uniting phrases of these other males (Nottebohm, 1967; Jellis, 1977). Probably, also at us marked cases are the certificate of non-standard learning of song types during their crystallization (formation, an establishment).
It is known, that at experiment of song learning on song sparrows (Melospiza melodia), having 5-11 types of songs in repertoire of male, very small percent belongs to individuals, which reproduced combinations of elements from various song types of teachers (of older males), and even some song types have been completely invented by them. But such song forms were individual and are not socialized (not shared) in group of males, that created conditions of ignoring unique (Burt, Beecher, 2000). Similar experiments can confirm ability of sparrow birds (Passeriformes) to improvisation at singing song types. And M. Beecher and his colleagues (Beecher, 1996) says, that development of sparrow song can differ in natural (field) and laboratory conditions.
It is interesting to notice, a case of improvisation is found in our researches at execution (performance) of types of songs (mixing, a combination of different phrases (or parts of songs) from one repertoire) by male of chaffinch, which inhabited on suburb (edge of territory) of population in Curonian spit (in sites with more rare vegetation of young alder) and had the maximal (six) quantity of types species specific songs (fig. 7).
Conclusions
Thus, in a population of chaffinch on Curonian spit has been revealed the typological organization of species song and insignificant microgeographical variability of songs of one type in the basic quantitative (time-and-frequency) parameters. Distinctions on qualitative characteristics of some song types of chaffinch are found at a combination of individual and geographical variability of these song types (in the form of syllables (elements) on sonograms, on syntax of song structures - sequence of phrases, elements, their combinations).
The song of chaffinch has a genetic basis (species-specific features) (Thorpe, 1958; Nottebohm, 1967), but at training (song learning), improvisation at singing, and also at mistakes of copying at vocal training (learning) constantly replenishes a variety of types of songs in a population (Slater et al., 1979, 1984 ; Ince et al., 1980; Jellis, 1977). The variability of song types, which found out by us, can testify to genetic heterogeneity of a population and microevolutionary processes in chaffinch population. Fig. 6. A combination of phrases (parts) of song types, which have been recorded in different areas of Curonian spit. Not combined song types: 1а - song type Е2, 1b - song type М12, 2а - song type Е5, 2b - song type I2, 3а - song type О3 (record in area of biological research station "Fringilla"), 3b - song type С*13 (record in a settlement Sea or Morskoy). The combined song types: 1c - song type ЕМ, 2c - song type ЕI (record in a settlement Wood or Lesnoy), 3c - song type ОС* (record near biostation "Fringilla").
Fig. 7. Improvisation at singing (a combination of different types of songs) in repertoire of one male of chaffinch (record in 13-th point in territory of biological research station "Fringilla"): a - song type О5, b - song type С6, c - song type ОС6 (consist of phrases or parts of two different song types, which are included in repertoire of male).
Gratitude
Authors express deep gratitude to employees of a biological research station "Fishing" of zoology institute of the Russian Academy of Science on Curonian spit for the help and support in work, gratitude to professor G. N. Simkin for valuable advice at the analysis of a material. Work is executed at financial support of the Russian Federal Property Fund, the grant 04-04-49276. +
The literature
Baker M. C. 1975. Song dialects and genetic differences in White-crowned Sparrows (Zonotrichia leucophrys) // Evolution. № 29. 226-241.
Banin D. A., Byome I. R., Byome R. L. 1994. Seasonal variability of functional loading songs of birds // The Bulletin of the Moscow university. Vol. 16. Biology, 3. Moscow: 40-42.
Baptista L. F., King J. R. 1980. Geographical variation song and dialects of montane white-crowned sparrows // Condor. № 79. 356-370.
Barber D. R. 1959. Singing pattern of the common chaffinch, Fringilla coelebs Linn. // Nature. № 10. P. 129.
Beecher M. D. 1996. Birdsong learning in the laboratory and field // Ecology and evolution of acoustic communication in birds / Eds D.E. Kroodsma, E.H. Miller. Itaca. London. P. 61- 78.
Burt J. M., Beecher M. D. 2000. Social influences during song development in the song sparrow: a laboratory experiment simulating field conditions // Animal Behaviour. № 59. 1187-1197.
Catchpole C.K., Slater P.J.B. 1995. Birdsong. Biological themes and variation// Cambridge: Cambridge University Press. 248 p.
Hultsch H., Todt D. 1981. Repertoires sharing and song-post distance in nightingales (Luscinia megarhynchos B.) // Behav. Ecol. Sociobiol. № 8. 183-188.
Ince S. A., Slater P. J. B., Weismann C. 1980. Changes with time in the song of a populations Chaffinches // Condor. № 82. 285-290.
Jellis R. 1977. Bird sounds and their meaning. Cambridge. 256 p. Krebs J. R., Kroodsma D. F. 1980. Repertoires and geographical variation in bird song // Adv. In Study Behav. № 11. 143-177.
Lemon L. E. 1975. How birds develop song dialects // Condor. № 77. 385-406.
Marler P. 1952. Variation in the song of the Chaffinch Fringilla coelebs // Ibis. № 94. 458-472. Marler P. 1956b. The voice of the chaffinch and its function as a language // Ibis. № 98. 231-261. Marler P., Tamura M. 1962. Song dialects in three populations of White-crowned Sparrows // Condor. № 64. 368-377.
McGregor P. K. 1980. Song dialects in the corn bunting (Emberiza calandra) // Z. Tierpsychol. № 54. 285-297.
McGregor P. K., Krebs J. R. 1982. Song types in a population of great tits (Parus major): their distribution, abundance and acquisition by individuals // Behavioiur. № 79. 126-152.
Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds // Acoustic communication in birds / Eds. D.E. Kroodsma, E.H. Miller. New York. P. 147-208.
Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs.. P. 265-280.
Promptov A. N. 1930. Geographical variability of chaffinch song in connection with the common questions of seasonal flights of birds // Zool. journ. № 10 (3): 17-40.
Sick H. 1939. Ueber die Dialektbildung beirn Regenruf des Buchfinken // J. Ornit. № 87. 568-592.
Simkin G. N. 1972. About biological value of bird singing // The Bulletin of the Moscow university. № 1. Moscow: 34-43. Simkin G. N. 1983. The typological organization and population phylogeny of bird songs // Bulletin of Moscow community investigators of nature. Section of biology. V. 88. № 1. Moscow: 15-27.
Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. 207-218. Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects // Behaviour. № 88. 76-97.
Slater P. J. B., Ince S. A. 1979. Cultural evolution in chaffinch song // Behaviour. № 71. 146-166.
Thielcke G. 1961. Stammesgeschichte und geographische Variation des Gesanges unserer Baumläufer // Verh Ornithol. Ges Bayern. № 14. 39-74.
Thielcke G. 1969. Geographic variation in bird vocalizations // Bird Vocalizations / Eds. R.A. Hinde. London, New York. P. 311-340.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. 535-570.
Thompson W. L. 1970. Song variation in a population of indigo buntings // Auk. № 87. 58-71. Whitney C. L., Miller J. 1987. Song leaning in the wood thrush // Can. J. Zool. № 65. 1038-1042.
Typological organization and microgeographical variability of chaffinch song (Fringilla coelebs L.) in population of Curonian spit
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology
Faculty of biology, Moscow State University of M. V. Lomonosov
e-mail: beme@nm.ru
chaffinch@bk.ru
Summary
Song repertoire of many species sparrow (Passeriformes) has not only one, but some types of song. Variants or types of species specific song are individually various and distributed between individuals of a population. Traditionally the chaffinch song (Fringilla coelebs L.) shares on three parts: the row of whistle sounds (started singing), sounds of trill (an average part) and a final stroke. We carried songs of two or all three similar parts to songs of one type. In the investigated population of chaffinch (Fringilla coelebs L.) us it has been revealed 22 types of songs, 4 of which appeared rare (are found in 1-2 points) and 5 types of songs were combined (have consisted of phrases of different types of songs). The average size of chaffinch repertoire on Curonian spit of Baltic Sea has been represented by 2-3 types of songs (max - 6 types of songs, min - 1 type of song). Insignificant microgeographical variability of songs of one type has been revealed at their qualitative and quantitative analysis. At chaffinch the song has a genetic basis (species-specific features), but by training (learning), improvisation, mistakes of copying a variety of types of songs in a population constantly replenishes. The variability of song types considered by us can testify to genetic heterogeneity of a population and microevolutionary processes which in it occur.
Key words: song of bird, types of species specific song, song behavior, song variability, geographical variation of song, microevolution of bird's song, song cultures in population. +
Vocal variability and dialect forms of chaffinch song (Fringilla coelebs L.) in populations of the central part of the European Russia
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology,
faculty of biology, Moscow State University of M. V. Lomonosov
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
In early works geographical variability of song at many species of birds has been marked as a phenomenon of differentiation of song structures or song patterns (as the image on sonograms) in different local populations (Marler, 1952; Marler, Tamura, 1962; Sick, 1939; Poulsen, 1951). Local variants of vocalizations of birds can be considered as analogy of human speech and to name as dialects. The word "dialect" has taken place from linguists and means a local speech variant, characteristic for region. Borders of a dialect define lexical (word structure), morphological (an accent, structure) and phonetic (pronunciation) characteristics of the dictionary (Kurath, 1972).
Song dialect of birds is a variant of the traditional form of song, shared by members of a local population of birds and it forming borders of a dialect (separating from other variant of song pattern), within the limits of which there is a traditional training (song learning) of characteristic song components in the given population. But thus big song repertoires of some species of sparrow birds interfere (have problem) with objective definition of dialect borders (Thielcke, 1969; Kreutzer, 1974; Kroodsma, 1974; Baker, 1975; Baptista, 1975; Lemon, 1975; Payne, 1981; Mundinger, 1980, 1982). Chaffinch (Fringilla coelebs L.) is classical object of study of becoming of vocal repertoire (Thorpe, 1958; Marler, 1956; Nottebohm, 1967) and of geographical variability of species specific song in a population (Promptov, 1930; Sick, 1939; Thielcke, 1961; Simkin, 1983; Slater et al., 1984). Regional variability of song can be determined in qualitative aspect (the form of a syllable, syntax) and quantitative (time-and-frequency) parameters. The concept of a dialect should be closely (cautiously) and is carefully argued at establishment of regional variability of call (voice) and of song patterns.
Material and methods One of problems of our researches has consisted in revealing macrogeographical distinctions of chaffinch song (Fringilla coelebs L.) in different local populations, which are removed approximately on 1000 km from each other. In northwest (Curonian spit, the Kaliningrad region) and in central parts (Zvenigorod, Moscow, Michurinsk) of the European Russia have been made tape records of singing males (N=218) during the spring-and-summer period of 2005-2006. Sonograms of types of songs have been analyzed with the help of computer program Avisoft SASLab Light. In total have been analyzed about five thousand songs. Types of songs have been marked by Latin letters. At record, songs of one type have been met in different points of territory (considered that belong to repertoires of different males), therefore alongside with the letter have been designated by numbers in ascending order (for example, А1, А2, А3, etc.).
For record of songs have been used tape recorder Panasonic RQ-SX95F, condenser microphone Philips SBC ME570. To territorial distribution of individuals has been applied group approach (Simkin, 1983) and in parallel - points of record have been marked on map.
At the analysis of song sonograms of chaffinch basically have been applied two qualitative methods: revealing (detection) of phonetic distinctions (frequency of a sound, its form on sonogram) or of ways of a pronunciation of syllables of the phrases making songs; revealing of lexical distinctions (changes of phrases of songs as a whole) (Mundinger, 1982). Also have been spent the quantitative analysis of dialect forms of songs of one type and comparison of their basic time-and-frequency parameters (duration of songs - sec, number of elements in song type, duration of syllables in started singing, trills, in final stroke, the maximal, minimal and average frequency of songs - KHz, intervals between songs -sec).
Result and discussion In populations of the central part of the European Russia (N=65 of males) we allocate (distinguish) 15 types of song of chaffinch (Fringilla coelebs L.), which have been completely similar in structure or have been in part modified in the syllabic form (on sonograms) in comparison with songs of same types in samples of a northwest part of the European Russia (N=153 of males). For seven types of song from twenty two (tests on Curonian spit) analogies (similarities) have not come to light - probably, owing to their rarity. Many song types in the central part of the European Russia have been considered as combined phrases (parts) of syllabic patterns known to us, but frequently with the changed figure (forms) on sonogram.
In result, have been found 12 dialect forms of songs of one type, which appeared similar in base structure of elements, of phrases, but in different regions of Russia frequently had distinguished manners, ways of their performance at singing (phonetic aspect). The average size of chaffinch repertoire (Fringilla coelebs L.) in populations of the central part of the European Russia from statistical calculations has been submitted 1,93 ± 0,22 types of song (max - 4 types of song, min - 1 song type) at a capture (volume) on the average 24,7 ± 11,3 songs from one of male. As a bright example of vocal variability of chaffinch (Fringilla coelebs L.) it is possible to result (to show) some local variants of songs, which we have attributed to one type (fig. 1). Samples of types of song, which will be resulted (be shown), were not single in a population and have been recorded in repertoires more, than 1-2 individuals. In Zvenigorod (the Moscow region) (fig. 1.2) a started singing (whistle elements) of songs such as type V has shorter second subelements, which are in the top part concerning the first subelements. Elements of a trill are more complex (difficult) in Zvenigorod. A final stroke - three large base elements are present as at samples of northwest, and of the center of the European Russia. But on Curonian spit (fig. 1.1) before last element (7) of final stroke there are three small elements (4, 5, 6) by higher frequency. Probably, it - phonetic components of a dialect, which characterize style of singing (song culture) in a local population on the given song type. In samples of Zvenigorod (fig. 2.2) in songs such as type S the first phrase of a trill is unusual (includes 3 elements), the final stroke is twisting and begins by small elements (can be - two incomplete arches). We shall assume that it is dialect forms of song type S. It is necessary to note, that the given forms of songs have been met at record of males in several points of territory, and that is these copies (songs of one type) are not individual in their local populations. In samples also types of song have been found, which are difficult for differentiating among themselves because of their similarity (fig. 3).
We have attributed the given samples of songs to different song types (D, F, G), but nevertheless they are similar among themselves in base structure of elements (especially in the started singing and trill), which nevertheless are performance at singing by different manners (ways), that makes their distinct or different. Probably, it - the initial forms of the further modification, and it represent insignificantly transformed forms for precise revealing of type to which are attributed.
The song type J is difficult for the analysis because its structure can be various both in one local population, and in different regions. The started singing frequently is made by the phrase of "V-shape" elements with the subsequent powerful subelements in force of a sound. In a trill usually large "arc-similar" elements (hardly differ on structure in northwest and in the center of the European Russia), before it there can be a phrase of thin elements - it is sometimes are heard as started singing (fig. 4.2).
Table 1. Character of vocal variability of chaffinch songs (Fringilla coelebs L.) in populations of northwest and the center of the European part of Russia (N = 218) *
Vocal variabilitySong type (token)ABCDEFGIHJKLMNOPQRSTVWUTotalDialects1+++++5Subdialects2+++++++7European part of RussiaSample size of song type (n) in populationNorthwest185381171610226133 1208181124171428183Center8824351111225223393
The note: 1dialects - different phonetic norms of song types; 2subdialects - small phonetic distinctions of song types; * - sample size of chaffinch males (Fringilla coelebs L.) at record; the most widespread (frequently met) in populations types of songs (with sample n> 15) are allocated (signed) by a font.
Table 2. The basic time-and-frequency parameters of local variants of song type B (fig. 5)
Type
of song (token)Number of songs (n)Place of recordThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables in song typeLength of syllables in started singing, secLength of syllables in trill, sec *Length of syllables in final stroke, secIntervals between songs, sec1
phrase2
phraseВ58Curonian spit (Kaliningrad region)2,47
±0,1041,57
±0,147,19
±0,323,77
±0,1424,5
±0,930,056
±0,0030,043
±0,0030,14
±0,010,06
±0,0075,32
±1,9В4Zvenigorod (Moscow region)2,48
±0,361,68
±0,167,92
±0,313,83
±0,3622,25
±2,990,073
±0,0050,04
±0,0030,17
±0,0080,075
±0,018В810Michurinsk
(Tambov region)2,34
±0,151,62
±0,098,044
±0,534,12
±0,2922,3
±2,60,068
±0,0030,034
±0,0060,127
±0,0130,09
±0,0164,3
±0,59В710Michurinsk
(Tambov region)2,99
±0,111,67
±0,087,63
±0,083,79
±0,1827,4
±1,170,05
±0,0050,033
±0,0030,15
±0,0070,084
±0,0227,06
±0,19
The note: average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced of males chaffinch in the given points of record are specified; the strongest differences counted a difference of parameters >0,5 КHz in frequency and >0,02 sec in length (are allocated or distinguish by a font); * - the trill of song type B will consist of two phrases.
Table 3. The basic time-and-frequency parameters of local variants of song type С (fig. 6)
Type
of song (token)Number of songs (n)Place of recordThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables in song typeLength of syllables in started singing, secLength of syllables in trill, sec *Length of syllables in final stroke, secIntervals between songs, sec1
phrase2
phraseС320Curonian spit2,074
±0,1181,627
±0,1319,698
±0,414,0996
±0,17317,25
±0,850,1203
±0,0050,068
±0,0030,1155
±0,0080,071
±0,0056,423
±1,9С315Zvenigorod
(Moscow region)2,59
±0,161,73
±0,238,15
±0,513,93
±0,2318,7
±0,90,15
±0,0070,052
±0,0050,15
±0,0340,1
±0,01410,06
±1,7С213Michurinsk
(Tambov region)2,98
±0,152,01
±0,367,5
±0,74,02
±0,222
±1,410,13
±0,010,048
±0,0070,12
±0,0150,084
±0,0056,05
±5,17С74Moscow2,24
±0,21,68
±0,169,39
±0,14,44
±0,1716,25
±1,890,14
±0,0080,06
±0,0030,15
±0,0040,1
±0,0768,8
±5,66С#113Moscow2,73
±0,31,78
±0,18,096
±0,64,54
±0,4320
±1,730,22
±0,0060,076
±0,0030,12
±00,081
±0,0198,26Cо410Zvenigorod
(Moscow region)2,39
±0,121,41
±0,17,75
±0,244,48
±0,2112,3
±0,480,21
±0,03-0,12
±0,010,1
±0,0856,54C*112Curonian spit2,55
±0,0821,3
±0,377,751
±04,478
±019
±1,40,175
±0,0070,034
±0,010,084
±0,0010,0915
±0,00222,9
The note: average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type which were reproduced of males chaffinch in the given points of record are specified; the strongest differences counted a difference of parameters >0,5 КHz in frequency and >0,02 sec in length (are allocated or distinguish by a font); * - the trill of song type С will consist of two phrases.
Fig. 1. Dialect forms of song type V: 1 - song type V 1 (record on Curonian spit, the Kaliningrad region); 2 - song type V (record in Zvenigorod, the Moscow region).
Fig. 2. Dialect forms of song type S: 1 - song type S5 (record on Curonian spit, the Kaliningrad region); 2 - song type S (record in Zvenigorod, the Moscow region).
The final stroke also varies in a manner of execution (performance) at singing: it can have at the beginning two small "arc-similar" element "^^" (fig. 4.1, 4.2, 4.3), which in samples of the central part of the European Russia frequently are absent (fig. 4.4, 4.5, 4.6), an average element of the final stroke - such as "a low hill" (it is in all songs such as type J, and in the greater degree it characterizes); last element of the final stroke has precise variability in different regions: in the center of the European Russia it larger (fig. 4.4, 4.5, 4.6). Probably, the features of structure of elements marked by us reveal dialect forms of song type J in phonetics of a trill and of a final stroke.
Despite of stability in a manner of performance of song type B, in different local populations on sonograms it is possible to note some variability of the final stroke (in the latter case (fig. 5.4) the number of the first elements of a stroke is increased, and the phrase is heard as a trill). In different areas of the European Russia penultimate elements of the final stroke differ: in northwest (fig. 5.1) they such as two arches (7, 8), and in the central part (fig. 5.2, 5.3, 5.4) are submitted by one big arch (5). Probably, given phonetic distinction (concerning to features of a pronunciation of elements, their forms, frequency on sonogram) of syllables of the final stroke is components of a dialect (local song cultures). Fig. 3. Difficultly differing types of songs at chaffinch (record on Curonian spit, the Kaliningrad region): 1 - song type D3; 2 - song type F4; 3 - song type G3.
Song type В7 (fig. 5.4) also it is possible to consider as the dialect form of song type B, because the morphological characteristic (an accent, structure) of this local song variant differs from others (phrases are similar under the form of syllables, but they are sung by other style). Thus, dialect forms of songs of one type can meet within the limits of one local population.
