DYNAMICS AND SPATIAL DISTRIBUTION OF …chernelmte.extra.hu/Tomord98-07.pdfDYNAMICS AND SPATIAL...

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DYNAMICS AND SPATIAL DISTRIBUTION OF MIGRATORY BIRDS Results of Bird Ringing Project of Tömörd Bird Observatory 1998-2007 by JÓZSEF GYURÁCZ PÉTER BÁNHIDI University of West Hungary István Chernel Ornithological and Nature Conservation Society Szombathely 2008

Transcript of DYNAMICS AND SPATIAL DISTRIBUTION OF …chernelmte.extra.hu/Tomord98-07.pdfDYNAMICS AND SPATIAL...

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DYNAMICS AND SPATIAL DISTRIBUTION OF MIGRATORY

BIRDS

Results of Bird Ringing Project of Tömörd Bird Observatory 1998-2007

by JÓZSEF GYURÁCZ

PÉTER BÁNHIDI

University of West Hungary István Chernel Ornithological and Nature Conservation Society

Szombathely

2008

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To the memory of JAKAB VÖNÖCZKY-SCHENK

Reviewed by:

CSABA NÉMETH

English reviewed by:

PÉTER MOLNÁR

ISBN 978-963-9871-17-5

Published by: Institute of Biology, Faculty of Sciences and Technics, Savaria Campus University of West Hungary

9701 Szombathely, Károlyi Gáspár tér 4., P. Box: 170. [email protected]; www.nyme.hu

Printed by: Balogh és Társa, Szombathely

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Contents

Introduction 7 Study area and methods 9 Study site and fieldwork 9 Data processing and statistical testing 11 Results 12 1. Teal (Anas crecca) 12 2. Garganey (Anas querqudula) 12 3. Marsh Harrier (Circus aeruginosus) 12 4. Hawk (Accipiter gentilis) 12 5. Sparrowhawk (Accipiter nisus) 12 6. Buzzard (Buteo buteo) 17 7. Kestrel (Falco tinnunculus) 17 8. Quail (Coturnix coturnix) 17 9. Pheasant (Phasianus colchicus) 17 10. Water Rail (Rallus aquaticus) 17 11. Spotted Crake (Porzana porzana) 18 12. Moorhen (Gallinula chloropus) 18 13. Jack Snipe (Lymnocryptes minimus) 18 14. Snipe (Gallinago gallinago) 18 15. Woodcock (Scolopax rusticola) 18 16. Green Sandpiper (Tringa ochropus) 18 17. Wood Sandpiper (Tringa glareola) 17 18. Turtle Dove (Streptopelia turtur) 19 19. Long-eared Owl (Asio otus) 19 20. Short-eared Owl (Asio flammeus) 19 21. Nightjar (Caprimulgus europaeus) 21 22. Swift (Apus apus) 20 23. Kingfisher (Alcedo atthis) 20 24. Bee-eater (Merops apiaster) 20 25. Hoopoe (Upopa epops) 20 26. Wryneck (Jynx toruilla) 20 27. Grey-headed Woodpecker (Picus canus) 22 28. Green Woodpecker (Picus viridis) 22 29. Black Woodpecker (Dryocopus martius) 22 30. Great Spotted Woodpecker (Dendrocopos major) 22 31. Syrian Woodpecker (Dendrocopos syriacus) 22 32. Middle Spotted Woodpecker (Dendrocopos medius) 22

