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MODELLING OF THE SEASONAL DYNAMICS OF BIRD MIGRATION
Abstract and Figures
Busse P. 1996.Modelling ofthe seasonal dynamzes ofthe bzrd mzgratzon. Ring, 18, 1-2: 97-119. The basic model as to strueture of the seasonal migration pattem of the bird species are: (I) the pattem is the normai distribution altcred by environmental faetors, and (2) it is composed of a number of waves of migrants cases of whieh are explained variously. Aceording to the first model thc pattem can be described by means of average datc and standard deviation or by median day and percentiles. The second model needs other methods of prcsentation of the seasonal dynamie s pattcm. The aim of this papcr is to diseuss the problem how to describe thc scasonal dynamics of the bird migration. Analyses of many ycar migration data suggest that the second model is right. Thc modeli ing of thc pattems as a row of waves of migrants is discussed theoretically. This is illustrated by an example ofthe seasonal migration pattem (1961-1990) of Willow Warblcr at two bird ringing stations located at the southem Baltic coast. It is concluded that: (I) the pattem ofthe passage is rcpeated year by ycar with quitc high accuracy being, however, altercd by random fluctuations of environmental variabies, (2) the averagc seasonal migration dynamics can bc described by pooling yearly dynamics data into the to tal com-posed distribution, (3) subsequent waves can bc reconstructed from thc total pattem as a row of quasi-normal distributions by means ofiterative modelIing proccdure, (4) obtained reconstruction ofthe contents ofthe sea-sonal dynamics pattem can be used for the estimation of relative frcquencies of birds belonging to subsequent waves. These results could be uscful for dceper monitoring studies and for comparisons between passagc pat-terns at diffcrcnt bird stations, (5) the wavc-likc structurc of thc seasonal migration pattcm cause that attempts of thc statistical dcseription of thc pattem by means of average day and the standard dcviation as wcll as by median day and percentilcs are biologically mcaning1ess. INTRODUCTlON Seasonal dynamics ofbird migration is rarely studied as a separate problem. Usually it is shown as a descriptive data being a background for discussing other problems, e.g. influence of weather factors on migration or shift in migration time over a large territory. To discuss weather influence on migration the authors stress occurrence of pronounced dai1y peaks ofthe species number as well as days when the species is not observed there. These peaks and minima of migration are assumed there as deviations from an "ideal" course of migration pattern which is silently assumed to be a normai distribution of num-bers of migrating birds. The same assumptions are made when shift in the time of mi gra-tion is studied. In that case, however, mentioned irregularities of migration pattern are frequently taken into consideration and statistical problems whether mean date
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... The air pressure changes seems to be direct stimulus for the starting decision of birds ready to move. From the other side, subsequent waves seems to be fixed to the "timewindows" during the season (e.g. for the Willow Warbler, Pylloscopus trochilus -Piotrkowska 1995, the Blackcap, Sylvia atricapilla - Busse 1996, Kopiec 1997, Kopiec--Mokwa 1999. ...
... Observation data on diurnal passage of birds were collected for many years (since 1961 to 1991) along the Polish Baltic coast at a few sites using the same field methods. The visual observations were carried out during work of the Operation Baltic ringing stations (Busse andKania 1970, Busse andHalastra 1972) parallely to the ringing of birds. Observations of passing birds were performed 15 minutes per every hour from sunrise to sunset. ...
... The main analysis of this work -description of the seasonal dynamics of migration -was carried out with modified method used for that kind of analysis in earlier publications by Busse and Kopiec (Busse 1996), Kopiec (1997) and Kopiec-Mokwa (1999). For all years separately there were calculated percent values of daily numbers. ...