At the quantitative analysis of local variants of song type B (with the help of computer program Avisoft-SASLab Light) come to light some distinctions (differences) in values of the basic time-and-frequency parameters, but in the majority they are similar (tab. 2). From results of comparison of parameters of song type B it is visible (tab. 2), that the length of songs, number of their syllables, the maximal frequency, intervals between songs and length of elements of the second phrase of a trill have the values varying in wide enough range, frequently even beyond the insignificant difference established by us (differences are allocated or signed by a font). It is necessary to note, that the minimal frequency and length of syllables of the first phrase of a trill of songs of type B are appeared completely similar as in northwest (Curonian spit), and in the center of the European Russia (Zvenigorod, Michurinsk). For chaffinch (Fringilla coelebs L.) typically selective training (song learning) of species specific song in sensitive (perceiving) period (Marler, 1956; Thorpe, 1958; Nottebohm, 1967). Therefore young birds do not learn songs of other species, and they are guided by a time-and-frequency range of songs of the own species. Probably, as a result of such rigid genetic determination of species specific song (Simkin, 1983) is observed small variability of the basic time-and-frequency parameters of its different dialect forms (even at the account of a wide variation of values).
By full dialect of chaffinch song (Fringilla coelebs L.) in the certain territory it would be more correct to account the presence of dialect forms on all song types known to us in a population (originally 22 song types have been found), that, likely, it is almost impossible because of limitation of a sample of songs and constant mixing of vocal traditions of birds in different populations as a result of migrations (Slater, Ince, 1979, 1980; Espmark et al., 1989).
Fig. 4. Local variants of song type J (are recorded in different areas of Russia): 1 - song type J10 and 2 - song type J (record on Curonian spit, the Kaliningrad region); 3 - song type J (record in Zvenigorod, the Moscow region); 4 - song type J3 and 5 - song type J8 (record in Moscow); 6 - song type J12 (record in Michurinsk, the Tambov region).
Song type C (fig. 6) is the most widespread in all areas of the European Russia investigated by us. Probably, therefore the given song type has some updating (modifications, variants) in a manner of execution (performance) of phrases at singing: simple song type C (fig. 6.1, 6.2, 6.3, 6.4) is recorded in different areas of Russia; song type C* (fig. 6.5), song type С* (fig. 6.7) and song type C (fig. 6.6). Nevertheless, all these variants of songs on hearing are perceived as similar: the started singing - a number (line, row) of sharp (sometimes gnash or creaking) whistle sounds ("fuit-fuit-fuit"), a trill consist of two phrases - the first phrase is heard as a number (line) of more thin sounds, the second phrase - tone of a trill is more powerful ("til-til-til* tel-tel"), a final stroke - is short sharp rise and recession of a sound ("chi-kuik"). Fig. 5. Local variants of song type B (are recorded in different areas of Russia): 1 - song type В5 (record on Curonian spit, the Kaliningrad region); 2 - song type B (record in Zvenigorod, the Moscow region); 3 - song type В8 and 4 - song type В7 (record in Michurinsk, the Tambov region).
Whether it is possible these three forms of song type C (C, C#, C*) to account as dialect forms, even if they meet in one territory (for example, С7 and C#11 recorded in Moscow (fig. 6.4, 6.5), С3 and С*11 (fig. 6.1, 6.7) recorded on Curonian spit) - a complex (difficult) question. What origin of these variations (variants) of songs of one type - by wrong (incorrect) learning of birds and by further fastening of these songs in a population by training (song learning) of the following generations, or it is result of "introduction" of songs of migrants in local song culture - also it is possible to assume only. In Zvenigorod (the Moscow region) and Michurinsk (the Tambov region) there was a simplified variant of song type C (C0) (fig. 6.6) without the first phrase of a trill and having the simplified final stroke. Probably, it is result of improvisation at singing.
At comparison of the basic time-and-frequency parameters of local variants of song type C (tab. 3) the greatest difference had values of length of songs, the maximal frequency, number of syllables in song types, intervals between songs at singing (these parameters included rather wide range of distribution of the data). In other parameters (the minimal, average frequency, length of elements of a trill) come to light (are revealed) the most distinguished values (are allocated or signed by a font), but relative uniformity of the data nevertheless is observed.
Fig. 6. Local variants of song type C (are recorded in different areas of Russia): 1 - song type С3 (record on Curonian spit, the Kaliningrad region); 2 - song type С3 and 6 - song type Со4 (record in Zvenigorod, the Moscow region); 3 - song type С21 (record in Michurinsk, the Tambov region); 4 - song type С7, 5 - song type C#11 (record in Moscow); 7 - song type С*11 (record on Curonian spit, the Kaliningrad region).
Thus, similarity (in view of a variation of values in the certain range) of the basic time-and-frequency parameters of song type C (at presence of dialect forms) also can confirm their importance in transfer and maintenance (keeping, memory) of song traditions at chaffinch (Fringilla coelebs L.) in different populations of an area of distribution of this species. Many types of chaffinch song (Fringilla coelebs L.) in samples of the central part of the European Russia were considered by us as combined phrases of already known types of the song, which have been recorded on Curonian spit (the Kaliningrad region), but their elements have been modified on sonograms. It is possible to result (to show) some examples.
Fig. 7. Dialect forms of phrases of song types Т and S: 1 - song type Т (record on Curonian spit, the Kaliningrad region); 2 - the combined song type ТS (record in Michurinsk, the Tambov region); song type S (record on Curonian spit, the Kaliningrad region).
Trill elements of song type TS (fig. 7.2), which also met in samples of the center of the European Russia, are similar as to a trill of song type S (fig. 7.3.), and to a trill of song type Т (fig. 7.1) from a population on Curonian spit (but it is more similar - to a trill such as type S). The started singing - is from song type T. The final stroke - are similarities and to song type S, and to type T. But the first elements of the final stroke of song type TS of lower frequency, and its last element is shorter, than at song types Т and S. +
The started singing (whistle elements) and trill elements (the first phrase) of song type CМ 2 (fig. 8.2) are similar to song type С* (fig. 8.3) (to started singing and to the second phrase of a trill of "stick-similar" syllables). Elements of the second phrase of a trill of song type CM 2 are almost identical with a trill of song type M (fig. 8.1). The final stroke of song type CM has the general (common) structure both to song type M, and to song type С* (last element of which by "triangular" form, but it is not divided in subelements).
The started singing (whistle elements) and the trill of the combined song type GC (fig. 9.2) is similar to starting singing and to the trill of song type G (fig. 9.1), and the final stroke - similar to the final of song type С* (fig. 9.3) (but last element of this stroke of type GC hardly is separated on subelements).
Thus, in samples of different areas of Russia there can be the types of song combined from phrases of song patterns of other populations, but including the modified elements - it is possible to tell, their dialect forms. In samples of the central part of the European Russia the songs have been found which include phrases at once of two-three different types of song from other population of chaffinch (Fringilla coelebs L.) in the northwest of the European Russia. These phrases had a lot of similar in structure of elements, but nevertheless they differed in a manner of their execution (performance) at singing. If such songs of one type having the changed elements have the majority of individuals in a population, it is possible to assume about existence certain (determined) song cultures in these song types or in separate phrases.
Fig. 8. A combination of dialect phrases: 1 - song type М2 (record on Curonian spit, the Kaliningrad region); 2 - the combined song type СM 2 (record in Michurinsk, the Tambov region); 3 - song type С*13 (record on Curonian spit, the Kaliningrad region). +
Conclusions Many characteristics of singing of birds are precisely mentioned by social training traditions, receiving their vocal patterns by species specific imitation. Therefore the tradition, custom should be taken into account at the analysis of geographical variability of vocal behaviour.
In local populations are formed certain song cultures which are capable to change during time and can to make dialect forms on all area of distribution of a species. Steady song dialects during time - the phenomenon of conservatism of the vocal traditions, transmitted to the subsequent generations by means of vocal imitation. For many species the patterns of variability as purchases (changes) of vocal traditions are a product (result) of cultural evolution (Mundinger, 1980). Cultural evolution is a parameter for species with a historical variety of patterns of microgeographical variability, and also for species with differentiated syllables. But it is not a parameter in morphological variability (Lemon, 1975; Slater and Ince, 1979).
It is possible to allocate (to sign) 3 steps of hierarchy for the description of spatial distribution of the certain qualities of vocal variants (by way of reduction of scale of a category) (Mundinger, 1982):
1. Vocal norms (installation) (song institute) - regional populations of lexical variants;
2. Dialects represent local populations of the phonetic variants limited by the main ''isophone bundles'' (set of different phonetic norms) (''iso-'' - different, lat.);
3. Subdialects are the next or neighbour (close to each other) populations of phonetic variants having small set of isophone (that is a small variety of forms).
Fig. 9. Dialect forms of phrases of song types G and C: 1 - song type G6 (record on Curonian spit, the Kaliningrad region); 2 - song type GC (record in Michurinsk, the Tambov region); 3 - song type С*11 (record on Curonian spit, the Kaliningrad region).
Proceeding from samples, made by us in different regions of the European Russia (northwest and the central part), it is possible to draw a conclusion, that in some song types of chaffinch (Fringilla coelebs L.) exist subdialects (insignificant phonetic distinctions of syllables) - for example, on song types B, M, D, G, V, S, U, and dialects (different phonetic norms of types of song) - C, C# and C*, F, I, J, N, and there are also stable types of song (in samples of different regions are invariable) - song type A.
Vocal norms (installations), likely, can be judged by quantity of individuals in a population, which adhere to those or others song cultures (ways of singing) of different song types. If the majority of males sing any song type such certain style (special manner of performance), then it is will be vocal norm (installation) in the given song type (or in type of a phrase) in the given population. But it is difficult to define vocal norms of a population at repertoires of great volume (when song types much) (Kroodsma, 1974). Thus, in twelve types of song at chaffinch (Fringilla coelebs L.) the dialect forms (B C D F G I J M N V S U) have been found. It is interesting to note, that the dialect forms of songs or separate phrases (in the "combined" songs) of one type can have been met within the limits of one local population.
The basic time-and-frequency parameters of local variants of song types of chaffinch (Fringilla coelebs L.) at the quantitative analysis had insignificant macrogeographical variability that can speak about their importance in transfer and maintenance (keeping, memory) of specific song traditions of this species. The most stable parameters of local variants of songs of one type are appeared minimal and average (median) frequency (KHz), values of length (sec) and intervals of elements of a trill (sec).
Gratitude
Authors express sincere gratitude for the help in gathering a field material to employees of Zvenigorod biological research station of the Moscow State University and are deeply grateful to professor G. N. Simkin for numerous consultations and discussion of results of the carried out researches. Work is executed at financial support of the Russian Federal Property Fund, the grant 04-04-49276.
The literature Baker M. C. 1975. Song dialects and genetic differences in White-crowned Sparrows (Zonotrichia leucophrys) // Evolution. № 29. P. 226-241.
Baptista L. F., King J. R. 1980. Geographical variation song and dialects of montane white-crowned sparrows // Condor. № 79. P. 356-370.
Espmark Y. O., Lampe H. M., Bjerke T. K. 1989. Song conformity and continuity in song dialects of redwings Turdus iliacus and some ecological correlates // Ornis Scand. 1989. № 20. P. 1-12.
Kreutzer M. 1974. Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes) // Rev. Comp. Anim. № 8. P. 270-286.
Kroodsma D. E. 1974. Song learning, dialects, and dispersal in the Bewick`s Wren // Tierpsychol. № 35. P. 352-380.
Kurath H. 1972. Studies in Areal Linguistics. Bloomington. Indiana University Press. 127 p.
Lemon L. E. 1975. How birds develop song dialects // Condor. № 77. P. 385-406.
Marler P. 1952. Variation in the song of the Chaffinch Fringilla coelebs // Ibis. № 94. P. 458-472. Marler P. 1956b. The voice of the chaffinch and its function as a language // Ibis. № 98. P. 231-261. Marler P., Tamura M. 1962. Song dialects in three populations of White-crowned Sparrows // Condor. № 64. P. 368-377.
Mundinger P. C. 1980. Animal cultures and a general theory of cultural evolution // Ethol. Sociobiol. № 1. P. 183-223.
Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds // Acoustic communication in birds / Eds. D.E. Kroodsma, E.H. Miller. New York. P. 147-208.
Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs. P. 265-280.
Payne R. B. 1981. Population structure and social behavior: models for testing ecological significance of song dialects in birds. In "Natural Selection and Social Behavior: Recent Research and New Theory" / Eds R.D. Alexander, D.W. Tinkle. New York, Chiron. P. 108-119. Poulsen H. 1951. Inheritance and leaning in the song of the Chaffinch (Fringilla coelebs) // Behaviour. № 3. P. 216-228.
Promptov A. N. 1930. Geographical variability of chaffinch song in connection with the common questions of seasonal flights of birds // Zool. journ. № 10 (3): 17-40.
Simkin G. N. 1972. About biological value of bird singing // The Bulletin of the Moscow university. № 1. Moscow: 34-43. Simkin G. N. 1983. The typological organization and population phylogeny of bird's songs // Bulletin of Moscow community investigators of nature. Section of biology. V. 88. № 1. Moscow: 15-27.
Slater P. J. B., Ince S. A. 1979. Cultural evolution in chaffinch song // Behaviour. № 71. P. 146-166.
Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. P. 207-218.
Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects // Behaviour. № 88. P. 76-97.
Sick H. 1939. Ueber die Dialektbildung beirn Regenruf des Buchfinken // J. Ornit. № 87. P. 568-592.
Thielcke G. 1961. Stammesgeschichte und geographische Variation des Gesanges unserer Baumläufer // Verh Ornithol. Ges Bayern. № 14. P. 39-74.
Thielcke G. 1969. Geographic variation in bird vocalizations // Bird Vocalizations / Eds. R.A. Hinde. London, New York. P. 311-340.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. P. 535-570.
Vocal variability and dialect forms of chaffinch song (Fringilla coelebs L.)
in populations of the central part of the European Russia
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology,
faculty of biology, Moscow State University of M. V. Lomonosov
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Summary
Songs of birds can be considered as analogy of human speech. Thus to territorial distribution of song patterns frequently apply geographical methods of dialects. Share (differentiation), change of vocalization structure (songs, calls) of birds during time and space - is the known phenomenon in bioacoustics. But mechanisms of formation of dialect song forms of many bird's species are found insufficiently out. In given article local variants of song types of chaffinch (Fringilla coelebs L.), comparison of their basic time-and-frequency parameters are submitted. Attempt is made to clear a direction of cultural evolution of species specific chaffinch song.
Key words: birds song, species specific song, vocal norms (installation), development of song dialects, subdialects of species specific song, cultural evolution, local song culture, the changed phrases (parts) of songs, vocal polymorphism, types of species specific song and their variational line (row), the quantitative analysis, the qualitative analysis.
The typological organization of chaffinch song (Fringilla coelebs L.)
in population of Curonion spit
Astakhova О. A., Byome I. R.
Moscow state university of М. В. Lomonosov Biological faculty, department of vertebrate zoology
Russia, 119992 Moscow, Lenin mountains, 1/12
e-mail: chaffinch@bk.ru
beme@nm.ru
Chaffinch (Fringilla coelebs L.) is a traditional object of study by populational ecologists and bioacoustics. On this species the stages of becoming songs for the first time have been investigated, shown interpopulational and geographical distinctions of vocal repertoire, and also existence of vocal microgroupings has been proved (Sultanov, 1979, 1984; Gerasimova, 1981, 1983; Simkin, 1983; Joachim, Lauga, 1997). For chaffinch the mosaic structure of geographical variability of songs is characteristic that probably reflects genetic heterogeneity of separate settlements of birds, and also can be connected to processes of an exchange by individuals between the next populations.
Our researches have been carried out on territory of biological research station of zoology institute the Russian Academy of Science on Curonian spit of Baltic Sea (the Kaliningrad region), where chaffinch is a background nesting species, and its local population is characterized by high density. Study of populational variability of chaffinch songs earlier on a biological research station have been not spent (carried out). In different points of territory (area of biological research station "Fringilla") during the period from 15 April till 10 May 2005 have been made tape records of singing males (N = 102). Further, sonograms of songs have been analyzed with the help of computer program Avisoft SASLab Light. In total have been analyzed about three and a half thousand songs.
Traditionally the chaffinch song shares (divides) into three parts: the row of whistle sounds (started singing), sounds of trill (the basic part) and a final stroke. At the analysis of sonograms within the limits of each part the separate phrases consisting of similar elements in the form have been allocated (signed). We considered songs of one type that had two or all three similar parts.
In the investigated population of chaffinch 22 types of songs have been revealed, 4 from which appeared rare (are found in 1-2 points) and 5 types of songs were combined (have consisted of phrases of different types of songs). The average size of chaffinch repertoire on Curonian spit has been submitted by 2-3 types of songs (max - 6 types of songs, min - 1 type of song). In repertoire of one individual there could be both simple and complex (difficult) songs. At comparison of the basic time-and-frequency parameters in the investigated population of chaffinch insignificant individual and microgeographical variability of songs of one type is revealed.
Gerasimova R.I. 1981. Features of similarity and distinctions in individual song repertoires of chaffinch, incorporated into spatially separated groupings // Ecology and protection of birds. Theses of the report of VIII All-Union ornithological conference. Kishinev. P. 56.
Gerasimova R.I. 1983. To a question on formation of microdialects in partly isolated local groupings of chaffinch / In Behaviour of animals in communities. Materials of III All-Union conference on behaviour of animals. Moscow. Т. 2. P. 10-11.
Joachim J., Lauga J. 1997. Demes in habitat patches, the Chaffinch (Fringilla coelebs), example in south-western France / Presentation in conference.
Simkin G. N. 1983. The typological organization and population phylogeny of bird's songs // Bulletin of Moscow community investigators of nature. Section of biology. V. 88. № 1. Moscow: 15-27.
Sultanov E. G. 1985. Typological structure and geographical variability of songs at birds in Caucasus / Dissertation of biology science. - Baku: Academy of sciences at Azerbaijan, Institute of zoology. - 243 p.
Macrogeographical variability of chaffinch song (Fringilla coelebs L.)
in populations of the European part of Russia
Astakhova O. A.
Moscow state university of M. V. Lomonosov
biological faculty, faculty of vertebrate zoology
Russia, Moscow 119992, Lenin mountains, 1/12
e-mail: chaffinch@bk.ru
Chaffinch (Fringilla coelebs L.) is classical object of study of becoming of vocal repertoire (Thorpe, 1958; Marler, 1956; Nottebohm, 1967) and geographical variability of species specific song in a population (Promptov, 1930; Sick, 1939; Thielcke, 1961; Simkin, 1983; Slater et al., 1984). Macrogeographical variability relates to distinctions between populations, which are divided (shared) by the big distances or other barriers and have wide inhabitancies (areas) that complicates their crossing (mixing) genes or their vocal traditions (geographical populations within the limits of a species).
Song dialect of birds is a variant of the traditional form of song, divided (shared) by the members of a local population of birds and it forming borders of a dialect (separating from other variant song pattern) within the limits of which there is a traditional training (song learning) of song component, characteristic in this population . But thus big song repertoires of some sparrow bird's species interfere (have problem) with objective definition of dialect borders (Marler and Tamura, 1962; Thielcke, 1969; Kreutzer, 1974; Kroodsma, 1974; Baker, 1975; Baptista, 1975; Mundinger, 1980, 1982).
The purpose of our researches was definition of macrogeographical distinctions of song types of chaffinch (Fringilla coelebs L.) in different local populations, distance between which about 1000 km. In northwest and central parts of the European Russia have been made tape records of singing males during the spring-and-summer period of 2005-2006 (N=218 of males). Sonograms of song types have been analyzed with the help of computer program Avisoft SASLab Light. In total have been analyzed about five thousand songs. At the analysis of sonograms basically two qualitative methods have been used: revealing of phonetic distinctions (frequency of a sound, its form on sonogram) or ways of a pronunciation of syllables of the phrases making songs; revealing of lexical distinctions (changes of phrases or parts of songs as a whole) (Mundinger, 1982). But sometimes there could be difficulties at differentiation of new lexical variants of songs from extreme phonetic variants of known syllabic types (elements). At the quantitative analysis of songs of different populations distinctions of the basic time-and-frequency parameters have come to light. Duration of chaffinch songs different types could vary widely: 1,97±0,23 sec in northwest (Curonian spit, the Kaliningrad region) (N=279 of songs) and 2,59±0,29 sec in the center of the European Russia (Moscow, Zvenigorod, Michurinsk) (N=73 of songs). Frequency of song types in different populations of chaffinch appeared can more stable (northwest of the European Russia (Curonian spit): min=1,63±0,063 КHz, max=7,683±0,42 КHz, median=4,10±0,4 КHz; the center of the European Russia: min=1,7±0,17 КHz, max=8,06±0,58 КHz, median=4,096±0,29 КHz). Probably, this criterion of species specific song of chaffinch is important at training (song learning) and at transfer of vocal traditions of species, as it is characterized by small macrogeographical variability. The literature Kreutzer M. 1974. Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes) // Rev. Comp. Anim. № 8. P. 270-286.