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33. Lesser Spotted Woodpecker (Dendrocopos minor) 23 34. Skylark (Alauda arvensis) 23 35. Sand Martin (Riparia riparia) 23 36. Swallow (Hirundo rustica) 23 37. House Martin (Delichon urbica) 25 38. Tree Pipit (Anthus trivialis) 26 39. Meadow Pipit (Anthus pratensis) 27 40. Water Pipit (Anthus spinoletta) 28 41. Yellow Wagtail (Motacilla flava) 28 42. Grey Wagtail (Motacilla cinerea) 28 43. White Wagtail (Motacilla alba) 28 44. Wren (Troglodytes troglodytes) 28 45. Dunnock (Prunella modularis) 32 46. Robin (Erithacus rubecula) 34 47. Thrush Nightingale (Luscinia luscinia) 37 48. Nightingale (Luscinia megarhynchos) 38 49. Bluethroat (Luscinia svecica) 38 50. Black Redstart (Phoenicurus ochruros) 38 51. Redstart (Phoenicurus phoenicurus) 39 52. Whinchat (Saxicola rubetra) 41 53. Stonechat (Saxicola torquata) 42 54. Wheatear (Oenanthe oenanthe) 44 55. Blackbird (Turdus merula) 44 56. Fieldfare (Turdus pilaris) 48 57. Song Thrush (Turdus philomelos) 48 58. Redwing (Turdus iliacus) 51 59. Mistle Thrush (Turdus viscivorus) 53 60. Grasshopper Warbler (Locustella naevia) 53 61. River Warbler (Locustella fluviatilis) 55 62. Savi’s Warbler (Locustella luscinioides) 55 63. Moustached Warbler (Acrocephalus melanopogon) 56 64. Sedge Warbler (Acrocephalus schoenobaenus) 56 65. Marsh Warbler (Acrocephalus palustris) 58 66. Reed Warbler (Acrocephalus scirpaceus) 60 67. Great Reed Warbler (Acrocephalus arundinaceus) 62 68. Icterine Warbler (Hippolais icterina) 62 69. Barred Warbler (Sylvia nisoria) 64 70. Lesser Whitethroat (Sylvia curruca) 64 71. Whitethroat (Sylvia communis) 67 72. Garden Warbler (Sylvia borin) 69

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73. Blackcap (Sylvia atricapilla) 71 74. Pallas’s Warbler (Phylloscopus proregulus) 75 75. Yellow-browed Warbler (Phylloscopus inornatus) 75 76. Wood Warbler (Phylloscopous sibilatrix) 75 77. Chiffchaff (Phylloscopus collybita) 77 78. Willow Warbler (Phylloscopus trochilus) 80 79. Goldcrest (Regulus regulus) 82 80. Firecrest (Regulus ignicapillus) 85 81. Spotted Flycatcher (Muscicapa striata) 87 82. Red-breasted Flycatcher (Ficedula parva) 89 83. Collared Flycatcher (Ficedula albicollis) 90 84. Pied Flycatcher (Ficedula hypoleuca) 90 85. Long-tailed Tit (Aegithalos caudatus) 92 86. Marsh Tit (Parus palustris) 94 87. Willow Tit (Parus montanus) 96 88. Crested Tit (Parus cristatus) 96 89. Coal Tit (Parus ater) 96 90. Blue Tit (Parus caeruleus) 97 91. Great Tit (Parus major) 100 92. Nuthatch (Sitta europaea) 104 93. Treecreeper (Certhia familiaris) 105 94. Short-toed Treecreeper (Certhia brachydactyla) 107 95. Penduline Tit (Remis pendulinus) 109 96. Golden Oriole (Oriolus oriolus) 110 97. Red-backed Shrike (Lanius collurio) 110 98. Great Grey Shrike (Lanius excubitor) 113 99. Jay (Garrulus glandarius) 113 100. Starling (Sturnus vulgaris) 114 101. House Sparrow (Passer domesticus) 114 102. Tree Sparrow (Passer montanus) 114 103. Chaffinch (Fringilla coelebs) 116 104. Brambling (Fringilla montifringilla) 118 105. Serin (Serinus serinus) 120 106. Greenfinch (Carduelis chloris) 120 107. Goldfinch (Carduelis carduelis) 122 108. Siskin (Carduelis spinus) 122 109. Linnet (Carduelis cannabina) 123 110. Redpoll (Carduelis flammea) 123 111. Crossbill (Loxia curvirostra) 123 112. Scarlet Rosefinch (Carpodacus erytrinus) 123