The Wood Pigeon, Columba palumbus , is a common diurnal migrant across most of the Europe. The visual observations were carried out during autumn work of the Operation Baltic ringing stations, mainly at Mierzeja Wiślana (54°21’N, 19°19’E) parallely to the ringing of birds. Observations of passing birds were performed 15 minutes per every hour from sunrise to sunset. The autumn observation period was long enough to cover whole pigeons migration at the southern coast of Baltic (14 Aug. – 11 Nov.). Altogether 119,019 Wood Pigeons were observed within 20 seasons of observations. The goals of this work is (1) to describe the course of the Wood Pigeon autumn migration at the Polish Baltic coast as to numbers in long-term aspect and the seasonality of migration, (2) to propose further development in the study on wave structure of the bird passage. The main analysis of this work-description of the seasonal dynamics of migration - was carried out with modified and widened method used for that kind of analysis in earlier publications. In the present paper the wave structure is described by the estimated border days between supposed waves taking under consideration different properties of the passage dynamics. Here are used three groups of parameters: (1) total numbers of birds observed daily during several years, inter-year variability (as SD between year number values), and the coefficient of variation ( V ), (2) number of migration peaks that occurred in that date within years of the study (there were used two classes of peaks – High Peaks and Peaks ), (3) numbers of birds passing during recognized peak days. All estimations were done independently of others and only finally summarized using common ranking scheme. Such step by step procedure make estimation more and more reliable. Very high agreement between estimations based on three groups of parameters suggests strongly that the proposed estimation procedure is effective enough.
... W analizie sezonowej dynamiki przelotu poszczególnych gatunków zastosowano podejście zgodne z koncepcją falowości wędrówki (Busse 1996), według której przelot tworzą następujące po sobie fale wędrujących ptaków (wyróżniające się jako maksima), rozdzielone okresami mniejszego nasilenia wędrówki (minima). Przedstawiono relatywną liczbę osobników rozpatrywanego gatunku odłowionych danego dnia (%d t ) w całkowitej liczbie osobników danego gatunku schwytanych w danym sezonie. ...
... Koncepcja falowości wędrówki zakłada, że pod wpływem zaistnienia pewnych czynników (wewnętrznych i/lub zewnętrznych), ptaki migrujące w zsynchronizowany sposób rozpoczynają wędrówkę (lub jej konkretny etap) w danym miejscu i czasie, przez co tworzą one swoistą "falę wędrówkową" (Dolnik & Blyumental 1967, Busse 1996. Tego rodzaju synchronizację wędrówki udokumentowano m.in. ...
... Koniunkcja sprzyjających warunków pogodowych (i innych czynników egzogennych, np. zachowań socjalnych) oraz fizjologicznej (w tym energetycznej) gotowości migracyjnej wielu osobników skutkować mogłaby synchronizacją decyzji o podjęciu wędrówki (Dolnik & Blyumental 1967, Ketterson & Nolan 1985, Weber et al. 1998, Zehnder et al. 2001) i w konsekwencji utworzeniem się fali ptaków migrujących w zbliżonym czasie i przez konkretny obszar (Dolnik & Blyumental 1967, Busse 1996, Remisiewicz & Baumanis 1996. ...
Ornis Polonica 57, 2016: 39–57.
Autumn migration of passerines was studied basing on daily catches at the Rakutowskie Lake (central Poland) in 2010-2012. Among captured birds, the most abundant were long-distance migrants, which formed main migration peak. Less numerous partial and short-distance migrants created the second, later peak. The most abundant species showed either high (e.g. Reed Warbler Acrocephalus scirpaceus and Sedge Warbler A. schoenobaenus) or low (e.g. Barn Swallow Hirundo rustica and Long-tailed Tit Aegithalos caudatus) inter-seasonal migration dynamics repeatability. Among species representing three main migration strategies: long-distance, short-distance and partial migrants, we observed a distinctive similarity in migration curve shapes. This was manifested by the convergent course of dynamics curves and similar timing of migration waves, when comparing many pairs of species. In Reed and Sedge Warblers, which were characterized by the highest repeatability of migration, we found higher similarity of migration curves between the species in one season than within species in two consecutive seasons. This suggests mainly exogenous (e.g. weather-related) rather than endogenous control of migration timing. Key words: migration strategy, migration waves, bird ringing, bird banding.
... where: X y -number of birds in year y, X oy -moving average for year y, N -number of study years, M -total average. The moving weighted-average smoothed numbers for five following years (Busse 1973) and was calculated according to the formula by Busse (1996) ...