Kroodsma D. E. 1974. Song learning, dialects, and dispersal in the Bewick`s Wren // Tierpsychol. № 35. P. 352-380.
Marler P. 1956b. The voice of the chaffinch and its function as a language // Ibis. № 98. 231-261. Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds // Acoustic communication in birds / Eds. D.E. Kroodsma, E.H. Miller. New York. P. 147-208.
Promptov A. N. 1930. Geographical variability of chaffinch song in connection with the common questions of seasonal flights of birds // Zool. journ. № 10 (3): 17-40.
Simkin G. N. 1983. The typological organization and population phylogeny of bird's songs // Bulletin of Moscow community investigators of nature. Section of biology. V. 88. № 1. Moscow: 15-27.
Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects // Behaviour. № 88. 76-97.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. 535-570.
The analysis of phrase combinations and various ways of syllabic singing at vocal variability of song types of chaffinch (Fringilla coelebs L.)
O. A. Astakhova, I. R. Byome
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Explanation of birds song variability shows, that traditional patterns of vocal behaviour do not exist equally with morphological features (more likely, as separate aspect). It partly due result of cultural evolution, which represents the important factor - as though a traditional heredity is developed, but, apparently, it not a parameter in morphological variability (Lemon, 1975; Slater, Ince, 1979; Mundinger, 1980). Basic (base) structure of birds vocalization represents fundamental, stabilized, species specific characteristic feature of vocalization (Mundinger, 1979). Many local populations of birds are characterized by the population specific image (on sonogram) of a vocal pattern (dialect), which has certain lexical (dictionary structure), morphological (an accent, structure) and phonetic (pronunciation) features (Kurath, 1972).
Thielcke G. (1965, 1969) defined dialects of birds as vocal variants with mosaic distribution. This mosaic definition one of the basic, which are widely used, but the majority of researchers of microgeographical variability not precisely enough assert in studying vocal distribution, whether there is a mosaic pattern (Kreutzer, 1974; Kroodsma, 1974).
Regional vocal variability of birds can be determined in qualitative aspect (the form of a syllable, syntax) and in quantitative parameters (frequency - KHz, time parameters - sec, min). For many species specific patterns of variability as purchases (having) of vocal traditions are products of cultural evolution. Cultural evolution is a parameter for species of birds with a historical variety of patterns of microgeographical variability, and also for species with differentiated vocal syllables (Mundinger, 1980). Many characteristics of a vocal of birds are precisely mentioned (related) by social training traditions (song learning), receiving their vocal patterns by specific imitation. Therefore the tradition, custom should be taken into account at the analysis of geographical variability of vocal behaviour.
Material and methods
Methods of geographical dialects can be applied to songs of birds. For example, sonograms can be used for definition of vocal variants, which are geographically distributed. Frequently (usually) two qualitative methods define (Mundinger, 1982):
* revealing of phonetic distinctions (phoneme) (frequency of a sound - KHz, its form on sonogram - (a pronunciation) - syllabic variability - isophones;
* revealing of lexical distinctions (lexicon) (dictionary structure - change of phrases of songs) - a regional lexicon - isolexes.
However, it is difficult to differentiate (to distinguish) new lexical variants (types of songs, their parts) from extreme (changed) phonetic variants ("figures" of a sound on sonograms) of known syllabic types (that are changed forms of song elements, making a variational line (row) of syllables or elements of one type). As it is difficult to define (determine) a limit at transition of variational lines (rows) of one song type of birds in a variational line (row) of other dialect (modified) song type, which "is obviously close, related" to the first on the origin - so-called "related" vocal lines (rows) at the given species of birds - probably, developed on the basis of the general (common), uniform, initial "ancient" vocal form.
In this work we show the qualitative analysis of song types on above stated methods and we shall try to reveal some features of cultural evolution of chaffinch song (Fringilla coelebs L.) in populations of the European Russia.
Tape records of male song have been made in northwest (Curonian spit, Baltic Sea) (N=153) and in the central part of the European Russia (Zvenigorod, Moscow, Michurinsk) (N=65), distance between which approximately 1000 km. Sonograms of song types were analyzed with the help of computer program Avisoft SASLab Light. In total about five thousand songs are analyzed. Song types were designated (marked) by Latin letters. The songs of one type, which has been recorded in different points (places) of territory were designated (marked) by numbers in ascending order (A1, A2, A3, etc.)
Results and discussion
According to methods of the qualitative analysis of song types - had been signed (allocated) by us two basic kinds of their distinctions: lexical and phonetic.
Lexical variability of chaffinch song types (Fringilla coelebs L.) - distinction of dictionary structure, change of phrases of song types. Proceeding (basing) from samples, by us it has been marked by four ways of its formation (are submitted below).
Replacement of a phrase of one kind by another in songs of one type at singing
In variants of song types A (fig. 1.1) and I (fig. 1.2), recoded by tape in different points of territory of Curonian spit (biological research station "Fringilla") - the kind of final stroke, characteristic for these types, is replaced with other, usually making song type C (fig. 1.3).
Fig. 1. The songs of one type distinguished (differ) by final strokes (record on Curonian spit, the Kaliningrad region): 1a - song type А8, 1b - song type А5; 2a - song type I 2, 2b - song type I 5; 3 - song type C.
By one type we considered the songs, having two or all three similar parts: started singing (the row of whistle elements), trill elements, final stroke (which they as can be subdivided into phrases - the elements similar under the form).
Addition of a phrase to songs of one type at singing
In song type Е1 (fig. 2.1a) a phrase of final stroke is added, and in type М16 (fig. 2.2b) a phrase is duplicated in started singing. Trills in variants of song type M differ because at singing there is an accent on different components of elements. Two such forms of a trill in song type M meet by equal frequency in territory of Curonian spit and characterize individual variability of songs of one type.
Fig. 2. The songs of one type differing by an additional phrase (record on Curonian spit): 1a - song type Е1, 1b - song type Е2; 2a - song type М12, 2b - song type М16.
Removal of a phrase from songs of one type at singing
Song type К2 (fig. 3.1a) it is possible to consider as the uncompleted form of song type I (fig. 3.1b), and song type Q (fig. 3.2a) - such as uncompleted song type D (fig. 3.2b) (on sonograms there is no final stroke). In "incomplete" song types last phrases on hearing are perceived as final stroke (in "full" forms they sound as trill) - therefore in separate, but rare types of songs have been allocated (signed).
Fig. 3. The samples of songs - similar on the first phrases, but they relate to different types (are recorded on Curonian spit in repertoires of different males): 1a - song type К2, 1b - song type В3; 2a - song type Q, 2b - song type D2.
Phrase combinations of different song types at singing
In repertoires of the some males there were the songs consisting of phrases of other types of songs (fig. 4). Similar cases are described from experiments song training when to caught male of chaffinch in sensitive period daily played different songs of two wild males. In the result, after a while the caught bird sang a song uniting phrases of songs of these two wild males (Nottebohm, 1967; Jellis, 1977). Probably, and by us the marked cases are the certificate of non-standard learning of song types during their crystallization (stabilization).
Such phenomena of lexical distinction of songs of one type within the limits of the certain population of some species sparrow many bioacoustics (scientists) explained from the point of view of natural variations in repertoire of the birds based on vocal generalization and individualization. The phenomena giving development of songs (developmental interaction) can include cooperating processes of imitation (copying) and improvisation (temporarily-dependent copying, a dispersion of young, drift, a regrouping of syllables, phrases) (Lemon, 1975; Burt, Beecher, 2000; Slater et al., 1984).
Fig. 4. Sonograms of the combined types of songs at chaffinch (Fringilla coelebs L.), recorded on Curonian spit (those song types - the phrases of which they consist, also are shown): 1a - song type А6, 1b - song type С5, 1c - song type AC; 2a - song type Е6, 2b - song type М1, 2c - song type ЕМ; 3a - song type О3, 3b - song type С*13, 3c - song type P (OC *).
By song birds are trained from old birds in the first year of a life (Thorpe, 1958), and training (song learning) occurs in most cases more, than from one individual. Various types of songs occur (origin) as a result: migrations, mistakes at training (copying) and improvisation. New song types can be considered as a new growth in a course of "cultural mutation". Distribution of song types in a population corresponds to the assumption, that birds copy them from other individuals of species at random, and that less than 15 % of these new growths influence creation of new song types. One individual of chaffinch male can have repertoire from 1-6 types of songs (Slater, Ince, 1979; Slater et al., 1980).
Thus, the examples of lexical distinctions of song types shown (resulted) by us can be based on mistakes of training and processes of improvisation at singing, that it is possible to name "cultural mutation" in limits of population song cultures. Such cases make small percent from all song types of a population.
Phonetic (syllabic) variability of chaffinch song types (Fringilla coelebs L.) - distinctions in a manner (ways, styles) of performances, pronunciations of syllables, of elements of one type - distinction of their forms on sonograms). We considered change of the form (of pronunciation) of elements of one type in different parts of songs: started singing (the row of whistle elements), a trill (an average part) and final stroke.
Syllabic variability in started singing of song types
Fig. 5. Sonograms of different types of songs, started singing of which will consist of similar elements under the form, but modified in repertoires at different males (record on Curonian spit): 1a - song type F7, 1b - song type G6; 2a - song type I 9, 2b - song type G1; 3 - song type D1.
Started singing of song types F (fig. 5.1a) and G 6 (fig. 5.1b) are similar in the base (initial) form of elements, but they are sung differently a little. Started singing (the row of whistle elements) in song types I (fig. 5.2a) and G 1 (fig. 5.2b) as are similar, but they also differ in a manner of performance at singing. The song type D (fig. 5.3) has original (distinguished) started singing, but in the general (common) structure of elements " v \ " is similar with started singing of song types (F, G, I) analyzed by us. Thus, even within the limits of a local population (Curonian spit) variability of syllables (elements) of one kind in different types of songs is observed at their different manner of performance (of singing). It is possible to result (to show) one more example of vocal variability of started singing in different song types (fig. 6).
Elements of started singing (the row of whistle elements) in these song types are arc-similar under the form, but can sound differently: song type С* (fig. 6.2), song type C# (fig. 6.5) have a little creaking (gnash) of a sound "vji-vji", that confirms other pronunciation. In song types Т (fig. 6.4) and S (fig. 6.3) the manner of performance of started singing is similar with the previous song types (C*, C#), but a sound is more thin - "fui-fui". At song type C (fig. 6.1) elements of started singing can sound most precisely, by high tone - "fuit-fuit-fuit". Thus, despite of similarity of elements in the form (on sonogram), and their similar sound at singing, exists their different pronunciation, a manner of performance, that is phonetic distinction. How these phonetic forms of started singing were fixed in song culture of a population - complex (difficult) question (mistakes at training of songs, folding or forming another song cultures in different local populations, and mixing song traditions at migration?).
Casual copying of song types with possible mistakes, should conduct (lead) to change of birds repertoire of populations during some time. The birds, which were born later, had less time for song training that could conduct (lead) to more erroneous copying (Slater et al., 1980). +
Fig. 6. Sonograms of types of songs, started singing of which will consist of similar elements under the form, but with different manners of their singing (performance): 1 - song type С1 (record in settlement Fishing, Curonian spit), 2 - song type C*, 3 - song type S (record in biostation "Fringilla", Curonian spit), 4 - song type T3 (settlement Fishing, Curonian spit), 5 - song type C* 11 (record in Moscow).
Syllabic variability of elements in trill of song types
Similarity of started singing (the row of whistle elements) in these song types (D, F, G) was already discussed, but if to pay attention on trill elements - it is possible to note their similar base form, but thus to agree - that they differently have been sung in separate song types. In general, the given song types - D (fig. 7.1), F (fig. 7.2), G (fig. 7.3) are similar among themselves, and sometimes their some variants it is difficult to differentiate as three song types, but, nevertheless, their phrases (parts) in many respects differ in a manner of performance of elements at singing. Probably, such phonetic distinctions of the same elements also give different types of songs during cultural evolution of populations (in space and in time).
The trill of these three forms of song type C (fig. 8) consists of two phrases. The first phrase - finer (thin) under the form ("angular") elements, on hearing as a row of thin sounds ("til-til-til"), the second phrase - the complex (difficult) elements consisting of two subelements, and on hearing as more powerful sound ("tel-tel"). On sonograms the given elements both of the first, and of the second phrase have variability, though the initial (base) form of them is uniform, but in different song types at singing have been sung by different styles. The different phonetics (pronunciation) of a trill can generate (give) dialects of songs of one type in different territories, forming vocal culture of a local population.
Fig. 7. The types of songs having similar trill elements, but with different ways of their singing execution (performance) at singing (are recorded on Curonian spit): 1 - song type D3; 2 - song type F7; 3 - song type G7.
Fig. 8. Vocal variability of trill syllables of song type C: 1 - song type С1 (record in a settlement Fishing, Curonian spit), 2 - song type С*11 (record in settlement Sea, Curonian spit), 3 - song type C* 11 (record in Moscow).
Elements of a trill of song type M (fig. 9) are similar in the base, but differ on a manner of performance at singing: in song types M 1 (fig. 9.1) and M 8 (fig. 9.2) the accent at singing goes on the bottom bend of elements, and in song type M 12 (fig. 9.3) - the accent on the top bend. Such phonetic variability of syllables of a trill of song type M is observed at individuals of chaffinch males (Fringilla coelebs L.) within the limits of one local population (on Curonian spit).
Thus, phonetic variability of syllables (elements) of song types represents their basic (initial) structure, which can be uniform, similar, but modified at singing different chaffinch males (Fringilla coelebs L.) both within the limits of one local population, and in populations in different territories. Fig. 9. Vocal variability of trill syllables of song type M (record on Curonian spit): 1 - song type М1, 2 - song type М8, 3 - song type М12.
Syllabic variability in final stroke of chaffinch song types (Fringilla coelebs L.)
The extreme phonetic variant of song type M (fig. 10.1) is similar to song type J (fig. 10.2). Final strokes are similar on last elements.
We have attributed (related) these songs to one type (fig. 11). But despite of similar general (common) structure, on sonograms precise syllabic variability comes to light, is especial in final stroke: at song type C * (fig. 11.2) last element (which by the "triangular" form at song type C (fig. 11.1)) is divided in three separate subelements. This interesting phonetic difference which changes a kind of a phrase as a whole so, changing also the lexical party (side) of the given song type.
Fig. 10. Vocal variability of final strokes of song types М8 and J6 (record on Curonian spit): 1 - song type М8, 2 - song type J6.
Fig. 11. Vocal variability of final strokes of song type C (record in Moscow): 1 - song type С7, 2 - song type C* 11.
On sonograms a final stroke (fig. 12) has a number of phonetic features: at song type V1 (fig. 12.1) before last element (7) - three small elements of higher frequency (4, 5, 6); the second small elements (2) in final strokes also differ under the form. These phonetic features influences lexicon of song type as a whole, because there is a change of a kind of a phrase. The given samples of song type V have been recorded in repertoires of more than 1-2 individuals of chaffinch (Fringilla coelebs L.) in their local populations.
The case is interesting, when in one population (a settlement Wood or Lesnoy, Curonian spit) songs of one type, but differently performance at singing, have been found at singing of two males (are recoded in the certain day with identical conditions). One of them sang this song type normally (as the majority in our sample) (fig. 13.1), and another (fig. 13.2) - with appreciable errors in phonetics of elements (in their pronunciation) that is reflected in the form on sonogram. In repertoire of everyone male their song form are repeated many times at record (that is already was characteristic for them).
Fig. 12. Vocal variability of final strokes of song type V, recorded in different areas of Russia: 1 - song type V1 (record on Curonian spit, the Kaliningrad region), 2 - song type V (record in Zvenigorod, the Moscow region).
Fig. 13. Normal (1) and (2) forms of song type S taught (learnt) with phonetic mistakes (record in settlement Wood, Curonian spit).
Whether incorrectly learnt form of song type S (fig. 13) will be kept by the subsequent new generation by song training or remain as individual "mutation" which at all will disappear? Whether the erroneous song form becomes sometime of the one of characteristics of song cultures, traditions for one of local populations, probably, already dialect (changed) for that form of this type, which once was norm? It is difficult to answer, it is possible to assume only. Conclusion Thus, as a result of lexical (concerning phrases or parts of songs) and phonetic (concerning to syllables, elements of song types) vocal variability in local populations of chaffinch (Fringilla coelebs L.) the certain song cultures are formed, which are capable to change during time and can make dialect forms on all area of distribution of a species. For many birds species the patterns of variability represent change of vocal traditions are result of cultural evolution (Mundinger, 1980). Frequently, extreme (changed) phonetic variants turn into lexical variants (phrases vary because of change of forms of elements). In these cases it is difficult to differentiate new lexical variants from extreme phonetic variants of known syllabic types.
Equally with a substantiation of distinctions and variability of song types of birds as product (result) of mistakes of copying, of improvisation at singing and of transfers of features of song traditions to the following generation during cultural evolution (Slater et al., 1979, 1984; Ince et al., 1980), there is an assumption of gradual phylogenetic complication of initial "ancient" (more simple on the structure) song types to more "perfect", complex (difficult) in structure song forms, existing in a population equally with first, on the basis of occurrence of their geographical variability (biomorphism) (Simkin, 1983).
Nevertheless, there are a number of generalizations (Mundinger, 1982; Kroodsma, Miller, 1982):
1. differentiation (division or sharing, distinctions on the basis of the general (common)) of syllables of song types is widely distributed in the nature (Whitney, Miller, 1987; Petronovich, Baptista, 1984; Thielcke, 1984+); 2. microgeographical researches are perspective in phonetic variability, macrogeographical variability includes regional distinctions and similarities in syllabic sets (in groups of types of elements) (that is lexical variability - change of separate song parts and of species specific patterns of song as a whole);
3. among sparrow birds spatial distribution of vocal regional variability can give some forms, the majority of them are general (common), in which various regions were characterized by qualitative distinctions of syllables of songs (different types of song patterns of a species) (Wiley, 1971; Kroodsma, 1974; Nottebohm, 1969; Bertram, 1970; Lemon, 1965, 1966; Avery, Oring, 1977; Grimes, 1974);
4. microgeographical studying of vocal variability focus attention to processes of cultural evolution, macrogeographical studying gives the result in biological evolution;
5. very specific variations (changes) in syntax (sequence, combination of syllables) have been found at some species of birds (Thielcke, Linsenmair, 1963; Kroodsma, 1980; Mundinger, 1975; Bitterbaum, Baptista, 1979);
6. "islands" distribution correlates (have connection) with increase in variability of song patterns; there were the birds - in regular intervals distributed, and during nesting had constantly high density, distinction of their songs - small (insignificant) (Thielcke, 1969).
Mechanisms of forming and development of song traditions of many species of birds remain interesting for clearing (precise) evolutionary directions of species specific song during time and space. Thus it is necessary to note, that at some species the vocalization are not related by social training (song learning), and it is completely hereditary, genetically predetermined. Nevertheless, and these hereditary vocalizations are capable to develop during evolution, can change in the structure. Therefore, probably, influence of heredity and of environment cannot be strengthened in their separate values (accents) - would be more correct to consider forming song of birds at their dynamical interaction.
The literature
Avery M., Oring L. W. 1977. Song dialect in the Bodolink (Dolichonyx oryzivorus) // Condor. № 79. P. 113-118.
Bertram B. 1970. The vocal behavior of the Indian Hill Mynah (Gracula religiosa) // Anim. Behav. Monogr. № 3. P. 81-192.
Bitterbaum E., Baptista L. F. 1979. Geographical variation in songs of California House Finches (Carpodacus mexicanus) // Auk. № 96. P. 462-474.
Burt J. M., Beecher M. D. 2000. Social influences during song development in the song sparrow: a laboratory experiment simulating field conditions // Animal Behaviour. № 59. P. 1187-1197.
Grimes L.G. 1974. Dialects and geographical variation in the song of the splendid sunbird (Nectarinia coccinigaster) // Ibis. № 116. P. 314-329.
Ince S. A., Slater P. J. B., Weismann C. 1980. Changes with time in the song of a populations Chaffinches // Condor. № 82. P. 285-290.
Kreutzer M. 1974. Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes) // Rev. Comp. Anim. № 8. P. 270-286.
Kroodsma D. E. 1974. Song learning, dialects, and dispersal in the Bewick`s Wren // Tierpsychol. № 35. P. 352-380.
Kroodsma D. E. 1980. Winter Wren singing behaviors: A pinnacle of song complexity // Condor. № 82. P. 180-188.
Kroodsma D. E., Miller E. H., Quellet H. 1982. Communication and behavior an interdisciplinary series. Vol. 2: Song leaning and its consequence / D.E. Kroodsma, E.H. Miller. London: Academic press, 347 p.
Kurath H. 1972. Studies in Areal Linguistics. Bloomington: Indiana Univ. Press, 127 p.
Lemon E. L. 1965. The song repertoire of Cardinals (Richmondena cardinals) at London, Ontario // Can. J. Zool. № 43. P. 559-569.
Lemon E. L. 1966. Geographic variation in the song of Cardinals // Can. J. Zool. № 44. P. 413-421.
Lemon L. E. 1975. How birds develop song dialects // Condor. № 77. P. 385-406. +
Mundinger P. C. 1975. Song dialects and colonization in the House Finch, Carpodacus mexicanus, on the east coast // Condor. № 77. P. 407-422.
Mundinger P. C. 1979. Call learning in the Carduelinae: Ethological and systematic considerations // Syst. Zool. № 28. P. 270-283.
Mundinger P. C. 1980. Animal cultures and a general theory of cultural evolution // Ethol. Sociobiol. № 1. P. 183-223.
Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds // Acoustic communication in birds. New York, P. 147-208.
Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg. P. 265-280.
Nottebohm F. 1969. The song of the Chingolo, Zonotrichia capensis, in Argentina: Description and evaluation of a system of dialects // Condor. № 71. P. 299-315. Petrinovich L., Baptista L. F. 1984. Song dialects, mate selection, and breeding success in white-crowned sparrows // Anim. Behav. № 32. P. 1078-1088. +
Simkin G. N. 1983. Typological organization and population phylogeny of song at birds // Moscow group of nature science (Department of Biology, Vol. 88, № 1). P. 15-27. +
Slater P. J. B., Ince S. A. 1979. Cultural evolution in chaffinch song // Behaviour. № 71. P. 146-166. Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. P. 207-218. Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects // Behaviour. № 88. P. 76-97. +
Thielcke G., Linsenmair K. E. 1963. Zur GeographiachenVariation des Gesanges des Zilpzalpa, Phylloscopus collybita, in Mittel und Südwesteuropa mit einem Vergleich des Gesanges der Fitis, Phylloscopus trochilus // Journ. ornithol. № 104. P. 372-402.
Thielcke G. 1965. Gesangsgeographische Variation des Gartenbaumläufers (Certhia brachydactyla) im Hinblick auf das Artbildungsproblem // Z. Tierpsychol. № 22. P. 542-566.
Thielcke G. 1984. Gesangslernrn beim Gartenbaumläufer Certhia brachydactyla // Vogelwarte. № 32. P. 282-297.
Wiley R. H. 1971. Song groups in a singing assembly of Little Hermits // Condor. № 73 P. 28-35.
Whitney C. L., Miller J. 1987. Song leaning in the wood thrush // Can. J. Zool. № 65. P. 1038-1042. ++
The analysis of phrase combinations and various ways of syllabic singing at vocal variability of song types of chaffinch (Fringilla coelebs L.)
O. A. Astakhova, I. R. Byome
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Summary
Questions of evolutionary development of vocalizations (songs, calls) in time and space of many species of birds already for a long time are mentioned in bioacoustics. Connection of geographical variability and structural features of vocalizations was one of the first preconditions of evolutionary hypotheses. Development song cultures at species of birds with the big repertoires, having social training traditions, remains till now in many respects not clear. In article some features of formation of new song types of chaffinch (Fringilla coelebs L.) and their territorial distribution are considered.
Key words: birds song, species specific song, individual variation of song, parts (phrases) of song, phonetic distinctions, lexical distinctions, the quantitative analysis, song traditions in population, polymorphism of song, development (evolution) of song.
The quantitative analysis of chaffinch song types (Fringilla coelebs L.) in populations of the European part of Russia
O. A. Astakhova, I. R. Byome
Department of vertebrate zoology,
Faculty of biology, Moscow State University of M. V. Lomonosov
e-mail: beme@nm.ru
chaffinch@bk.ru
The quantitative analysis of songs of birds is one of the basic ways of an estimation of structure song patterns, their set in repertoires of individuals and a typological comparison in song culture of populations. With a view of revealing individual and geographical features of song types on different time-and-frequency parameters from experiments in local populations of different species of birds many bioacoustics had been lead (had been carried out) calculations of their statistical data and the subsequent comparisons of values (Barber, 1959; Zablotskaya, 1981, 1982; Marler and Tamura, 1962; Konishi, 1964; Kroodsma, 1985; Bremond, 1968).
First of all, in researches the characteristic of bird repertoires (number of song types, their general (common) phrases or parts, qualitative structure was mentioned at the certain volume of sample, frequency of occurrence and a comparison of song patterns in repertoire of separate individuals and in a population, as a whole) (Brown, 1985; Krebs, 1976; Kroodsma, 1976; Whitney, 1987; Dobson and Lemon, 1975; Payne et al., 1981; Falls and Krebs, 1975; Lemon and Chatfield, 1971). Chaffinch (Fringilla coelebs L.) - one of the most widespread species of birds which were objects of studying song organizations (Hinde, 1958; Slater, 1980, 1983). As against many others, chaffinch has harmonious, frequently short, precisely organized and in the greater degree song genetically determined in a time-and-frequency range (Thorpe, 1958; Marler, 1952; Nottebohm, 1969, 1967). The size of repertoire of one individual is within the limits of 1 - 6 types of songs (Slater, 1981; Marler, 1956).
The problem of our work will consist in definition of the quantitative characteristic of repertoire and the typological organization of chaffinch song (Fringilla coelebs L.), revealing of individual and geographical features of song patterns.
Material and methods
In different areas of the European part of Russia in spring-summer period of 2005-2006 years. Records of singing males have been made. For record of songs used tape recorder Panasonic RQ-SX95F, condenser microphone Philips SBC ME570. Further, sonogramms of songs were analyzed with the help of computer program Avisoft-SaSLab Light. In total it has been analyzed about five thousand songs. Types of songs were marked by Latin letters. At the quantitative analysis measured both structure of song type as a whole, and its separate phrases (in three parts of songs - started singing, a trill, a final stroke) on different frequency-time parameters: length (duration), maximal, minimal and average (median) frequency, quantity (amount) of elements (syllables), length (duration) of syllables, intervals between syllables, intervals between phrases.
During statistical processing the primary data basically defined arithmetic-mean value and a standard (arithmetic-mean) deviation (with the help "Master of functions" of Microsoft Excel). At comparison frequency-time parameters are allocated (distinguished) a range, within the limits of which the difference of values was considered insignificant (if values difference equal or smaller 0,5 КHz in frequency and 0,02 sec in length). The characteristic of song types and repertoires of males with the greater sample of songs (n> 14) was more significant.
Results and discussion
Song repertoire includes song types, which is capable to reproduce male of chaffinch (Fringilla coelebs L.) at singing. Song type - individual (within the limits of one population) or geographical (in populations on different territories) song variant of a species. It is usually these song patterns in its individual becoming are precisely determined in repertoire of an adult individual after a stage of crystallization (stabilization) of species specific song. The repertoire of young individuals frequently is established not completely, because after sensitive period of training (learning) song types are still plastic in the structure, can change or not appear at all under influence of "communicative" group (Thorpe, 1958; Marler, 1956; Nottebohm, 1967).
On the basis of experiments in different local populations the statistical analysis of the size of chaffinch repertoire (Fringilla coelebs L.) has been lead (has been carried out). Those individuals from which n>14 songs have been recorded, were taken into account only. In northwest of the European Russia from the general (common) sample are allocated males, recorded in different areas of Curonian spit of Baltic sea (settlement Sea or Morskoy, settlement Fishing or Ribachiy, area of biological research station "Fringilla", settlement Wood or Lesnoy) and having on the average n > 20 reproduced (output) songs. In the central part of the European Russia at processing records of chaffinch (Fringilla coelebs L.) also were taken into account males with volume of sample of songs on the average n > 20. In the table repertoires of individuals from different areas (Zvenigorod, Moscow, Michurinsk) are shown (tab. 1).
On the basis of the primary data arithmetic-mean value and a standard deviation of number of males song types have been calculated. In result appeared, that the average size of chaffinch repertoire (Fringilla coelebs L.) in different regions of the European Russia included approximately 2 types of songs (tab. 1).
Table 1. The average size of chaffinch repertoire (Fringilla coelebs L.) in different areas of the European Russia
Northwest of European Russia (Curonian spit) _
x ± õ
Central part of European Russia _
x ± õ
Area of research biostation "Fringilla"2,5 ± 1,67Zvenigorod
(Moscow region)1,68 ± 0,67Settlement Fishing (Ribachiy)2,33 ± 1,5Moscow2 ± 0,63Settlement Sea (Morskoy)1,83 ± 0,4Michurinsk
(Tambov region.)2,1 ± 0,66Settlement Wood (Lesnoy)1,8 ± 0,45 Total2,115 ± 0,35Total1,93 ± 0,22
The note: the volume of sample of songs from one male in the center of the European Russia has on the average made 24,7 ± 11,3; average volume of song sample from one male in northwest of the European Russia 26,8 ± 12,25.
The maximum quantity of song types, which are included in chaffinch repertoire - six, is found in samples of Curonian spit (area of biological research stations "Fringilla") (tab. 1). The minimal size of repertoire - 1 song type. Nevertheless, quite often there were repertoires with other quantity of song types (tab. 2).
Table 2. A percentage comparison of chaffinch repertoires (Fringilla coelebs L.) in populations of the European Russia
Repertoire size
(quantity of song types)Quantity of males with the given size of repertoirePercent from the common number of individuals in a population11822 %24656,1 %389,76 %4 - 644,9 %
The note: in sample of populations of the European Russia (N = 137) those were taken into account chaffinch males, from which have been recorded on the average n>20 songs (them appeared n = 76).
Very seldom in populations of chaffinch (Fringilla coelebs L.) there were the repertoires consisting of 4-6 song types (4 male - 4,9 %). The repertoire from 3 song types - also was rare (8 male - 9,76 %). The often phenomenon - 2 song types in repertoire (46 male - 56,1 %). Even the repertoire from one song type met less often (18 male - 22 %) (probably because the majority recorded males with such repertoire have not come in the account because of small song sample, that is n <14).
It is necessary to note, that it is the generalized statistical calculations. In the nature there is all is very relative. Sometimes it is possible to record one chaffinch male during hours, and in its repertoire only 1-2 song types are found, though the bird can reproduce and a lot song patterns (it is allowable, 3-4). But it is possible to conduct record and 10 - 15 minutes - and to grasp thus whole repertoire of an individual (4, 5, 6 types of songs? And it is possible, only and, really, - 1-2 types). The rare song type in repertoire can appear once (time at one o'clock, at two o'clock, in day, week?). Whether it will get in sample of the researcher? (Kroodsma, 1974; Slater, 1983).
Interestingly typological comparison of chaffinch songs (Fringilla coelebs L.) in local populations. In samples on Curonian spit of Baltic Sea (N=153) we allocate (distinguish) 22 song types which were marked by Latin letters (A B C D E F G H J I K L M N O P Q R S T U V) (fig. 1). At record songs of one type came across in different points of territory (was considered, that belong to repertoires of different males), therefore equally with the letter were designated by numbers in ascending order (for example, А1, А2, А3, etc.).
In populations of the central part of the European Russia (N=65) 15 different song types are found, from which 12 song types were similar in the structure to corresponding types on Curonian spit, but frequently had the modified elements on sonogramms, therefore were considered as dialect forms (B C D F G I J M N S U V) (fig. 3). The song type A was stable in the structure in different regions of Russia. It has not been found analogues six song types (H K Q P R T), and types W, Y appeared new in relation to known to us song patterns on Curonian spit. Fig. 1. The most widespread types of songs of chaffinch (Fringilla coelebs L.) in a population of northwest of the European Russia (Curonian spit): 1 - song type A, 2 - song type S, 3 - song type М12, 4 - song type F, 5 - song type C, 6 - song type C *, 7 - song type I, 8 - song type O, 9 - song type R.
Table 3. The quantitative characteristic of the most widespread song types of chaffinch (Fringilla coelebs L.) (the example of population in Curonian spit)
Parameters
of songsSong type (token) _
x ± õАSMFCIORDuration of song, sec1,96
± 0,252,29
±0,251,875
±0,162,24
±0,31,96
±0,111,86
±0,221,56
±0,152,05
±0,0861,97
±0,23Min frequency, KHz1,6
±0,161,605
±0,051,718
±0,121,57
±0,0381,59
±0,121,56
±0,0651,64
±0,171,72
±0,131,63
±0,063Max frequency, KHz7,63
±0,197,51
±0,157,6
±1,257,18
±0,38,36
±0,97,8
±0,477,15
±0,287,44
±0,77,683
±0,42Median (average) frequency, KHz4,22
±0,23,84
±0,064,06
±0,384,01
±0,114,2
±0,124,1
±0,224,3
±0,094,05
±0,214,1
±0,14Number of syllables14,9
±1,8619,45
±1,4817,7
±1,715,6
±1,716,03
±1,411,9
±1,3512,05
±1,6117,5
±0,9515,6
±2,7Duration of syllables, sес:0,04
±0,0070,12
±00,046
±0,0170,11
±0,020,14
±0,030,14
±0,0110,06
±0,0040,039
±0,0070,087
±0,045started singingtrill0,11
±0,0140,06
±0,0040,08
±0,0060,096
±0,040,066
±0,010,083
±0,0040,072
±0,0040,16
±0,020,09
±0,0321 phrase2 phrase0,1
±0,0010,11
±0,0170,06
±0,0050,09
±0,03final stroke0,12
±0,0360,12
±0,0070,041
±0,0160,13
±0,020,072
±0,0070,15
±0,0430,15
±0,0350,19
±0,030,12
±0,05Intervals between syllables, sec:0,047
±0,0090,083
±0,00030,048
±0,0160,067
±0,0050,061
±0,020,063
±0,0080,03
±0,0050,057
±0,017start singingtrill0,054
±0,0170,026
±0,0020,0405
±0,010,052
±0,010,026
±0,0090,0485
±0,0050,044
±0,0070,03
±0,0060,04
±0,0111 phrase2 phrase0,035
±0,10,042
±0,0150,03
±0,0070,03
±0,021final stroke0,05
±0,0140,033
±0,0070,055
±0,0090,059
±0,0140,035
±0,0050,069
±0,00380,08
±0,0110,05
±0,0410,054
±0,016Intervals between phrases, sес:0,05
±0,0120,054
±00,047
±0,0020,057
±0,030,05
±0,0060,059
±0,020,052
±0,0090,033
±0,0040,05
±0,008start singing
and trilltrill
and final stroke0,074
±0,0150,066
±0,0160,04
±0,0090,06
±0,0070,047
±0,0220,056
±0,0120,048
±0,0120,056
±0,0160,056
±0,011Intervals between songs, sес:6,54
±2,87,21
±3,116,6
±0,375,2
±3,567,13
±2,57,58
±2,187,45
±4,177,66
±0,0166,92
±0,81Number of measured song (n)152411345733683734,88
±19,56
The note: average value and a standard deviation of parameters of song types are specified; the songs of one type, which has been recorded in different points of territory, but not having any differences in the structure, were taken into account; in the last column - the generalized values on all measured song types (are signed by a font).
Besides, in samples of the center of the European Russia there were the types of songs combined from 2-3 phrases (parts of songs), which we already found in populations of northwest of the European Russia, but their elements have been modified (dialects). We spelled (called) such types of songs as those types, from phrases of which they have consisted (for example, NIE, SNE, ВF, TS, СM, etc.) Nevertheless, these song forms it is possible to allocate (distinguish) as independently existing types of song describing song culture of a local population, and at record they met in different points of territory (were not individual in a population) (fig. 4).
To have more precise representation about song types, which we discuss, it is possible to result (show) sonograms of their samples (fig. 1) and some statistical data of their basic time-and-frequency parameters (tab. 3).
At close (attentive) viewing of sonogramms it is possible to allocate (distinguish) a number of features in structure of songs, characteristic only for the certain type. It is necessary to remind, that we considered songs are similar on two or all three song parts relate to songs of one type (started singing or the row of whistle elements, a trill and a final stroke), which also can include one or two phrases (the elements similar under the form). Fig. 2. Degree of a variation of values of parameters of different song types of chaffinch (Fringilla coelebs L.) on Curonian spit of Baltic Sea (north of Europe): 1 - length (duration) of song, 2 - the minimal frequency, 3 - the maximal frequency, 4 - average (median) frequency, 5 - number of syllables, 6 - length (duration) of syllables in started singing, 7 - length of syllables of a trill (1 phrase), 8 - length (duration) of syllables of a trill (2 phrase), 9 - length of syllables of a stroke, 10 - intervals between syllables (started singing), 11 - intervals between syllables of a trill (1 phrase), 12 - intervals between syllables of a trill (2 phrase), 13 - intervals between syllables of a stroke, 14 - intervals between phrases (started singing and a trill), 15 - intervals between phrases (a trill and a stroke), 16 - intervals between songs.
From table 3 it is visible, that values of parameters of song types are various, but nevertheless are in the certain range that can confirm the precise organization and in many respects a genetic basis of species specific song of chaffinch (Fringilla coelebs L.). Despite of the big similarity of values of parameters of song types, nevertheless it is interesting to find out, on what from them variability was the greatest. For this purpose it is necessary to calculate factor of a variation (V, %) each parameter. The factor of a variation represents a parity (ratio) of a standard deviation and average value, shows a degree of a variation of an attribute. It is possible to make the diagram on character of variability of parameters of different types of songs (fig. 2, 5).
The most varying attributes (33 - 50 %) different song types appeared (on Curonian spit, north of Europe) (fig. 2): length (duration) of syllables in started singing (6), trill (7, 8) and a final stroke (9) (and, started singing and a final stroke were more variability). Significant stability (small variability) (3 - 5 %) has come to light at minimal (2), average (median) (4) and the maximal (3) frequencies. Other parameters of different song types had an average variation (10 - 30 %), which can enter into a category of homogeneous values, but among these data there are also those, which variability was the greatest - intervals between syllables in started singing (the row of whistle elements) (10), intervals between syllables of a trill (11) and intervals between elements of a stroke (13).
Thus, songs of chaffinch (Fringilla coelebs L.) of different types can be stable in frequency, but are more individual in values of length (duration) of syllables (elements) and intervals between them.
The factor of a variation (V, %) (fig. 2, 5) shows a degree of a variation (variability) of this or that attribute (sign), proceeding from his(its) average arithmetic value (x) and a standard deviation (õ) (tab. 3), which shows as far as on the average each value of an attribute (sign) of the general (common) sample "deviates" from average value of this attribute. The more value of factor of a variation (V, %) is the more variability of values of parameter. Such attitude (relation) can be expressed by the general (common) formula (Nimenya, 2003):
õ
V = --- × 100%
x
On the basis of the given schedules (fig. 2, 5) we can make the conclusions on a variation of time-and-frequency parameters of different song types of chaffinch (Fringilla coelebs L.) from two geographical populations of the East Europe:
а) Similarities on a variation of values of quantitative parameters of chaffinch songs (Fringilla coelebs L.) in two geographical populations: • poorly (weak) varied (factor of variability (V) 3-9 %) in all two geographical populations of chaffinch are minimal, maximal and average (median) frequency (KHz) - it is probable, the frequency range of species specific song of chaffinch is genetically determined; • the average variation (factor of variability (V) 10-30 %) in all two geographical populations of chaffinch was observed at such parameters as length of songs (sec), the number of syllables in a song, intervals between songs (sec), intervals between syllables of a trill (2 phrase) (sec), intervals between phrases (started singing and a trill) (sec), intervals between a trill and a final stroke (sec) - is probable, it also genetically determined, more or less stable attributes (signs) of species specific song of chaffinch, but capable in the greater degree to change, vary, than frequency of songs (КHz) at individuals of chaffinch at any geographical population;
• the strong variation (factor of variability (V) 33-50 %) in all two geographical populations of chaffinch was at parameters of song types: length of syllables in started singing (sec), length of syllables of a trill (1 phrase) (sec), length of syllables of a final stroke (sec) - these parameters of songs are capable to change strongly in a population of chaffinch at song training (learning) and, probably, at genetic variability as well to create different song forms (types of species specific song, distinguished on structure and the form of elements).
b) Distinctions on a variation of values of quantitative parameters of chaffinch songs (Fringilla coelebs L.) in two geographical populations:
• the length of syllables of a trill (2 phrase) (sec) is average varied (factor of variability (V) 10-30 %) in populations of the center European Russia, but in northern population is strongly varied (factor of variability (V) 33-50 %) - probably, chaffinch was a nesting species on Curonian spit (north), had the big population density and many the differentiated types of species specific song (22 song types in sample), therefore elements of a trill could be various under the form and quantitative parameters; while in populations of the center European Russia can be many migrants, flying groups of chaffinch, which adapt in many respects to local song cultures of nesting chaffinch and have poorly differentiated under the form and time-and-frequency parameters elements (syllables) of a trill (especially 2-nd phrases at songs with a complex (difficult) trill on structure);
• intervals between syllables in started singing (sec), intervals between syllables of a trill (1 phrase) (sec), intervals between syllables of a final stroke (sec) is average vary (factor of variability (V) 10-30 %) in northern population, but is strongly vary (factor of variability (V) 33-50 %) in the center European Russia it is possible, nesting chaffinch better kept of song tradition in the north (Curonian spit) and the given attributes (signs) of species specific song were more stable at variability, than among the majority of flying chaffinch (migrants) of the center European Russia.