... The waves of migration were separated by dates on which the numbers of birds caught were clearly lower than on the adjacent days (the passage minima). A detailed description of dividing migration dynamics into waves is given in papers by Busse (1996) and Kopiec--Mokwa (1999). In the next stage, the dynamics curve was three times smoothed by the moving weighted-average, as in the case of multi-year dynamics. ...
Autumn Migration Dynamics and Biometrical Differentiation of the Dunnock (Prunella modularis) Passing the Southern Baltic Coast
The aim of this study is to determine biometrical differentiation among Dunnocks caught at the two ringing sites (Bukowo-Kopań and Mierzeja Wiślana) located on the southern Baltic coast. The distance between those two stations covers 190 km. The material was collected during autumn fieldwork of the Operation Baltic in 1961-2003. The material used for biometrical analysis comprises only immature birds from the period of the most intensive migration, when the numbers of caught individuals allowed to compare the results for both stations. The seasonal dynamics at both sites was pooled for 43 years of catching. Medians of autumn migration for the stations were significantly different. A shift of the median for the eastern site (Mierzeja Wiślana) by 6 days after the median for the western site (Bukowo-Kopań) suggested different origins of birds migrating through the stations. The analysis of standard deviations for the studied biometrical parameters confirms an intra-seasonal change in proportions of birds probably originating from different areas in Europe.
... This is very well confirmed by the long-term data (since 1961) of Operation Baltic in Poland, e.g. ten years of data for many species ringed/observed -Busse and Halastra 1981; thirty years of data for the Willow Warbler Pylloscopus trochilus -Piotrkowska 1995 and for the Blackcap Sylvia atricapilla -Busse 1996, Kopiec 1997, Kopiec-Mokwa 1999 and fourteen years of data for the Robin Erithacus rubecula -Nowakowski et al. 2005, as well as data from other areas, e.g. a nineteen-year study on many species from Hungary -Gyuracz et al. 2017, or a ten-year study of many species from Palestine - Awad et al. 2017. There are many, many examples of multi-wave distributions. ...
Cavina E., Bucchi R. and Busse P. 2018. The general pattern of seasonal dynamics of the autumn migration of the Wood Pigeon Columba palumbus in Italy . Ring 40: 3-18.
Given the scarcity of studies on the migration of the Wood Pigeon through Italy, the first systematic observations by a network of hunters, as citizen researchers, can be presented as a starting point for more in-depth analyses. Observations from the years 1998-2006 are analysed and presented in a generalized form. During this period more than 100 observation sites, covering most of Italy, were active for about 40 days every autumn. Migration over Italy was described in terms of the timing and intensity of migration. Special attention was directed to the long-term number dynamics and seasonal dynamics of the passage. The most intensive migration was observed within northern Italy, while lower intensity is visible more to the south of the peninsula. Following tendencies in numbers of observed migrants within the ten years of the study, we can find positive tendencies in most of the northern provinces, while three negative trends are visible in central Italy. The study of the seasonal pattern, in terms of the number dynamics of the passage and the frequency of pronounced peak days, strongly suggests that there are five or six waves of pigeons passing through Italy in different parts of the autumn that are quite stable between years. Every year the time of the passage includes a few peak days of migration.
... This is a very rough division of Robins into few groups used in this exemplary analysis and is not equivalent to a real wave division, similar to those used in other biometrical analyses (e.g. Busse 1972, Busse and Maksalon 1978, Maksalon 1983 or analyses of seasonal migration dynamics (Busse 1996, Remisiewicz and Baumanis 1996, Kopiec 1997. Figures II-72 and II-73 show how well pronounced the differences in wing/tail distributions are if more compact groups of migrants are analysed. ...
Foreword: The first bird station I saw in my life was the Bradwell Bird Observatory, Great Britain. I vis-
ited it invited by Bob Spencer (who He was I do not need to explain even to younger ringers) in
autumn 1959. Two days of the bird catching and 173 birds caught. For young ringer who has never
ringed one full-grown bird, but pulli only, that experience was great.