Fig. 5. A degree of a variation of values of parameters of different song types of chaffinch (Fringilla coelebs L.) in the center of the European Russia: 1 - length of songs (sec), 2 - minimal frequency (КHz), 3 - maximal frequency (КHz), 4 - average (median) frequency (КHz), 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 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).
For full representation of song cultures in different local populations of chaffinch (Fringilla coelebs L.) it is necessary to show samples of the most widespread song types in territory of the center of the European Russia, which can not coincide with a degree of their spreading in other populations (fig. 3).
It is impossible to miss from attention and the so-called combined types of songs. In the first researches on Curonian spit (Kaliningrad region) these song forms were individual (unique), and in the subsequent experiments in the center of the European Russia - song types combined of modified elements (in relation to already known) appeared in the majority and in many respects characterized song culture of a local population (fig. 4, tab. 5). During the quantitative analysis of song types comparison of their basic time-and-frequency parameters has been lead (has been carried out) in view of individual (within the limits of one population) and geographical (in populations in different territories) variability (tab. 4).
In table 4 distinctions of parameters were considered within the limits of each song type (the font allocated (distinguished) the greatest by quantity of distinction at songs of one type). At comparison of values of different song types by the common there were distinctions for such parameters as length (duration) songs (sec), number of syllables and intervals between songs (sec). More individual (not for all song types) distinctions in the maximal frequency (types of songs G, C, Е, В), length (duration) of syllables in started singing (song type С), trills (types of songs G, C, B), a final stroke (song types A, G, E, I), intervals between syllables and phrases (song types A, Е, I, C) were characteristic. Values of parameters, which entered into limits of an insignificant difference (had small distinction) - minimal and average (median) frequency (KHz), intervals between syllables (elements) of phrases (sec) were characteristic for songs of one type, but the marked similarities came to light at different types (having numbers song variants), measured in parallel, independently from each other. If to pay attention to distinctions of values at individual and geographical variability (tab. 8) it is possible to note their different parity (ratio) even within the limits of one song type. So, sometimes individual variability of parameters of songs of one type (which, apparently, should be the least) had a lot of distinctions, than values of the same parameters of songs at geographical variability which could and not have at all distinctions (measurement of song types from the different local populations removed from each other frequently on the big distances) - for example, at types of songs A and G - length (duration) of a stroke (sec), the maximal frequency (KHz) - at types of songs Е, I, G, intervals between a trill and a stroke (sec) - at song type C. Nevertheless, song types in most cases distinctions or coincided at individual and geographical variability, or prevailed at geographical variability.
Thus, the majority of distinctions has been found in values of length (duration) of songs (sec), numbers of syllables and intervals between songs (sec) of one type (it has come to light for all song types, which have been analyzed), the least distinctions - minimal, average (median) frequency (KHz), intervals between syllables and phrases (sec). Other parameters in the distinctions were individual for each type (tab. 4). Geographical and individual variability had no precise parity (ratio) in distinctions (songs of one type in one territory could differ more, than songs of the same type in different local populations), but presence of distinctions on the certain parameters frequently coincided.
Conclusions
The quantitative analysis of songs - one of complex (difficult), relative, but accessible and precise ways of the description individual and population structures of song organization of different bird species.
Fig. 3. The most widespread types of songs of chaffinch (Fringilla coelebs L.) in populations of the central part of the European Russia: 1 - song type C, 2 - song type C*, 3 - song type I, 4 - song type J, 5 - song type A, 6 - song type B.
As a result of statistical calculations it has been found, that the average size of chaffinch repertoire (Fringilla coelebs L.) in different areas of research made approximately 2 song types. In different populations, song culture of chaffinch can be characterized by different song types and their different percentage parity (ratio) - for example, those song types, which were in the majority in one population, could make small percent in another population - and, some song types were as dialect forms (modified in the form of elements on sonogramms). The "combined" types of songs (consisting of phrases of different song types) could be as individual in one population (were considered as "a cultural mutation"), and more widespread in other local populations, describing the song culture in the greater degree.
Chaffinch songs (Fringilla coelebs L.) of different types could be stable in frequency (KHz), but more individual (differ) in values of length (duration) of syllables (elements) (sec) and intervals between them (sec). Fig. 4. Sonogramms of "combined" types of chaffinch songs (Fringilla coelebs L.), recorded in different regions of the European Russia: 1 - song type AC, 2 - song type E I, 3 - song type EM, 4 - song type ОС6 (are recorded on Curonian spit of Baltic sea, north of Europe); 5 - song type BF, 6 - song type N I E, 7 - song type SNE (are recorded in the center of the European Russia). +
Table 5. A percentage parity (ratio) of the combined song types of chaffinch
(Fringilla coelebs L.) in populations of the European Russia
Place of record
(region of Russia)Song type
(token)Number of song types in samplePercent from the common number of song types in a populationVolume of sample
type songs
(n)Northwest of
European
Russia
(Curonian
spit)Setllement Wood
(Lesnoy)E I10,55 %183
E M10,55 %Area of
biostation
"Fringilla"A C10,55 %O C610,55 %O I 510,55 %O C*10,55 %Total63,28 %Central part of European RussiaZvenigorod
Moskow, MichurinskN I E33,2 %93
ZvenigorodS N E44,3 %Moskow, MichurinskD E22,15 %MichurinskB F66,45 %O E11,075 %T S22,15 %C T22,15 %C M22,15 %G C22,15 %Total2425,8 %
The note: a percentage parity (ratio) expected proceeding from the common number of song types in samples of populations in northwest of the European Russia - 183 song types, in the central part of Russia - 93 song types (that is repertoires of male with small samples were taken into account also); thus it is necessary to understand, that songs of one type from everyone male had sample on the average n>20.
Table 4. Results of comparison of parameters of different song types of chaffinch (Fringilla coelebs L.) in populations of the European Russia at individual and geographical variability Parameters of songs
(quantity of distinctions)Song type (token)АGCEIB Ind.1
var. Geog.2
var.Ind. 1
var.Geog.2
var.Ind.1
var.Geog.2
var.Ind. 1
var.Geog.2
var.Ind.1
var.Geog.2
var.Ind.1
var.Geog.2
var.Duration of song, secnumber2178814194320736% *20,217,513,315,418,91614,31515,8712,7318,7514,3Min frequency, KHznumber3%2,5Max frequency, KHznumber125498411126%0,958,37,712,26,7214,358,73,614,3Median (average) frequency, KHznumber113242% *0,92,551,73,173,6Number of syllablesnumber1958813183218736%18,2712,513,315,417,615,110,71014,312,7318,7514,3Duration of syllables,
sес:started singingnumber122513218412 %0,93,33,86,810,97,1456,357,276,254,8trill1 phrasenumber544226321%4,8106,73,852,75,0410,7101,82 phrasenumber768125%11,711,510,810,0811,9final strokenumber154521431553%14,4108,33,81,353,361511,99,097,14Intervals between syllables, sec:start singingnumber34732852% *2,97,75,8810,7106,359,094,8trill1 phrasenumber10541213%9,612,56,71,97,1455,452 phrasenumber4512%7,74,26,254,8final strokenumber12617223%0,955,0455,63,612,57,14Intervals between phrases, sес:start singing
and trillnumber633211161%5,87,557,1458,710,92,4trill
and final strokenumber62328521462%5,8553,810,84,27,1453,1710,912,5Intervals between songs, sесnumber958914113220513% *8,712,513,317,318,99,2410,71015,879,096,257,14Quantity of compared pairs of song (n)2178915195320737Quantity of parameters
distinctions104 4060 527411928 20126 5516 42
The note: % * - percent from the common number of distinctions on all parameters of the given song type at individual (geographical) variability; 1 Individual variability - variants (versions) of songs of one type, which has been recorded in different points of certain territory or in repertoire of one male; 2 Geographical variability - variants (versions) of songs of one type, which has been recorded in different territories; differences of parameters, a difference between which > 0,5 КHz in frequency and > 0,02 sec in length were taken into account; empty cells of the table - the data of this parameter were within the limits of an insignificant difference; values of parameters with a plenty of distinctions are allocated (signed) by a font +.
Songs of one type less differed on values minimal, average (median) frequencies (KHz), intervals between syllables and phrases (sec). The greatest quantity of distinctions has fallen (were) to such parameters of songs of one type as length (duration) of songs (sec), number of syllables, intervals between songs (sec). Geographical and individual variability could coincide on presence of distinctions in the same parameters of songs of one type, but had no conformity in a percentage parity (ratio).
Literature
Barber D. R. 1959. Singing pattern of the common chaffinch, Fringilla coelebs Linn. // Nature. № 10. P. 129.
Bremond J.-C. 1968. Recherche sur ia se´mantique et ies e´le´ments vecteurs d´information dans les signaux acoustiques du rouge-gorge Erithacus rubecula // Terre Vie. № 2. P. 109-220.
Brown D. 1985. The role of song and vocal imitation among common Crows (Corvus brachyrbyncbos) // Z. Tierpsychol. № 68. P. 115-136.
Dobson С. W., Lemon R. E. 1975. Re-examination of the monotony-threshold hypothesis in bird song // Nature. № 257. P. 126-128.
Falls J. B., Krebs J. R. 1975. Individual recognition by song in white-throated sparrows II: Effects of location // Can. J. Zool. № 53. P. 1412-1420.
Jellis R. 1977. Bird sounds and their meaning. Cambridge, 256 p.
Hinde R. A. 1958. Alternative motor patterns in Chaffinch song // Anim. Behav. № 6. P. 211-218.
Konishi М. 1964. Effects of deafening on song development in two species of juncos // Condor. № 66. P. 85-102.
Krebs J. R. 1976. Habituation and song repertoires in the great tit // Behav. Ecol. Sociobiol. № 1. P. 215-227.
Kroodsma D. E. 1976. Reproductive development in a female songbird: different stimulation by quality of male song // Science. № 192. P. 574-575.
Kroodsma D. E. 1985. Development and use of two song forms by the Eastern phoebe // Wilson Bull. № 97. P. 21-29.
Lemon R. E., Chatfield C. 1971. Organization of song in cardinals // Anim. Behav. № 19. P. 1-17.
Marler P. 1952. Variation in the song of the Chaffinch Fringilla coelebs // Ibis. № 94. P. 458-472.
Marler P. 1956b. The voice of the chaffinch and its function as a language // Ibis. № 98. P. 231-261.
Marler P., Tamura M. 1962. Song dialects in three populations of White-crowned Sparrows // Condor. № 64. P. 368-377.
Nottebohm F. 1969a. The "critical period" for song leaning in birds // Ibis. № 111. P. 386-387.
Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs. P. 265-280.
Payne R. B., Thompson W. L., Fiala K. L., Sweany L. L. 1981. Local song traditions in indigo buntings: cultural transmission of patterns across generations // Behaviour. № 77. P. 199-221.
Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. P. 207-218.
Slater P. J. B. 1981. Chaffinch song repertoires: observations, experiments and a discussion of their significance // Z. Tiërpsychol. № 72. P. 177-184.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. P. 535-570.
Zablotskaya M. M. 1981. Acoustic communication of Acanthis flammea flammea L. // Academy of sciences of the USSR. Puscheno, 27 p. (Centre of science of biological researches, Institute of biological physics, a record library of animal's voices).
Zablotskaya M. M. 1982. Acoustic communication of Acanthis cannabina L. // Academy of sciences of the USSR. Puscheno, 40 p. (Centre of science of biological researches, Institute of biological physics, a record library of animal's voices).
Whitney C. L., Miller J. 1987. Song leaning in the wood thrush // Can. J. Zool. № 65. P. 1038-1042.
The quantitative analysis of chaffinch song types (Fringilla coelebs L.) in populations of the European part of Russia
O. A. Astakhova , I. R. Byome
Department of vertebrate zoology,
faculty of biology, Moscow State University of M. V. Lomonosov
e-mail: beme@nm.ru
chaffinch@bk.ru
Summary
Time-and-frequency characteristics of sparrow songs (Passeriformes) can be tracked both in a sound equivalent, and as the image on sonograms. In given article generalizing aspects of the statistical analysis chaffinch song (Fringilla coelebs L.) from different local populations are submitted. The medium-sized estimation of male repertoire, a degree of a variation of different time-and-frequency parameters of chaffinch song types is resulted. Also drew conclusions of comparison of parameters of songs of one type at individual and geographical variability. The data it is possible to use at discussion of evolutionary development song of chaffinch.
Vocal variability of chaffinch song (Fringilla coelebs L.) as a condition of cultural evolution in local populations
O. A. Astakhova, I. R. Byome
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Singing for the majority of sparrow birds - are incontrovertible feature of their life, determining and building a reproductive cycle preparation and course (Lukanus, 1907; Darling, 1938, from Simkin, 1972). At detailed studying song repertoire of many sparrow species it was revealed, that the individual has not one, and some types of songs. Thus spring singing represents the multifunctional phenomenon (Simkin, 1972) and can bear (carry) various values (Banin, Byome, 1994). On I. Altum (Altum, 1868, from Simkin, 1972) the singing - is not only means of attraction female, but also means of intimidation of the contender and delimitation of nested territory. Variants or types of species specific songs are individually various and distributed between individuals of a population (Slater et al., 1980; McGregor and Krebs, 1982).
In early works song geographical variability of many species of birds was marked as a phenomenon of differentiation song structures or song patterns (as the image on sonograms) in different local populations (Marler, 1952; Marler and Tamura, 1962; Sick, 1939; Poulsen, 1951). Local variants of bird vocalizations can be considered as analogy of human speech and to name dialects. The word "dialect" has taken place from linguists and means a local speech variant, characteristic for region. Borders of a dialect define lexical (dictionary structure), morphological (an accent, structure) and phonetic (pronunciation) the characteristic of the dictionary (Kurath, 1972). Explanation of bird song variability shows, that traditional patterns of vocal behaviour do not exist equally with morphological features (more likely, as separate aspect). It partly due result of cultural evolution of birds which represents the important factor, as though a traditional heredity developed, but, apparently, it not a parameter in morphological variability (Lemon, 1975; Slater, Ince, 1979; Mundinger, 1980).
Basic (base, the basic) the structure of bird vocalization represents fundamental, stabilized, species specific characteristic feature of vocalization (Mundinger, 1979). Many local populations of bird are characterized by the population-specific image (on sonograms) a vocal pattern (dialect). Thielcke (1965, 1969) defined dialects of birds as vocal variants with mosaic distribution. This mosaic definition one of the basic which are widely used, but the majority of microgeographical variability of bird song researchers not precisely enough assert (approve) in studying vocal distribution, whether there is a mosaic pattern (Kreutzer, 1974; Kroodsma, 1974). Chaffinch (Fringilla coelebs L.) is classical object of studying of becoming of vocal repertoire (Thorpe, 1958; Marler, 1956; Nottebohm, 1967) and of geographical variability species specific songs in a population (Promptov, 1930; Sick, 1939; Thielcke, 1961; Simkin, 1983; Slater et al., 1984). As against many others, chaffinch (Fringilla coelebs L.) has harmonious, frequently short, precisely organized and in the greater degree song genetically determined in a time-and-frequency range (Thorpe, 1958; Marler, 1952; Nottebohm, 1969, 1967). The size of song repertoire of one individual is within the limits of 1 - 6 types of songs (Slater, 1981; Marler, 1956).
Regional variability of bird song can be determined in qualitative aspect (the form of a syllable, syntax) and quantitative (time-and-frequency) parameters. The concept of a dialect should be closely (cautiously) and is carefully argued in an establishment of regional variability signal and song patterns of birds. Methods
One of problems of our researches has consisted in revealing macrogeographical distinctions of chaffinch song (Fringilla coelebs L.) in different local populations which are removed approximately on 1000 km from each other. In northwest (Curonian spit, Kaliningrad region) and central parts of the European Russia (Zvenigorod, Moscow, Michurinsk) have been made tape recorder records of singing males during the spring-and-summer period of 2005-2006 (N=218 of males). For record of songs used tape recorder Panasonic RQ-SX95F, condenser microphone Philips SBC ME570. Sonograms of song type were analyzed with the help of computer program Avisoft SASLab Light. In total it is analyzed about five thousand songs. To one type considered the songs, having two or all three similar parts: started singing (the row of whistle elements), trill elements, a final stroke (they as can be subdivided into phrases - the elements similar in the form). Types of songs were marked by Latin letters. At the analysis of song sonograms basically applied two qualitative methods: revealing of phonetic distinctions (frequency of a sound, its form on sonogram) or ways of a pronunciation of syllables of the phrases making songs; revealing of lexical distinctions (changes of phrases of songs, as a whole) (Mundinger, 1982). As the quantitative analysis of dialect song forms of one type and comparison of their basic time-and-frequency parameters (duration of all songs, number of elements in song type, duration of syllables in a trill, the maximal, minimal and average (median) frequency of songs, intervals between songs) was carried out (was spent).
Results
Dialect song forms of one type
Methods of geographical dialects can be applied to songs of birds. For example, sonograms can be used for definition of vocal variants which are geographically distributed.
Song dialect of birds is a variant of the traditional song form, divided (shared) by the 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 the given population. But thus big song repertoires of some sparrow species interfere with objective definition of dialect borders (Thielcke, 1969; Kreutzer, 1974; Kroodsma, 1974; Baker, 1975; Baptista, 1975; Lemon, 1975; Payne, 1981; Mundinger, 1980, 1982).
In populations of the central part of the European Russia (N=65 of males) we allocate (distinguish) 15 types of chaffinch songs (Fringilla coelebs L.), which were completely similar in structure or are in part modified in the syllabic form (on sonograms) in comparison with songs of corresponding types in samples of a northwest part of the European Russia (N=153 of males). For seven types of songs from twenty two (from sample on Curonian spit) analogies has not come to light - probably, owing to their rarity. Many song types in the central part of the European Russia were considered as combined of phrases of syllabic patterns known to us, but frequently with the changed figure on sonogram.
In result, 12 dialect song forms of one type, which appeared similar in base structure of elements, phrases have been found, but in different regions of Russia frequently had distinguished manners, ways of their performance at singing (phonetic aspect). The average size of chaffinch repertoire (Fringilla coelebs L.) in populations of the central part of the European Russia from statistical calculations has been submitted 1,93 ± 0,22 types of songs (max - 4 types of songs, min - 1 song type) at a record on the average 24,7 ± 11,3 songs from one male.
As a bright example of chaffinch vocal variability (Fringilla coelebs L.) it is possible to show some local variants of songs which we have attributed to one type (fig. 1, 2). Samples of song types which will be resulted were not single in a population and more than 1-2 individuals have been recorded in repertoires. Fig. 1. Dialect forms of song types V: a - song type V (record on Curonian spit, the Kaliningrad region), b - song type V1 (record in Zvenigorod, the Moscow region).
In Zvenigorod (fig. 1b) started singing (the row of whistle elements) of songs such as type V has shorter second subelements which are in top concerning the first subelements. Elements of a trill are more complex (difficult) in Zvenigorod (fig. 1b). A final stroke - three large base elements are present as at samples of northwest, and the center of the European Russia. But in songs on Curonian spit (fig. 1a) before last element (7) of stroke are three small elements (4, 5, 6) higher frequency (KHz). Probably, it is phonetic components of a dialect, which characterize style of singing (song culture) in a local population on the given type of songs. Fig. 2. Local variants of song types C (are recorded in different areas of Russia): a - song type C (record on Curonian spit), b - song type C* (record in Moscow), c - song type С* (record on Curonian spit).
Song type C (fig. 2) - the most widespread in all areas of the European Russia investigated by us. Probably, therefore the given song type has some forms on a manner of performance of phrases at singing: simple song type C (fig. 2a) and song type С* (fig. 2c), recorded on Curonian spit; the song type C# (fig. 2b) is recorded in Moscow. Nevertheless, all these variants of songs on hearing are perceived is similar: started singing - a number (line) sharp (sometimes creaking) of whistle sounds ("fuit-fuit-fuit"), two-phrase trill - the first phrase is heard as a number (line) of more thin sounds, the second phrase - tone of a trill is more powerful ("til-til-til* tel-tel"), a final stroke - short sharp rise and recession of a sound ("chi-kuik").
Whether it is possible these three forms of song type C (C, C*, C#) to count dialect even if they meet in one territory (for example, C (fig. 2a) and С* (fig. 2c), recorded on Curonian spit) - a question complex (difficult). What origin of these variations of songs of one type - by wrong learning and their further fastening in a population training (song learning) of the following generations, or is result of "introduction" songs of migrants in local song culture - also it is possible to assume only.
Fig. 3. Difficultly differing types of songs of chaffinch (record on Curonian spit, the Kaliningrad region): a - song type D, b - song type F, c - song type G.
In samples song types which are difficult for differentiating among themselves, because of their similarity (fig. 3) have been found. We have attributed the given samples of songs to different types (D, F, G), but nevertheless they are similar among themselves in base structure of elements (especially in started singing and trill), which nevertheless are performance at singing by different manners, ways that does (makes) their distinct. Probably, it is the initial forms of the further modification being in themselves insignificantly transformed for precise revealing of type to which are attributed.
Thus, dialect forms of chaffinch song types (Fringilla coelebs L.) could meet within the limits of one local population in the same territory. Probably, it is result of constant mixing of vocal traditions at migrations (Slater, Ince, 1979, 1980; Espmark et al., 1989). Variability of phrases (motifs) of song types (lexical variability)
According to methods of the qualitative analysis of song types, we had been allocated two basic kinds of their distinctions: lexical and phonetic. Lexical variability of chaffinch song types (Fringilla coelebs L.) will consist in distinction of dictionary structure, phrases of song types.
In variants of song type A (fig. 4), recorded in different points of territory Curonian spit (area of biological research station "Fringilla"), characteristic for these song types the kind of a final stroke is replaced with other, usually making song type C (fig. 2a).
Song type W (fig. 5b) (it is recorded in Zvenigorod, the Moscow region) it is possible to consider as a combination of phrases of song types found on Curonian spit: started singing (the row of whistle elements) from song type B (fig. 5a) (1-st phrase), a final stroke - from song type Н (fig. 5c) (last phrase).
Fig. 4. The songs of one type, distinguished by strokes (record on Curonian spit): a - song type A, b - song type А*.
Fig. 5. A combination of phrases of song types B and H: a - song type B (record on Curonian spit), b - song type W (record in Zvenigorod, the Moscow region), c - song type Н (record on Curonian spit).
Trill elements of song type W (fig. 5b) under the form are similar to those of song type B (2-nd phrase of a trill), but finer on the size. Probably, the trill of song types W (fig. 5b) and B (fig. 5a) is dialect. In repertoires of the some males there were the "combined" songs consisting of phrases of other song types (fig. 6 and 7). Similar cases are described from experiments of song training (learning), when caught young chaffinch male (Fringilla coelebs L.) in first year of his life, in sensitive period, daily played different songs of two wild males. In the result, after a while the caught bird sang a song uniting phrases of songs two wild males (Nottebohm, 1967; Jellis, 1977). Probably, and us the marked cases are the certificate of non-standard learning of song types during their crystallization.
Fig. 6. Sonogram of the combined song type of chaffinch (Fringilla coelebs L.), recorded on Curonian spit (those types of songs of which phrases will consist also are shown): a - song type Е, b - song type I, c - song type E I.
The song type EI (fig. 6c) includes phrases of song type Е (fig. 6a) and song type I (fig. 6b). The song type ОС* (fig. 7c) will consist of phrases of song type O (fig. 7a) and song type С* (fig. 7b). All these samples of chaffinch songs (Fringilla coelebs L.) have been recorded in population of Curonian spit (both in repertoire of one individual, and at singing of different males).
Fig. 7. Improvisation in a combination of phrases at singing of one chaffinch male: a - song type O, b - song type C*, c - song type ОС*.
Such phenomena of lexical distinction of songs of one type within the limits of the certain population of some species of sparrow, bioacoustics explained many from the point of view of natural variations in repertoire of the birds based on vocal generalization and an individualization. The phenomena giving development songs (developmental interaction) can include cooperating processes of imitation (copying) and improvisation (temporarily - dependent copying, a dispersion young, drift, a regrouping of syllables, phrases) (Lemon, 1975; Burt and Beecher, 2000; Slater et al., 1984).
New song types can be considered as a new growth in a course of "a cultural mutation". Distribution song types in a population corresponds to the assumption, that birds copy them from other individuals of a species at random, and that less than 15 % of these new growths influence creation of new types of songs. One individual of chaffinch male (Fringilla coelebs L.) can have repertoire from 1-6 types of songs (Slater, Ince, 1979; Slater et al., 1980).
Thus, the examples of lexical distinctions of song types shown by us can be based on mistakes of training (song learning) and processes of improvisation at singing that it is possible to name a cultural mutation in limits of population song cultures. But such cases make small percent from all types of songs of a population.
Variability of elements (syllables) of song types (phonetic variability)
Phonetic (syllabic) variability of types chaffinch songs (Fringilla coelebs L.) consists in distinctions of a manner (ways, styles) performances, pronunciations of syllables, elements of one type - in distinction of their forms on sonograms). We considered change of the form (pronunciation) of elements of one type in different parts of songs: started singing (the row of whistle elements), a trill (an average part) and a final stroke.
Syllabic variability of started singing (the row of whistle elements) it is possible to show by the example of song type C (fig. 2). At all three forms of this song type elements of started singing arc-similar, but sound differently: at song type C* (fig. 2b) and С# (fig. 2c) have a little creaking a sound "vji - vji" (it is reflected in more rough form of syllables on sonogram), and at song type C (fig. 2a) they sound most precisely, high tone - "fuit-fuit-fuit", that confirms other pronunciation (phonetic distinction).
Fig. 8. Dialect forms of phrases from types of songs B and F: a - song type B (Curonian spit), b - song type BF (record in Michurinsk, the Tambov region), c - song type F (Curonian spit, the Kaliningrad region). +
Also elements of started singing in song type B (fig. 8a) (record on Curonian spit) and the "combined" song type BF (fig. 8b) (record in Michurinsk, the Tambov region) are modified in phonetic (have different style of singing, but the similar form on sonogram), therefore were considered dialect. Elements of a trill in difficultly differentiated types of songs D, F, G (fig. 3) are similar in the base, initial form, but nevertheless are performances by different manners, styles at singing.
It is possible to note phonetic variability of a trill in song type C (fig. 2). The second phrase of trill in song type С* (fig. 2c) as against others song forms will consist from "stick-similar" elements. And the first trill phrase of song type C# (fig. 2b) includes wider indistinct syllables. Syllabic variability of a stroke is distinctly visible by the example of song type V (fig. 1) and song type C (fig. 2), that already was partly discussed in section about dialect songs. The case of variability of a stroke in song type C# (fig. 2b) is especially interesting - last element (which at song type C (fig. 2a) the "triangular" form) is separated on three separate subelements. This phonetic difference changes a kind of a phrase as a whole so, and also changes the lexical party (side) of song type. It is sometimes difficult to differentiate new lexical variants from extreme phonetic variants of known syllabic types.
Thus, phonetic variability of syllables (elements) of song types consists, that their base (initial, the basic) structure can be uniform, similar, but modified at singing of different chaffinch males (Fringilla coelebs L.) both within the limits of one local population, and in populations in different territories.
Time-and-frequency parameters of local variants of song types
At comparison of the basic time-and-frequency parameters of local variants of song type C (tab. 1) the greatest difference had values of length of songs, the maximal frequency, number of syllables in song types, intervals between songs at singing (these parameters included rather wide range of distribution of the data). In other parameters (the minimal, average (median) frequency, length of trill elements) come to light the most distinguished values (are allocated by a font), but relative uniformity of the data nevertheless is observed.
Table 2. The quantitative analysis of dialect phrases of song types B and F
Type of song
(place of record)
Parameters of phraseВ
(Curonian spit)ВF 5
(Michurinsk)F
(Curonian spit)1 phrase
(started singing)2 phrase
(trill)В - similar elements (started singing)F- similar
elements
(trill)1 phrase
(started singing)1
phrase2
phraseThe length of phrase, sec1,13
±0,090,32
±0,050,62
±0,130,24
±0,021,17
±0,251,27
±0,11Max frequency, КHz6,57
±0,464,626
±0,327,3
±0,265,9
±0,527,44
±0,226,25
±0,2Min frequency, КHz2,017
±0,132,067
±0,13,1
±0,32,4
±0,121,9
±0,171,722
±0,14Median (average) frequency, КHz4,306
±0,243,17
±0,095,44
±0,23,8
±0,174,31
±0,1724,2
±0,09Number of syllables in phrase type11,43
±0,795,29
±0,765,2
±0,85
±06,4
±1,347,6
±0,7The length of syllables, sec0,058
±0,010,037
±0,0050,08
±0,0140,04
±0,0080,15
±0,010,12
±0,028Intervals between syllables,
sec0,048
±0,010,037
±0,0040,05
±0,010,02
±0,0050,05
±0,0080,06
±0,012
The note: specified are average value and a standard deviation of parameters of dialect phrases from statistical calculations for all songs of one type, which were reproduced chaffinch males in the given points of record; the strongest differences of phrases of one type counted a difference of parameters >0,5 KHz in frequency and >0,02 sec in length (are allocated by a font).
Table 1. The basic time-and-frequency parameters of local variants of chaffinch song type С
Type of song (token)Sample size (songs)The name of record placeThe length of song, secMin frequency, КHzMax frequency, КHzMedian (average) frequency, КHzNumber of syllables
in song type
Length of syllables of start singing, secLength of syllables trill, sec*Length of syllable of final stroke, sec*Intervals between songs,
sec1
phrase2
phraseС20Curonian spit2,074
±0,1181,627
±0,1319,698
±0.414,0996
±0,17317,25
±0,85 0,1203
±0,0050,068
±0,0030,1155
±0,0080,071
±0,0056,423
±1,9С#2Moscow2,73
±0,31,78
±0,18,096
±0,64,54
±0,4320
±1,730,22
±0,0060,076
±0,0030,12
±00,081
±0,0198,26C*3Curonian spit2,55
±0,0821,3
±0,377,751
±04,478
±019
±1,4 0,175
±0,0070,034
±0,010,084
±0,0010,915
±0,00222,9
The note: specified are average value and a standard deviation of parameters of song types from statistical calculations for all songs of one type, which were reproduced chaffinch males in the given points of record; the strongest differences counted a difference of parameters >0,5 KHz in frequency and >0,02 sec in length (are allocated by a font); * - the trill of song type C will consist of two phrases (fig. 2).
For chaffinch (Fringilla coelebs L.) selective training (song learning) of species specific songs in sensitive period is typically (Marler, 1956; Thorpe, 1958; Nottebohm, 1967). Therefore young birds do not learn songs of other species, and are guided by a time-and-frequency song range of their species. Probably, as a result of such rigid genetic determination species specific songs (Nottebohm, 1967; Simkin, 1983) is observed small variability of the basic time-and-frequency parameters of its different dialect forms (even at the account of a wide variation of values).
Results of the quantitative analysis of dialect forms of phrases of one type (tab. 2) are interesting. At comparison of the basic frequency time parameters of the first phrases of song types B and BF (fig. 8a and 8b) comes to light a significant difference in their length, the maximal, minimal and average (median) frequency. If to pay attention to the given measurements of a F-similar trill of song type BF (fig. 8b), in comparison with dialect it a phrase of started singing in song type F (record on Curonian spit) (fig. 8c), come to light similarities in such parameters as the minimal and average (median) frequency, intervals between syllables of a phrase.
Proceeding from the analysis carried out by us, it is possible to draw a conclusion, that the length of dialect phrases and their maximal frequency basically are various, and the minimal frequency and intervals between syllables appeared in the majority are similar. Values of others time-and-frequency parameters can to vary widely, but within the limits of the certain range.
It is necessary to note, that at the greater distinction of syllables form of dialect phrases on sonograms (up to occurrence of new syllabic patterns), the big distinctions of their basic time-and-frequency parameters are observed (for example, the first phrases in started singing of song types B (fig. 8a) and BF (8b)). Discussion
Many characteristics of birds singing are precisely depends from social training (song learning) traditions, receiving their vocal patterns by species specific imitation. Therefore, the tradition, custom should be taken into account at the analysis of geographical variability of vocal behaviour of birds (Mundinger, 1982; Kroodsma, Miller, 1982). In local populations of birds are formed certain song cultures, which are capable to change during time and to make dialect forms on all an area of species distribution. Steady dialects of birds in time - the phenomenon of conservatism of the vocal traditions, transmitted to the subsequent generations by means of vocal imitation. For many species patterns of variability as purchases of vocal traditions are a product of cultural evolution of birds (Mundinger, 1980). Cultural evolution is a parameter for species with a historical variety of patterns of microgeographical variability of vocalizations, and also for species of birds with differentiated syllables of songs. But it is not a parameter in morphological variability of birds (Lemon, 1975; Slater and Ince, 1979).
It is possible to draw a conclusion, that as a result of lexical (concerning phrases of songs) and phonetic (concerning to syllables, elements of song types) vocal variability in local populations of chaffinch (Fringilla coelebs L.) are formed certain song cultures, which are capable to change during time and to make dialect forms on all an area of distribution of a species.
At chaffinch (Fringilla coelebs L.) the song has a genetic basis (species-specific features) (Thorpe, 1958; Nottebohm; 1967), but by training (song learning), improvisation, mistakes of copying a variety of song types in a population constantly replenishes (Slater et al., 1979, 1984; Ince et al., 1980; Jellis, 1977).
Equally with a substantiation of distinctions and variability of song types of birds as product of mistakes of copying, improvisation at singing and transfers of features song traditions to the following generation during cultural evolution (Slater et al., 1979, 1984; Ince et al., 1980), there is an assumption of gradual phylogeny complication initial "ancient" (more simple on the structure) types of songs in more "perfect", complex (difficult) on structure song forms existing in a population equally with first, on the basis of occurrence of their geographical variability (biomorphism) (Simkin, 1983).
The basic time-and-frequency parameters of local variants of chaffinch song types (Fringilla coelebs L.) at the quantitative analysis had insignificant macrogeographical variability, that can speak about their importance in transfer and maintenance of species specific song traditions. The most stable parameters of local variants of songs of one type appeared minimal and average (median) frequency, values of length and intervals of trill elements.
Proceeding from samples, made by us in different regions of the European Russia (northwest and the central part), it is possible to draw a conclusion, that on some song types of chaffinch (Fringilla coelebs L.) exist subdialects (insignificant phonetic distinctions of syllables) - for example, on song types B, M, D, G, V, S, U, and dialects (different phonetic norms of types of songs) - C, C* and C#, F, I, J, N, and there are also stable types of song (in samples of different regions are invariable) - song type A.
Vocal norms (installation), likely, can be judged by quantity (amount) of individuals in a population, who adhering those or others song cultures (ways of singing) of different types of songs. If the majority of males sings any song type such style (a manner of performance), then it is will be vocal norm (installation) on this song type (or on type of a phrase) in the given population. But it is difficult to define vocal norms of a population at repertoires of great volume (when types of songs much) (Kroodsma, 1974).
Full dialect of chaffinch song (Fringilla coelebs L.) in the certain territory would be more correct to count presence of dialect forms on all known to us song types in a population (originally 22 types have been found), that, likely, it is almost impossible, because of sample limitation and constant mixing of vocal traditions of different populations as a result of migrations (Slater and Ince, 1979, 1980; Espmark et al., 1989).
The literature
Baker M. C. 1975. Song dialects and genetic differences in White-crowned Sparrows (Zonotrichia leucophrys). - Evolution, 29: 226-241.
Banin D. A., Byome I. R., Byome R. L. 1994. Seasonal variability of functional loading songs of birds. - The Bulletin of the Moscow university. Vol. 16, Biology, 3. Moscow: 40-42.
Baptista L. F. 1975. Song dialects and demes in sedentary populations of the White-crowned Sparrow (Zonotrihcia leucophrys nuttalli). - Univ. Calif., Berkeley, Publ. Zool., 105: 1-52.
Espmark Y. O., Lampe H. M., Bjerke T. K. 1989. Song conformity and continuity in song dialects of redwings Turdus iliacus and some ecological correlates. - Ornis Scand., 20: 1-12.
Ince S. A., Slater P. J. B., Weismann C. 1980. Changes with time in the song of a populations Chaffinches. - Condor, 82: 285-290.
Jellis R. 1977. Bird sounds and their meaning. Published by the British Broadcasting corporation. Cambridge, 256 p. Kreutzer M. 1974. Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes). - Rev. Comp. Anim., 8: 270-286.
Kroodsma D. E. 1974. Song learning, dialects, and dispersal in the Bewick ` s Wren. - Tierpsychol., 35: 352-380.
Kroodsma D. E., Miller E. H., Quellet H. 1982. Communication and behavior an interdisciplinary series. Vol. 2: Song leaning and its consequence / D.E. Kroodsma, E.H. Miller. - London: Academic press, 347 p. Kurath H. 1972. Studies in Areal Linguistics. - Indiana Univ. Press, Bloomington.
Lemon L. E. 1975. How birds develop song dialects. - Condor, 77: 385-406.
Marler P. 1952. Variation in the song of the Chaffinch Fringilla coelebs. - Ibis, 98: 458-472. Marler P. 1956a. The voice of the chaffinch and its function as a language. - Ibis, 98: 231-261. Marler P., Tamura M. 1962. Song dialects in three populations of White-crowned Sparrows. - Condor, 64: 368-377.
McGregor P. K., Krebs J. R. 1982. Song types in a population of great tits (Parus major): their distribution, abundance and acquisition by individuals. - Behavoiur, 79: 126-152.
Mundinger P. C. 1979. Call learning in the Carduelinae: Ethological and systematic considerations. - Syst. Zool., 28: 270-283.
Mundinger P. C. 1980. Animal cultures and a general theory of cultural evolution. - Ethol. Sociobiol., 1: 183-223.
Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds. In: Kroodsma D. E. Miller E. H. (eds) Acoustic communication in birds, vol. 2, New York: Academic Press, P. 147-208.
Nottebohm F. 1969. The song of the Chingolo, Zonotrichia capensis, in Argentina: Description and evaluation of a system of dialects. - Condor, 71: 299-315.
Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs. - P. 265-280.
Payne R. B. 1981. Population structure and social behavior: models for testing ecological significance of song dialects in birds. In "Natural Selection and Social behavior: Recent Research and New Theory" (R.D. Alexander and D.W. Tinkle, eds.), Chiron, New York: 108-119. Poulsen H. 1951. Inheritance and leaning in the song of the Chaffinch (Fringilla coelebs). - Behaviour, 3: 216-228.
Promptov A. N. 1930. Geographical variability of chaffinch song in connection with the common questions of seasonal flights of birds. - Zool. journ. № 10 (3): 17-40.
Simkin G. N. 1972. About biological value of bird singing. - The Bulletin of the Moscow university. № 1. Moscow: 34-43. Simkin G. N. 1983. The typological organization and population phylogeny of bird songs. - Bulletin of Moscow community investigators of nature. Section of biology. V. 88. № 1. Moscow: 15-27.
Slater P. J. B., Ince S. A. 1979. Cultural evolution in chaffinch song. - Behaviour, 7: 146-166.
Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals. - Behaviour, 75: 207-218. Slater P. J. B. 1981. Chaffinch song repertoires: observations, experiments and a discussion of their significance. - Z. Tiërpsychol., 72: 177-184.
Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects. - Behaviour, 88: 76-97.
Sick H. 1939. Ueber die Dialektbildung beirn Regenruf des Buchfinken.- J. Orn., 87: 568-592.
Thielcke G. 1961. Stammesgeschichte und geographische Variation des Gesanges unserer Baumläufer. - Verh Ornithol. Ges Bayern, 14: 39-74.
Thielcke G. 1965. Gesangsgeographische Variation des Gartenbaumläufers (Certhia brachydactyla) im Hinblick auf das Artbildungsproblem. - Z. Tierpsychol., 22: 542-566.
Thielcke G. 1969. Geographic variation in bird vocalizations. In "Bird Vocalizations" (R. A. Hinde, ed.), Cambridge Univ. Press., London and New York: 311-340.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs. - Ibis, 100: 535-570.