In 1960 first Polish bird station started pioneering bird catching – a group of students from the
University of Warsaw started to learn bird migration at the Polish Baltic coast. Our minds were
fresh, we knew nothing about bird migration but we were enthusiastic – after successful catching
in 1960 (one thousand of ringed birds), we started a project of setting up several bird ringing sta-
tions along the Polish Baltic coast, called later the „Operation Baltic”. Because the bird migration
is a huge and very complicated phenomenon, thus the main idea was „a NETWORK working ac-
cording to the standard methods”. Since that time the Operation Baltic has been working for 40
years and a number of birds ringed within the project exceeded well 1 200 000. Most of them were
not only ringed but measured, fat scored and weighed. Other data were collected as well. Large
number of birds forced development of working routine in such a manner that we are able to cope
with huge rushes of birds and safely handle them during collecting a lot of data. These data are the
main objective of the fieldwork – bird ringing must make science and not only fun.
Visiting many bird stations, from Britain and Italy to Russia and from Finland and Sweden to
Israel, I found that the main problem of international co-operation is incompatibility of methods
and local routines of work, which make exchange and efficient use of data files very difficult. And
once more the idea of „a NETWORK working according to the standard methods” seems to be
a proper solution. Such trials were made within „ESF European-African Songbird Migration Net-
work” working three years on the SW bird migration flyway in the mid 90ties and, finally, Manual
of Field Methods was published. In 1996 a group of ornithologists from northern and central
Europe established the SEEN („SE European Bird Migration Network”) that focuses on the SE fly-
way that has been poorly studied yet. Necessity of common methodical programme induced prepa-
ration of comprehensive manual that could give not only methods of the fieldwork, but also some
methods of evaluation of collected material.
I am deeply grateful to all people who contributed to this manual and discussed its contents. I
would be very happy to hear from the Readers what was useful for you and what should be added
or changed in a future development of the Bird Station Manual.
I hope that the Manual will be useful not only for bird stations but for many ringers who ring
the birds not only for fun, but whose ambition is to contribute to Science as well. Obviously, not all
parts of the manual will be applicable in an individual ringing, but always it is worth to think
a while when we encounter new, sometimes apparently strange ideas – especially when we plan to
start a new project
... Równie¿ obserwowana zmiennoae dynamiki sezonowej przestaje byae przeszkod¹ przy analizie danych z bardzo d³ugich okresów czasu to, co w skali pojedynczych sezonów wydaje siê byae dyskusyjne i niepewne (np. falowoae przelotu) na takim materiale jawi siê jako w znacznym stopniu prawdopodobne i uzasadnione (KANIA 1981, BUSSE 1996, KOPIEC 1997, KOPIEC-MOKWA 1999. Tak¿e problemy metodyczne przy opracowaniu danych biometrycznych, tak istotne przy wykorzystaniu sk¹pych ilociowo materia³ów, staj¹ siê mniej dokuczliwe, gdy dysponujemy danymi obfitymi i zbieranymi w ró¿nych latach. ...
The Polish bird migration research began in 1931, when within Govern-mental Zoological Museum the Bird Migration Research Station was launched as the Polish Bird Ringing Centre. After the IInd World War the Centre continued its work as Ornithological Station, at the beginning in Warsaw and then in Gdañsk. Till 1960 bird ringing in Poland concentrated on nestlings and only later, with organizing a research programm, called Operation Baltic, a heavy point shifted to ringing of migratory birds. The programm introduced a revolutionary standard to bird migra-tion studies worldwide: a long-term studies running under strictly standardized methodology and within the simultaneously working network of bird stations. The idea was further developed when the Operation Baltic initiated in 1996 establishing the SEEN (SE European Bird Migration Network) the international network for the bird migration studies covering several countries from Northern and Central Europe to Syberia in the east and Middle East in the south. The Operation Baltic introduced several original methods both field ones and data elaboration procedures. Among the field methods some should be mentioned: tail and wing-formula measure-ments, fat scoring on live birds and a flat orientation cage for studies of directional preferences of nocturnal migrants. Elaboration methods are adapted to use of vario-us data collected and being collected by the Operation Baltic and the SEEN. They cover, among others, evaluation of wing-formula measurements using wing-shape indices, standardization of body mass for a fat load, analyses of the biometrical data, with location of breeding grounds of migrants, as well as statistical analysis of the orientation cage data. Since the beginning of work of the Operation Baltic complexity of the study methods and large-scale, continental size migration patterns were taken under consideration as the basis of the research. An unique, in the World scale, database on bird migration, running system of collecting of the field data and a wide
... For each day the percent value was calculated in relation to the average daily number of Robins caught (immatures and adults jointly) in the analysed season at a given station and then used to draw the migra- tion dynamics. The obtained dynamics were divided into migration waves according to modelling procedure proposed by Busse (1996). This method is based on cutting the source data for each season according to the minima of the dynamics. ...