+
Degree of prevalence of different song types of chaffinch (Fringilla coelebs L.)
in populations of the European Russia
O. A. Astakhova, I. R. Byome
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: beme@nm.ru
chaffinch@bk.ru
Abstract
At studying song organizations of chaffinch (Fringilla coelebs L.) are found out unique typological parities (ratio) of song in different local populations of a species-specific area. The degree of prevalence of different chaffinch song types can not correspond in percentage proceeding from their general (common) number in a population. Thus in different areas of Russia are forming original (different) song cultures of chaffinch, consisting of the certain set and a parity (ratio) of song types, frequently including so-called dialect song forms, that have been observed. The complex interrelation of geographical variability and structural variability of chaffinch song in many respects gives a support at an evolutionary view in the given aspect.
Keywords: species-specific song of chaffinch, different song types, song cultures, degree of prevalence of song types in populations.
Introduction
At many species of sparrow (Passeriformes) males have repertoires from two and more variants of species-specific song (Hartshorne, 1973). Some variants or types of songs are individually various and divided (shared) between individuals of a population (Slater, 1981; McGregor, Krebs, 1982). According to the proof from many laboratory researches the distributed (allocated) song type arises through vocal imitation (Kroodsma, 1978). It is interesting to clear, as this process operates in distribution of different types of songs of chaffinch (Fringilla coelebs L.). The samples of the songs, which have been recorded in several local populations of chaffinch in territory of distribution of a species are investigated.
One of problems (tasks) has consisted in revealing geographical variability of chaffinch song. Thus attempt to establish macrogeographical distribution and a variety of song types is done (made). Most the general (common) and widely widespread types of songs of chaffinch are compared in structural variability in local territory to variability of the certain area of distribution of a species. At studying microgeographical distribution of chaffinch song in local territory, divided (shared) of the general (common) types of songs into small distance between males comes to light. There are questions: as far as different song types of chaffinch are distributed independently from each other, and whether there are rare songs in an equal parity (ratio) in different territories? Individuals of chaffinch males in the repertoire can have 1-6 types (variants) of species-specific song. Usually the song of chaffinch will consist of three parts: started singing (it is heard as the row of whistle sounds of high tone), elements of trill (sounds are quickly poured each other and usually tone hardly is lower) and a sharp short stroke in the end (Marler, 1952; Thorpe, 1958, Nottebohm, 1969).
Material and methods
In northwest (Curonian spit, the Kaliningrad region) and central (Zvenigorod, Moscow, Michurinsk) parts of the European Russia have been made tape recorder records of singing males of chaffinch during the spring-and-summer period of 2005-2006 (N=218 of males). Different local populations of chaffinch (Fringilla coelebs L.), researched by us, are removed approximately on 1000 km from each other.
Further, sonograms of songs were analyzed with the help of computer program Avisoft-SaSLab Light. In total it has been analyzed about five thousand songs. The songs similar in two or all three parts (started singing, a trill, and a final stroke) we considered as one type. Types of songs were marked by Latin letters.
In samples on Curonian spit (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). At record, songs of one type met in different points of territory (was considered, that belong to repertoires of different males), therefore alongside with the letter were designated by numbers in ascending order (for example, А1, А2, А3, etc.).
In populations of the central part of the European Russia (N=65 of males) 15 different types of songs (A B C D F G I J M N S U V W Y), from which 12 types of songs were similar in the structure to corresponding types on Curonian spit are found, but frequently they had the modified elements on sonograms, therefore were considered as dialect forms (B C D F G I J M N S U V). 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. Results and discussion
It is possible to make a percentage parity (ratio) of the typological organization of chaffinch song (Fringilla coelebs L.) in local populations (table 1).
Both in northwest, and in the center of the European Russia the most widespread appeared song type C (fig. 2.2, 2.3). The second on prevalence in local populations the song type I (fig. 2.5) was. Song type R (fig. 2.9) and song type M (fig. 2.6, 2.7) are more often, than others met at record only on Curonian spit. But song culture of a population of chaffinch is characterized by different types of songs, which have the big sample size: A, F, S, O, T, J (fig. 1, 2). Fig. 1. Most frequently met types of songs of chaffinch (Fringilla coelebs L.) in different regions of the European Russia: without an asterisk - types of songs from a population of northwest of the European Russia (Curonian spit); * - types of songs from samples of the central part of the European Russia; % - percent from the general (common) number of song types in a population.
To have more precise representation about types of chaffinch songs, which we discuss, it is possible to cite (to show) sonograms their samples (fig. 2).
Fig. 2. The most widespread types of songs of chaffinch (Fringilla coelebs L.) in a population of northwest of the European Russia (Curonian spit): 1 - song type A, 2 - song type C, 3 - song type C *, 4 - song type F, 5 - song type I, 6 - song type М1, 7 - song type М12, 8 - song type O, 9 - song type R, 10 - song type S, 11 - song type J.
Table 1. A percentage comparison of chaffinch song types (Fringilla coelebs L.) in different regions of the European Russia
Type of song
(token)Northwest of European Russia
(Curonian spit)
Center of European Russia
(Moscow, Zvenigorod, Michurinsk)
Number of song types in samplePercent from the common number
of song type in a populationNumber of song types in samplePercent from the common number
of song type in a populationA189,84 %88,6 %B52,73 %88,6 %C3820,8 %2425,8 %D116,01 %33,2 %Е73,82 %F168,74 %55,4 %G105,5 %11,07 %H63,28 %I2212,02 %1111,83 %J137,1 %1212,9 %K31,64 %L10,5 %M2010,93 %22,15 %N84,4 %55,4 %O189,84 %P1R2413,1 %Q10,55 %S179,3 %22,15 %T147,7 %U84,4 %33,2 %V21,1 %22,15 %W33,2 %Volume of sample individuals (n)15865Volume of sample song types (n)18393
The note: the volume of sample includes all recorded males in populations (the bottom line); a percentage parity (ratio) expected proceeding from the general (common) number of song types in samples of populations in northwest of the European Russia - 183 song types, in the central part of European Russia - 93 song types (that is repertoires of males with small samples were taken into account also); thus it is necessary to understand, that songs of one type from everyone male had sample on the average (n> 20); the most widespread (frequently met) in populations song types (with sample n> 15) are allocated (distinguished) by a font.
At close (attentive) viewing of sonograms (fig. 2) it is possible to allocate (to find) a number (row) of features in structure of songs, characteristic only for the certain song type. It is necessary to remind, that we considered to songs of one type, what are similar on two or all three parts of songs (started singing or whistle elements, a trill and a final stroke), which (every song part) also can include one or two phrases (the elements similar under the form). The whistle elements (started singing) of chaffinch songs (Fringilla coelebs L.) can be as arc-similar, and narrow linear forms. But on hearing all of them are perceived by higher tone, as if a row of whistle sounds "fuit-fuit-fuit". The trill with equal frequency can consist both of one phrase, and from two phrases. Elements of a trill can be completely different in form, consist of two and more subelements. But if to look narrowly (more attentive), nevertheless it is possible to notice, that in many respects on the structure elements are similar - only their primary basis in different types differently performance at singing. The same is possible to tell about a final stroke - usually sharp on the hearing, consisting from stick-similar of elements on sonograms, has much in common at different types of songs, but also is individual for everyone song pattern.
For full representation of song cultures of different local populations of chaffinch (Fringilla coelebs L.) it is necessary to cite (to show) samples of the most widespread types of songs in territory of the center of the European Russia, which can not coincide with a degree of their prevalence in other populations (fig. 3).
Fig. 3. The most widespread types of songs of chaffinch (Fringilla coelebs L.) in populations of the central part of the European Russia: 1 - song type C, 2 - song type C*, 3 - song type I, 4 - song type J, 5 - song type A, 6 - song type B.
Song type C (fig. 3.1) (its version type C* (fig. 3.2)) and song type I (fig. 3.3) appeared in the majority both in samples in northwest, and in the center of the European Russia (tab. 1). But song type I (fig. 3.3) - the dialect form in relation to songs of this type on Curonian spit. The song type J (fig. 3.4) also is distributed enough in the center of the European Russia, but in northwest this song type was average on occurrence. Types of songs A, B (fig. 3.5, 3.6) also have average prevalence. These are the basic song patterns, which frequently met in populations of all two regions of Russia where research (fig. 1). Rare song types of chaffinch are V, K, Q, L (on Curonian spit) (fig. 4, 5). In the center of the European Russia rare types of songs could be and what are distributed enough in northwest of Russia (G, M, S) (fig. 4, 6), and any types of songs, in comparison with Curonian spit, and did not meet at all (E, H, O, R, T) (fig. 4, 7). It can speak about different and the certain originality of song cultures of different local populations of chaffinch.
Fig. 4. Seldom met song types of chaffinch (Fringilla coelebs L.) in different regions of the European Russia: without an asterisk - types of songs from a population of northwest of the European Russia (Curonian spit); * - types of songs from samples of the central part of the European Russia; % - percent from the general (common) number of song types in a population.
Thus, some song types of chaffinch, the most widespread in one local population, could be rare in samples of other population. Probably, as a result of migrations there is a constant mixing vocal traditions (Slater, Ince, 1979, 1980; Espmark et al., 1989).
Than it is possible to prove a rarity of these types of chaffinch songs (fig. 5, fig. 6, fig. 7.1 and 7.3)? Probably, such song structure was not in the greater degree characteristic for species specific song of chaffinch (Fringilla coelebs L.), therefore it had no "support" for the big distribution to "communicative" groups of a population. But can be, it is simple song types of migrants from other populations, which song culture includes their majority.
Conclusions
Geographical variability of song is obvious at many species of birds. The typological parity (ratio), the certain set of song patterns in local populations of birds with the big repertoire (including it is a lot of types of songs) is possible to characterize as mosaic distribution (Thielcke, 1965, 1969), but a number (line, row) of researchers do not support the similar description of bird song variability in territory. It is possible to track change of bird song from a place to a place and to draw a conclusion, that geographical variability - result of accumulation of mistakes of copying at training of songs (song learning) from generation to generation (Catchpole, 1995). But big song repertoires of birds interfere with objective definition of dialect borders within the limits, of which there is a traditional training of song components, characteristic this population (Kroodsma, 1974; Kreutzer, 1974).
In local populations of birds are formed certain (determined) song cultures, which are capable to change in during time and to make vocal dialect forms of birds on all an area of distribution of a species. Steady in time dialects of birds - the phenomenon of conservatism of the vocal traditions, transmitted to the subsequent generations by means of vocal imitation. For many species of bird's patterns of variability as purchases of vocal traditions is product of cultural evolution (Mundinger, 1980).
Fig. 5. Sonograms of rare song types of chaffinch (Fringilla coelebs L.) (are recorded on Curonian spit, the Kaliningrad region): 1 - song type K, 2 - song type L, 3 - song type V, 4 - song type Q.
For comparison we shall result (show, cite) individual types of chaffinch songs (Fringilla coelebs L.) in populations of the central part of the European Russia (fig. 6) (at record met in repertoire of 1-2 males). References Catchpole С. K. 1995. Bird song: biological themes and variations. Cambridge: Cambridge University Press. 248 p.
Espmark Y. O., Lampe H. M., Bjerke T. K. 1989. Song conformity and continuity in song dialects of redwings Turdus iliacus and some ecological correlates // Ornis Scand. № 20. P. 1-12.
Hartshorne C. 1973. Born to sing. Bloomington: Indiana University Press. 132 p.
Kreutzer M. 1974. Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes) // Rev. Comp. Anim. № 8. P. 270-286.
Kroodsma D. E. 1974. Song learning, dialects, and dispersal in the Bewick`s Wren // Tierpsychol. № 35. P. 352-380.
Fig. 6. Rare song types of chaffinch (Fringilla coelebs L.) populations of the Central part of the European Russia: 1 - song type G, 2 - song type M 1 (record in Michurinsk, the Tambov region), 3 - song type S, 4 - song type V (record in Zvenigorod, the Moscow region).
Fig. 7. Song types of chaffinch (Fringilla coelebs L.), found only in samples on Curonian spit (the Kaliningrad region): 1 - song type Н, 2 - song type O, 3 - song type Е, 4 - song type Т (types of songs Н and Е were rare - 3-5 % of prevalence from all types of songs in a population).
Kroodsma D. E. 1978. Aspects of learning in the ontogeny of bird song: where, from whom, when, how many, which and how accurately? / Eds. Burghardt G., Bekoff M. New-York: Garland. P. 215-230.
Marler P.1952. Variation in the song of the Chaffinch Fringilla coelebs // Ibis. № 98. P. 458-472. McGregor P. K., Krebs J. R. 1982. Song types in a population of great tits (Parus major): their distribution, abundance and acquisition by individuals // Behaviour. № 79. P. 126-152.
Mundinger P. C. 1980. Animal cultures and a general theory of cultural evolution // Ethol. Sociobiol. № 1. P. 183-223.
Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds / Еds. Kroodsma D.E., Miller E.H. New-York: Academic Press. P. 147-208.
Nottebohm F.1969a. The "critical period" for song leaning in birds // Ibis. № 111. P. 386-387.
Slater P. J. B., Ince S. A. 1979. Cultural evolution in chaffinch song // Behaviour. № 71. P. 146-166.
Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. P. 207-218. Slater P. J. B. 1981. Chaffinch song repertoires: observations, experiments and a discussion of thier significance // Z. Tiërpsychol. № 72. P.177-184.
Thielcke G. 1969. Geographic variation in bird vocalizations / Eds. Hinde R.A. London, New-York: Cambridge University Press. P. 311-340.
Thielcke G. 1965. Gesangsgeographische Variation des Gartenbaumläufers (Certhia brachydactyla) im Hinblick auf das Artbildungsproblem // Z. Tierpsychol. № 22. P. 542-566.
Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. P. 535-570. ++
Results of the comparative analysis of chaffinch song (Fringilla coelebs L.) in geographical populations of the south of Ukraine (Crimea) and the European Russia
Astakhova О. A., Byome I. R.
Moscow state university of М. В. Lomonosov biological faculty, department of vertebrate zoology
Russia, 119992 Moscow, Lenin mountains, 1/12
e-mail: chaffinch@bk.ru
beme@nm.ru
Vocal variability of chaffinch (Fringilla coelebs L.) in different local and geographical (located on the big distances) populations is already for a long time the found out fact and widespread enough subject of studying in bioacoustics (Marler, 1952; Thielcke, 1969; Slater et al., 1984, Simkin, 1983; Bergmann, 1993). But character of the typological organization of "song culture" of population in different chaffinch subspecies - European (Fringilla coelebs coelebs L.) and Crimean (Fringilla coelebs solomkoi Menz.), living in geographical populations on distance of 1500-1850 km is interesting. From samples of chaffinch songs in the south of Ukraine (Crimea) (N=26 males) 16 separate types of songs have been revealed, from which only one song type (В) (6,25 %) appeared the general (common) or shared (completely similar) with analogous to it song type in populations of chaffinch in the European Russia. Thus 9 types of songs (56,25 %) were the general (common) or shared, but modified ("dialect") in the general (common) structure and in the form of elements in relation to corresponding (same) types of songs in the European Russia (song types Е, O, Т (18,75 %) - in relation to Curonian spit of Baltic sea (northwest, the Kaliningrad region) and song types V, Y, BF, CM, GC, OE (37,5 %) - in relation to the central part of the European Russia (Zvenigorod, Moscow, Michurinsk). The others 6 song types of chaffinch (37,5 %) appeared poorly similar (original) in the qualitative aspect (MI (N), MI (B), ME (I) (A), O*I, M (O) I, CME) and are named on phrases (parts) or as a whole of those types of songs, which "reminded" in the structure (the form of elements). Presence of the general (common) or shared (similar) song types of chaffinch shows coherence (interconnected, mutual) of geographical populations and their constant interaction (in the greater degree, at migrations), but on the basis of it there is also a formation local "song culture" (the certain composition of song types, their percentage parity (ratio)) in local populations, differentiation of song types in different variants at song learning ("mistakes" of copying) and improvisation at singing by young individuals of birds (Mundinger, 1982; Catchpole, 1995). Therefore different subspecies of chaffinch can be characterized distinguished by "song cultures" (different ways or manners of singing of species specific song), though "social" cultural traditions and a song of birds can be considered as "parallel" in the existence to the morphological attributes (signs), particularly determining a rank of a species (subspecies) (Lemon, 1975), but, nevertheless, song structure also is capable to reflect a degree of common (general) or separation in different geographical populations, can to mark stages of formation of populations in a separate species.
At the quantitative analysis of chaffinch songs has come to light, that in the south of Ukraine (Crimea) average (median) frequency (КHz) of song types is much higher, than in populations of the European Russia: in northwest (Curonian spit, the Kaliningrad region) (N=279 of songs) median=4,10±0,4 КHz, in the central part of the European Russia (Zvenigorod, Moscow, Michurinsk) (N=73 of songs) median=4,096±0,29 КHz, in the south of Ukraine (Crimea) (N=85 of songs) median=4,63±0,23 КHz. Probably, it is connected to warmer climate in the south, which promotes the greater song activity of birds. Thus the minimal and maximal frequency (KHz) of chaffinch songs has remained more stable in the values: in northern population (Curonian spit) min=1,63±0,063 КHz, max=7,683±0,42 КHz, in the center of the European Russia min=1,7±0,17 КHz, max=8,06±0,58 КHz and in a southern population (Crimea) min=1,85±0,14 КHz, max=7,98±0,74 КHz. It is necessary to pay attention to length (duration) (sec) of species specific song of chaffinch in different geographical populations: in the north 1,97±0,23 sec, in the center of the European Russia 2,59±0,29 sec, in the south of Ukraine (Crimea) 2,26±0,55 sec. In northern population of chaffinch (Curonian spit) a species specific song are shorter (sec) and lower frequency range (KHz), than in the center of the European Russia and in the south of Ukraine (Crimea). Probably, it is connected by that on the north colder climate, where the metabolism of an organism can be or lower, or pass with the greater economy of expenses of forces (energy) at realization of songs of chaffinch in his (its) northern population.
Literature
1. Marler P. 1952. Variation in the song of the Chaffinch Fringilla coelebs // Ibis. - № 94. - P.458-472. 2. Thielcke G. 1969. Geographic variation in bird vocalizations // Bird Vocalizations / Eds. R.A. Hinde. London, New York. - P. 311-340.
3. Slater P. J. B., Clement F. A., Goodfellow D. J. 1984. Local and regional variations in chaffinch song and the question of dialects // Behaviour. - № 88.- 76-97.
4. Catchpole C.K., Slater P.J.B. 1995. Birdsong. Biological themes and variation// Cambridge: Cambridge University Press.- 248 p.
5. Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds // Acoustic communication in birds / Eds. D.E. Kroodsma, E.H. Miller. New York. - P. 147-208.
6. Bergmann H.-H. Der Buchfink: Neues Über einen bekannten Sänger / Hans-Heiner Bergmann [Zeichn.: F. Müller]. - Wiesbaden: Aula-Verl, 1993. - 142 p.
Chaffinch (Fringilla coelebs L.) song in populations of the East Europe
O. A. Astakhova
Moscow state university of M. V. Lomonosov
biology faculty, department of vertebrate zoology
Russia, 119992 Moscow, Leninskye Gory 1/12
e-mail: chaffinch@bk.ru
Abstract
At studying of song organizations of chaffinch (Fringilla coelebs L.) are found out unique parities of song types in different local populations of a species-specific area. Thus in different areas of the East Europe are forming original (different) song cultures of chaffinch - the certain set of song types including so-called "dialect song forms". The complex interrelation of geographical variability of chaffinch song in many respects gives a support at an evolutionary view in the given aspect.
Keywords: species-specific song of chaffinch, different song types, song cultures, degree of prevalence of song types in populations.
Introduction
At many species of sparrow (Passeriformes) males have repertoires from two and more variants of species-specific song [1]. Some variants (types) of song are individually various and divided (shared) between individuals of a population [2, 3]. According to the proof from many laboratory researches song type arises through vocal imitation [4]. It is interesting to clear, as this process operates in distribution of different song types of chaffinch (Fringilla coelebs L.). Song of chaffinch is genetically determined. The samples of the songs which have been recorded in several local populations of chaffinch are investigated. Individuals of chaffinch (males) in the repertoire can have 1-6 types (variants) of species-specific song. Usually the song of chaffinch will consist of three parts: started singing (it is heard as the row of whistle sounds of high tone), elements of trill (sounds are quickly poured each other and usually are tone hardly lower) and a sharp short stroke at the end [5-7]. Shown the distribution of the song types of chaffinch in the populations of Eastern Europe removed at 1500-1800 km.
Material and methods
In northwest (Curonian spit, the Kaliningrad region) and central (Zvenigorod, Moscow, Michurinsk) parts of the European Russia have been made tape recorder records of singing males of chaffinch during the spring-and-summer period of 2005-2006 (N=218 males). Different local populations of chaffinch (Fringilla coelebs L.), researched by us, are removed approximately on 1000 km from each other.
Further, sonograms of songs were analyzed with the help of computer program Avisoft-SaSLab Light. In total it has been analyzed about five thousand songs. The songs similar in two or all three parts (started singing, a trill, and a final stroke) we considered as one type. Types of songs were marked by Latin letters.
In May, 2007 in the south of Ukraine (Crimea) (are removed approximately on 1500 km from the European Russia) have been made tape records of singing males of chaffinch, concerning (relate) to separate subspecies is chaffinch crimean (Fringilla coelebs solomkoi Menz.) - sample contained approximately 26 males, from which we tried to fix full song repertoire of each chaffinch individual. Results and discussion
Song types of chaffinch distinguished from each other in song structure (syntax), phonetics or a pronunciation of vocal elements, but similar in beat, duration, a way of differentiation on a part, phrases, in the common (similar) frequency range (which appeared stable and, most likely, is genetically determined in all populations of chaffinch as species) [8].
In samples on Curonian spit (the Kaliningrad region) (N=158 males) we allocate (distinguish) 22 types of song (A B C D E F G H J I K L M N O P Q R S T U V). At record, songs of one type met in different points of territory (was considered, that belong to repertoires of different males), therefore alongside with the letter were designated by numbers in ascending order (for example, А1, А2, А3, etc.).
In populations of the central part of the European Russia (N=65 males) 15 different types of song (A B C D F G I J M N S U V W Y), from which 12 types of song were similar in the structure to corresponding types on Curonian spit are found, but frequently they had the modified elements on sonograms, therefore were considered as dialect forms (B C D F G I J M N S U V). Local variants of vocalizations of birds can be considered as analogy of human speech and to name "dialects" [9, 10]. 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. From samples of chaffinch songs in the south of Ukraine (Crimea) (N=26 males) 16 separate types of songs have been revealed, from which only one song type (В) (6,25 %) appeared the common or shared (completely similar) with analogous to it song type in populations of chaffinch in the European Russia. Thus 9 types of songs (56,25 %) were the general (common) or shared, but modified ("dialect") in the general (common) structure and in the form of elements in relation to corresponding (same) types of songs in the European Russia (song types Е, O, Т (18,75 %) - in relation to Curonian spit of Baltic sea (northwest, the Kaliningrad region) and song types V, Y, BF, CM, GC, OE (37,5 %) - in relation to the central part of the European Russia (Zvenigorod, Moscow, Michurinsk). The others 6 song types of chaffinch (37,5 %) appeared poorly similar (original) in the qualitative aspect (MI (N), MI (B), ME (I) (A), O*I, M (O) I, CME) and are named on phrases (parts) or as a whole of those types of songs, which "reminded" in the structure (the form of elements) (Figure 1).
At the quantitative analysis of chaffinch songs has come to light, that in the south of Ukraine (Crimea) average (median) frequency (КHz) of song types is much higher, than in populations of the European Russia: in northwest (Curonian spit, the Kaliningrad region) (N=279 of songs) median=4,10±0,4 КHz, in the central part of the European Russia (Zvenigorod, Moscow, Michurinsk) (N=73 of songs) median=4,096±0,29 КHz, in the south of Ukraine (Crimea) (N=85 songs) median=4,63±0,23 КHz. Probably, it is connected to warmer climate in the south, which promotes the greater song activity of birds. Thus the minimal and maximal frequency (KHz) of chaffinch songs has remained more stable in the values: in northern population (Curonian spit) min=1,63±0,063 КHz, max=7,683±0,42 КHz, in the center of the European Russia min=1,7±0,17 КHz, max=8,06±0,58 КHz and in a southern population (Crimea) min=1,85±0,14 КHz, max=7,98±0,74 КHz. It is necessary to pay attention to length (duration) (sec) of species specific song of chaffinch in different geographical populations: in the north 1,97±0,23 sec, in the center of the European Russia 2,59±0,29 sec, in the south of Ukraine (Crimea) 2,26±0,55 sec (Table 1). In northern population of chaffinch (Curonian spit) a species specific song are shorter (sec) and lower frequency range (KHz), than in the center of the European Russia and in the south of Ukraine (Crimea). Probably, it is connected by that on the north colder climate, where the metabolism of an organism can be or lower, or pass with the greater economy of expenses of forces (energy) at realization of songs of chaffinch in his (its) northern population.
Figure 1. Chaffinch (Fringilla coelebs L.) song in populations of the East Europe
At the analysis of Table 2 it is possible to draw conclusions:
1. 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). Subdialects of chaffinch song types in the greater degree are found in those populations which are in closer communications (connections) (Figure 1).
2. 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) were observed (revealed) a lot of original types of songs (few similar, dialect, not having analogues), determining a degree of "alien" of song cultures (their distinction) at given populations of chaffinch.
3. Originality (allogeneity, distinguish) of song cultures 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 another).
Figure 2. "Netted" evolution of chaffinch song
Figure 3. The correlations of chaffinch song in populations of Eastern Europe
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) (Figure 2).
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 [11-13]. 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 specific 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
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. Table 1. Comparison of basic parameters of song types of chaffinch (Fringilla coelebs L.)
in different geographical populations of the East Europe (Astakhova, Byome, 2007).
Parameters of
song types
Geographical population Average values of parameters of song types
Duration of song, secMin frequency, KHzMax
frequency, KHzMedian
(average) frequency, KHzDegree of distribution
of song types, %Number of syllables
in songsDuration of start singing, secDuration of trill, secDuration of final stroke, secIntervals between songs, sесThe Northwest of the European Russia (Curonian spit)1,97
±0,231,63
±0,0637,683
±0,424,10
±0,146,82
±515,6
±2,670,47
±0,260,931
±0,4140,4
±0,16,92
±0,81The center of the European Russia (Moscow, Zvenigorod, Michurinsk)2,59
±0,291,7
±0,178,06
±0,584,096
±0,296,83
±6,6220,11
±4,60,88
±0,340,99
±0,320,62
±0,247,3
±1,9The south of Ukraine
(Crimea)2,26
±0,551,85
±0,147,98
±0,744,63
±0,236,6
±517,5
±4,860,72
±0,21,02
±0,560,34
±0,16,53
±1,68Total
(the common for song of species)2,27
±0,361,73
±0,127,91
±0,584,28
±0,226,75
±5,5417,74
±4,040,69
±0,270,98
±0,430,45
±0,156,92
±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 songs, in populations of the center of the European Russia - 73 songs of different types, from samples in the south of Ukraine (Crimea) - 85 songs; differences of parameters, a difference between which >0,5 КHz in frequency and >0,3 sec in length of song parts (are allocated (distinguished) by a font) were taken into account.
Table 2. Degree of a generality of chaffinch songs types (Fringilla coelebs L.) in different geographical populations of Europe
Character of
variability
song types Geographical population Percentage parity (ratio) of song types in populations (%)The 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**29,24,2**6,256,25*Subdialects 225,9*14,8**29,225**2537,5*Dialects 118,5*22,2**20,820,8**37,531,5*Not having similar 329,7*59,3**20,850**31,2524,75*
The note: dialects1 - different phonetic norms of song types; subdialects2 - small phonetic distinctions of song types; * - percent of the dialect (modified) or not modified (shared) 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 not modified (shared) 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 not modified (shared) types of songs in the given population in relation to song types of northwest of the European Russia; not having similar - percent of special types of the songs not having analogues and determining a degree of an originality of song culture of one population in relation to another (last line); the greatest values (30 - 50 %) are allocated (distinguished) by a font.
Geographical variability of song is obvious at many species of birds. The typological parity (ratio), the certain set of song patterns in local populations of birds with the big repertoire (including it is a lot of types of songs) is possible to characterize as mosaic distribution [14], but a number (line, row) of researchers do not support the similar description of bird song variability in territory. It is possible to track change of bird song from a place to a place and to draw a conclusion, that geographical variability - result of accumulation of mistakes of copying at training of songs (song learning) from generation to generation [15]. But big song repertoires of birds interfere with objective definition of dialect borders within the limits, of which there is a traditional training of song components, characteristic this population [16, 17].
In local populations of birds are formed certain (determined) song cultures, which are capable to change in during time and to make vocal dialect forms of birds on all an area of distribution of a species. Steady in time dialects of birds - the phenomenon of conservatism of the vocal traditions, transmitted to the subsequent generations by means of vocal imitation. For many species of bird's patterns of variability as purchases of vocal traditions is product of cultural evolution. 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, instead 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), phyletical (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. References [1] Hartshorne C. (1973) Born to sing. Bloomington: Indiana University Press., 132.
[2] Slater P. J. B. (1981) Chaffinch song repertoires: observations, experiments and a discussion of thier significance. Z. Tiërpsychol, 72, 177-184.
[3] McGregor P. K., Krebs J. R. (1982) Song types in a population of great tits (Parus major): their distribution, abundance and acquisition by individuals. Behaviour, 79, 126-152.
[4] Kroodsma D. E. (1978) Aspects of learning in the ontogeny of bird song: where, from whom, when, how many, which and how accurately? / Eds. Burghardt G., Bekoff M. New-York: Garland, 215-230.
[5] Marler P. (1952) Variation in the song of the Chaffinch Fringilla coelebs. Ibis, 98, 458-472. [6] Thorpe W. H. (1958) The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs. Ibis, 100, 535-570.
[7] Nottebohm F. (1969a) The "critical period" for song leaning in birds. Ibis, 111, 386-387.
[8] Simkin, G.N. (1983) The typological organization and populational phylogeny of birds song. The bulletin of Moscow society of nature researchers. Section of Biology, 88, 15-27. [9] Mundinger P. C. (1980) Animal cultures and a general theory of cultural evolution. Ethol. Sociobiol,. 1, 183-223.
[10] Mundinger P. C. (1982) Microgeographic and macrogeographic variation in acquired vocalizations of birds / Еds. Kroodsma D.E., Miller E.H. New-York: Academic Press., 147-208.
[11] Thielcke G. (1969) Geographic variation in bird vocalizations / Eds. Hinde R.A. London, New-York: Cambridge University Press., 311-340.
[12] Slater P. J. B., Ince S. A. (1979) Cultural evolution in chaffinch song. Behaviour, 71, 146-166.
[13] Slater P. J. B., Clement F. A., Goodfellow D. J. (1984). Local and regional variations in chaffinch song and the question of dialects. Behaviour, 88, 76-97.
[14] Thielcke G. (1965) Gesangsgeographische Variation des Gartenbaumläufers (Certhia brachydactyla) im Hinblick auf das Artbildungsproblem. Z. Tierpsychol, 22, 542-566.
[15] Catchpole С. K. (1995) Bird song: biological themes and variations. Cambridge: Cambridge University Press, 248.
[16] Kroodsma D. E. (1974) Song learning, dialects, and dispersal in the Bewick`s Wren. Tierpsychol., 35, 352-380.
[17] Kreutzer M. (1974) Stereotypie et varianttions dans les chants de proclamation territoriale chez le Troglodyte (Troglodytes troglodytes). Rev. Comp. Anim., 8, 270-286. ++
Combinative variability at different levels of the organization of a matter of the Nature: chromosomes of cells of an organism and vocal realization of birds (Aves)
Astakhova О. A.
Moscow state university of М. В. Lomonosov biological faculty, department of vertebrate zoology
Russia, 119992 Moscow, Lenin mountains, 1/12
e-mail: chaffinch@bk.ru
beme@nm.ru
There are such characteristics of a matter of the Nature, as variability and stability (tradition, norm). Probably, these opposite processes as if other qualities of different forms of energy also create "movement", development. Thus the given properties of a matter of the Nature can be considered at different levels of its organization. In this article we consider and we compare such biological essences as if chromosomes of cells of an organism and species-specific songs of birds. It appears that for these forms of energy of the Nature also variability and the certain norm are characteristic. Variability (Variation) - property of alive organisms and their displays to exist in various forms (phenotypic variability). That is found out differences of the certain attributes or properties of an organism are made hereditary (genetic) signs and environment variability. Genetic variability of organisms is caused by distinctions in a set of genes, but environment variability (or modification) are defined (determined) by reaction of the given organisms (with the identical set of genes) to influence of external conditions. Limits (scope) of environment variability are defined (determined) by norm of reaction (Biological dictionary, 1986).
Norm of reaction of an attribute (of a genotype) - in limits of which the phenotype can change without change of a genotype. The limit of norm of reaction, its "scope" at individuals of any species, depends on a level of the organization of an organism, and from features of an investigated attribute, and also depends on a genotype of an individual. It is usually forms of plural external (phenotype) display of one genotype - polymorphism (different shades of covers, different minor forms of a beak, a skeleton, bodies and specializations of cells, different variants or types of the general (of the common) species-specific songs in a population of birds) (the Biological dictionary, 1986).
Cultural (vocal, song, signal) tradition (Cultural norms or traditions) - vocal (song, signal) norms in structure and a pronunciation of vocal patterns of the majority of individuals of birds in the certain population of a species and establishment from year to year (in the certain interval of time and in the certain territory are formed different vocal traditions of birds of the certain species, representing different parities (ratio) (%) of different types or variants of species-specific songs, vocalizations of a species as a whole) (Mundinger, 1982).
Types of vocalizations (songs, signal) birds (Types of vocalizations) - the variants of one species-specific vocal pattern different to each other: 1) in morphology (different sound of parts, phrases of songs - an accent on different components), 2) in phonetics of songs (sound of separate elements - comes to light in distinction of their forms, structures on sonogram), 3) in some cases in syntax (a set and an arrangement or combination of similar phrases, parts of songs) (Catchpole, 1995).
Thus, we can compare existence of a cellular (molecular) level of the organization of a matter of the Nature and a cultural (social) level (by the example of vocal realization of birds), especially marking their ability to form different combinations from former conditions, creating new forms of a matter of the Nature, but containing initial elements.
The song of chaffinch (Fringilla coelebs L.) has a genetic basis (species-specific features) (Thorpe, 1958; Nottebohm, 1967), but by training, at improvisation at singing, and also at mistakes of copying at vocal training, a variety of types of songs in a population constantly are replenished (Slater et al., 1979, 1984; Ince et al., 1980; Jellis, 1977). The variability of song types found out by us can testify to genetic heterogeneity of a population and microevolutionary processes occurring in it. At the analysis recorded songs of chaffinch on Curonian spit of Baltic sea (the Kaliningrad region), cases of formation of new song forms have been marked as a result of a combination of phrases of different types of songs. Combinative variability of species-specific songs of birds have found out in the researches (on sonograms) by many scientists-bioacoustics (for example, in titmouses - Paridae which combined songs from different desires or sounds), were observed the mixed types of vocalizations (Hailman, Ficken, 1996).
Various alleles of different genes form every possible combinations in separate genotypes. Combinative genes have been carried out at crossingover and an independent divergence of different pairs chromosomes during meyoz, and also in sexual process. So arises combinative variability of a species. Different variants of the phenotypes which have arisen on the basis of different combinations of genes, are exposed to natural selection. It is possible to tell, that the genofund of any species possesses the certain reserve of combinational variability constantly filled up with new mutations and being the necessary precondition for a survival of a species in the changing environment (from N. N. Jordanian, 2001).
Thus it is necessary to note, that different genetic and chromosomal mutations (sharp changes of structure of chromosomes, genes) are possible (probable): translocation - a mutual exchange of sites between two different, not homology each other, chromosomes; deletion - loss by a chromosome of its any site; inversion - turn inside a chromosome of its any site on 1800; duplication - doubling of those or other sites of a chromosome; transposition - rearrangement of genes inside a chromosome and others. Similar forms of "mutations" can arise in song patterns of birds at vocal realizations - an exchange of parts between different types of species-specific songs ("translocation"), removal of the certain part from song pattern ("deletion") and formation of new types of songs is possible, doubling the same part in the certain types of songs at singing birds ("duplication"), rearrangement of an arrangement of parts in song pattern of a bird ("transposition") is possible (Astakhova, Byome, 2009б).
The cultural mutation (a vocal mutation - a "wrong", non-standard pronunciation of a vocal pattern) in populations of birds also can arise as a result of easing the control of occurrence of original non-standard vocal "phenotypes", but as display of the general (common) genotype, in populations or in the certain "communicative" groups of birds on the part of social "pressure" of the majority of individuals at vocal realization to each other by means of face-to-face singing, easing of restriction in a variety of vocal types of species.
1. Astakhova O. A., Byome I. R. 2009б. The analysis of combinations of phrases and various ways of singing of syllables at vocal variability of song types of chaffinch (Fringilla coelebs L.) // Successes of modern biology. In press.
2. The biological encyclopedic dictionary. 1986. Editor M. S. Giljarov; the Editorial board: A.A.Baev, G. G. Vinberg, G. A. Zavarzin, etc. - Moscow: Publishing house the Soviet encyclopedia, 831 p.
3. The Jordanian N. N. Evolution of a life: Studies grant for Institutions. - M.: the Publishing center "Academy", 2001. - 432 p.
4. Hailman J. P., Ficken M. S. 1996. Comparative analysis of vocal repertoires with reference to Chickadees. P. 136-159. In Ecology and evolution of acoustic communication in birds (Eds Kroodsma D., Miller E.). Cornell University Press, Comstock Publishing Associates. Ithaca, London. +
5. Ince S. A., Slater P. J. B., Weismann C. 1980. Changes with time in the song of a populations Chaffinches // Condor. № 82. 285-290.
6. Jellis R. 1977. Bird sounds and their meaning. Cambridge. 256 p. 7. Slater P. J., Ince S. A., Colgan P. W. 1980. Chaffinch song types: their frequencies in the population and distribution between repertoires of different individuals // Behaviour. № 75. P. 207-218
8. Thorpe W. H. 1958. The leaning of song patterns by birds, with especial reference to the song chaffinch Fringilla coelebs // Ibis. № 100. 535-570. +
9. Catchpole C. K., Slater P. J. B. 1995. Birdsong. Biological themes and variation. Cambridge: Cambridge University Press. 248 p.
10. Nottebohm F. 1967. The role of sensory feedback in development of avian vocalizations // Proc. Int. Ornithol. Cong. 14. Oxford-Еdinburg: Blackwell scient. Publs.. P. 265-280.
11. Mundinger P. C. 1982. Microgeographic and macrogeographic variation in acquired vocalizations of birds (Еds. Kroodsma D.E., Miller E.H.). Academic Press, New-York. P. 147-208.
Assumptions of evolution of chaffinch song (Fringilla coelebs L.)
Astakhova О. A.
Moscow state university of М. В. Lomonosov biological faculty, department of vertebrate zoology
Russia, 119992 Moscow, Lenin mountains, 1/12
e-mail: chaffinch@bk.ru
beme@nm.ru
Existence of any biological essence is usually accompanied by development or evolution ("movement" in time and space), by variability or by differentiation on distinguished set, but on similar forms in the base. These laws of biological substance have been revealed still by Ch. Darwin, E. Mayr. But whether it is possible to attribute (relate) singing of birds to similar biological essence as if, for example, coloring of covers, the form of a body, features of a structure of a skeleton, a digestive path, bodies of breath of different organisms, to their genetic and physiological characteristics? Singing of birds (Aves) is usually represented as if display of work of nervous system, mentality (its rudiments at birds), as if display of "spiritual" forces of an organism. It appears, the song of birds is capable to be differentiated in the different remote territories, is hereditary to be transferred in the features (in full or in part at different species), is capable to make set of distinguished forms (or different types, but similar in the base) that can speak about their the general (common) initial ancestor vocal form which as if a result of evolution and differentiation is original for each vocal line of different species and larger groups of birds.
In many references on bioacoustics of birds the point of view about formation of original vocal traditions of birds in different local and geographical populations is described - there is a distinction between sets of different types of species-specific song in the geographical populations remote from each other. Even if types species-specific song of chaffinch (Fringilla coelebs L.) can differ inside a geographical population, but they are in aggregate capable to reflect in the structure any features of a climate, of character of vegetation of district - and, thus, to form special local vocal (song) culture (ways of singing species-specific song), which distinguish 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 generation to next generation can develop (to be formed) different vocal (song) traditions, probably, giving rise to new forms (norms) of species-specific song or to distinguished vocalizations of new species of birds.
Singing of chaffinch (Fringilla coelebs L.) - the feature (sign) is paralleled 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 base 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). It is possible, more frequently evolution occurs in time, instead 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 lines, 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.
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 ways (special manners or patterns of acts) and can quickly "forget", change former (previous) course. 2
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Olesya Astakhova
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