Variation in the Extent of Greater Wing Coverts Moult in Robins ( Erithacus rubecula ) Migranting in Autumn Through the Polish Baltic Coast
Intra-seasonal variation in the number of unmoulted coverts in immature Robins caught during 2001-2003 autumn migrations was studied at two ringing stations (Mierzeja Wiślana and Bukowo-Kopań) located at the Polish Baltic coast. To determine the number of unmoulted greater wing coverts we counted immature-type coverts with light spots at tips. In the analyses data on the number of spotted coverts from ca 11 000 migrating individuals were used. We analysed the number of spotted coverts in each season and at each ringing station separately. Based on migration dynamics we distinguished migration waves and the number of spotted coverts were compared among the waves. Trends of seasonal changes in mean values of this parameter were assessed. To explain these tendencies, percentage distributions of wing spots for each wave were presented, with distinguished three categories: 0-3 (low), 4-5 (medium) and 6-8 (high) number of spotted coverts. Next, we compared distribution of this parameter among waves by Kruskal-Wallis and post-hoc Dunn's tests.
Late waves generally differed in the distributions of spotted coverts' number from the earlier ones in all seasons and at all stations. Our results showed the same tendencies within a season in all cases: the mean number of unmoulted coverts fluctuated in September, but starting from the end of this month and in October the trend was clearly increasing. This was due to changes in frequencies of Robins assigned to the distinguished categories - in September birds with medium number of spotted coverts constituted over 50% of all migrants, while in late September and/or October waves individuals with high number of spotted coverts predominated. Both in 2002 and 2003, the moment of the shift in this domination from birds with medium to those with high number of unmoulted coverts was synchronised between the two stations.
These intra-seasonal differences in moult advancement can be explained by two overlapping phenomena - subsequent migration over the Baltic coast of populations with different moult characteristics and by less advanced moult of birds from later broods. Correspondence of the observed trends in moult advancement with literature data on migration timing of Robins of different breeding origin and winter quarters indicates that the populational differences play an important role in the observed variation.
Busse P. 2018. Estimation of local heading patterns of nocturnal Migrants using orientation cages. Ring 40: 31-58
The main aim of this paper is to compare the results of two data evaluation procedures
used for presenting the data from the orientation cage field tests. Both procedures accept
multimodality of the data and multimodality of the headings of an individual bird as well
as migrating population. The goal is to reach acceptable level of migration patterns pres-
entation in biological sense, taking under consideration a flexibility of the real movements,
depending on specific weather and landscape parameters. Such knowledge is absolutely
necessary for estimating migration bottle-necks and the long-term studies on influence of
the climate changes on migration patterns. The material used for the comparison of the
procedures was collected in years 2001-2007 by the team of the Bulgarian Ringing Station
Kalimok (44 o 00’N, 26 o 26’E) within the frame of the SEEN (SE European Bird Migration
Network) activity and kindly shared for evaluation. The data were obtained using the stan-
dard SEEN methods, with the standard Busse’s cage working procedure of the field tests.
The material contains data on four species of nocturnal migrants living in different habi-
tats: the Great Reed Warbler, Acrocephalus arundinaceus (ACR.ARU), the Sedge Warbler,
A. schoenobaenus (ACR.ENO), the Willow Warbler, Phylloscopus trochilus (PHY.LUS) and
the Whitethroat, Sylvia communis (SYL.COM). There are confirmed earlier conclusions
that so called „classic” unimodal procedure is not applicable to the orientation cage data
resulted from any field procedure. There are available two evaluation procedures that base
on the same general assumptions: multimodality of distributions that reflects combination
of several unimodal partial distributions, that can be described both using sophisticated
Bayesian „Calculation” method and much simpler „Estimation” procedure. Results of both
procedures are enough close to each other that they can be used for describing local and
general heading patterns of migration of the nocturnal migratory movements studied us-
ing orientation cages.
Biometrics Variation and Directional Preferences of Immature Robins ( Erithacus rubecula ) Caught in Northern Italy during Autumn Migration in 2005
Inter-seasonal changes of biometry and preferred migration directions of Robins were studied according to data collected during autumn migration in northern Italy at the Arosio Bird Observatory (45°43'N, 9°12'E). Altogether 598 immature Robins were caught and 187 orientation tests were performed. Wing, tail and tarsus length, wing shape and weight were analysed in subsequent five migration waves distinguished according to migration dynamics. General pattern of migration as well as graphs with distribution of preferred directions in subsequent waves were prepared. In the case of tail length and weight their average values in subsequent waves were significantly different. Decrease of wing length was noted along the season. On the contrary, increasing trend was observed in the case of tail length and wing shape. Results of orientation tests showed that SSE direction was predominant (34%). SW direction was not clearly marked and its percentage was 23%. Distribution of directions slightly changed in subsequent migration waves. Noted results suggest passage of Robins heading to the Mediterranean basin and Apennine winter quarters. Obtained inter-seasonal changes of biometry and preferred directions can be an effect of differences in migration time between this groups or gradual inflow of more northern populations what the authors discuss here.
Extended English summary is on pp. 264-267 of the text
Previous papers on Song Thrush migration through Polish Baltic coast have shown that these birds are originated from wide territory of Fennoscandia, Baltic Republics and western part of Russia. They migrate toward four winter-quarters distributed in Europe from Atlantic coast up to the Balkans. This paper presents biometrical dataq collected during Operation Baltic work in the period 1961-1970 (7792 individuals measured) and preliminary analysis of results. Measurements were collected at 5 ringing stations by means of standard Operation Baltic methods. Since 1965 Song Thrushes were aged, but in this paper all individuals are treated together as biometrical age-depended differences are slight in this species (0.6-0.8% for wing-length, wing index l, weight and 2.6& for tail length). Very complicated pattern of biometrical differentiation found in the studied species suggest that classic methods of biometrical analysis based on simple looking for statistical differences between averages characterizing some bird groups are not sufficient and more advanced techniques should be developed.
Material collected by the method of visual observation and catching birds in nets during their spring migration on the Hel Peninsula in 1963-1967 and on Mierzeja Wślana (vistula Spit) in 1966-1967 is presented. The three main parts of the paper provide the author's methodological conceptions the working out material collected by above-mentioned methods, a description of migration dynamics, and a biometric analysis of the birds trapped. The degrees of generalisation based on such material are presented and a suitable system of symbols is proposed in the course of methodological considerations. The biometric part contains an analysis of the population composition of migrating birds. Population differentiations have been found between successive waves of migrants and between birds flying over the two observation points. The variability of biometric parameters of populations in the consecutive years is also discussed.
Analysis and comparison oj' the dynamics oj' autumn migration oj' Willow Warbler (Phylloscopus trachitus) at Bukowo, Hel and Vistula Spito Diploma work
- L Piotrkowska
Piotrkowska L. 1995. Analysis and comparison oj' the dynamics oj' autumn migration oj' Willow Warbler
(Phylloscopus trachitus) at Bukowo, Hel and Vistula Spito Diploma work. Bird Migration Research Station. Univ. ofGdansk.
Same aspects oj'Song Thrush migration at the Polish Ballic coast
- P Busse
- L Maksalon
Busse P., Maksalon L. 1978. Same aspects oj'Song Thrush migration at the Polish Ballic coast. Not. Om. 19,
1-4: 1-14.
Autumn migration oj' Song Thrush thraugh
- L Maksalon
Maksalon L. 1983. Autumn migration oj' Song Thrush thraugh Polish Baltic coast. Not. Om. 24: 3-29.