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113. Bullfinch (Pyrrhula pyrrhula) 123 114. Hawfinch (Coccothraustes coccothraustes) 125 115. Yellowhammer (Emberiza citrinella) 127 116. Ortolan Bunting (Emberiza hortulana) 129 117. Reed Bunting (Emberiza schoeniclus) 129 118. Corn Bunting (Emberiza calandra) 131 Summary and discussion 133 Acknowledgements 137 References 138 Appendix 139

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Introduction Bird migration is one of the most exciting phenomena of our living world, and therefore it is not surprising that it has been the subject of investigation since ancient times. Each year an estimated 50,000 million birds travel somewhere in the world. Short-distance migrants may just leave their mountainous breeding areas to seek milder winter conditions in the nearby foothills whereas long-distance migrants often cover tens of thousands of kilometres (ALERSTAM 1990). The principal migratory routes for many species of passerines migrating from the Europe to African wintering sites are fairly well known. What is missing is a detailed analysis of environmental factors controlling bird migration as well as the temporal and spatial course of the migratory journeys (BAIRLEIN 1997, TIELEMAN 2007).

Palearctic-African migratory bird populations have to defeat several ecological barriers, like seas and deserts during the migration in post-breeding season and spring. During the migration driven natural selection population-specific strategies (e.g. habitat selection, food preference, fat deposition, migratory direction, migratory speed and dynamic, intra- and inter-specific competition, moult timing, etc.) and control mechanisms (e.g. photoperiodicity of hormonal system, nervous system and orientation, etc.) have evolved. The migration dynamics involves the evolution of the number, time and duration of the stopover of the migratory population on a territory for feeding or resting (BERTHOLD 1993, RAPPOLE & JONES 2002).

Many species of European bird species are currently undergoing rapid declines, but some birds have increasing and expansive populations (BIRDLIFE 2004). The causes of these changes are not clear but factors like habitat fragmentation, agricultural technology, hunting and global climate change known to be seriously affecting certain European bird populations. Monitoring is essential for the identification of ecological and conservation problems affecting birds. According to the EC Birds Directive on the protection of European bird population, all European Community countries have an obligation to monitor bird population. In Annex V of the directive, specific reference is made to use bird ringing for the monitoring of population levels of migratory bird species. Birds are particularly good subjects for population monitoring because they are sensitive indicators of habitat changes. Annual counts of birds tell us how numbers are changing, but bird ringing is needed if we want to understand the mechanism of the observed changes.

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Bird ringing for scientific purposes started in Hungary in 1908, when JAKAB VÖNÖCZKY-SCHENK released White Storks that were ringed with metal rings engraved with numbers. Bird ringing and bird migration studies in particular were revolutionized in the mid-20. century when mist-nets first became generally available. Mist-nets placed besides suitable vegetation readily capture passerines like warblers in their mesh. The BirdLife Hungary started a project called „Actio Hungarica” (AH) for investigation of bird migration in 1974 (SZENTENDREY et al. 1979). Actio Hungarica has been involved in the netting of migrating songbirds at a network of bird ringing camps in Hungary. The netting effort was strictly standardized and the habitat was not allowed to change, so changes in the numbers of birds captured should reflect changes in the migrating populations. The Tömörd Bird Observatory was founded in 1998 and it joined to the Actio Hungarica and the South-East Bird Migration Network (SEEN) that focuses SE flyway that has been poorly studied yet (BUSSE 2000). Different types of useful information can be recorded when birds are caught for the purpose of ringing. These include the age and sex of the bird, a variety of measurements which can be used to characterize different populations, the amount of fat stored by migratory birds, the state of feather moult as well as the habitat in which the species was captured. Single birds can be recaptured by other ringers, resighted or recovered by members of the public in a variety of different ways: hit by cars, found dead, caught by cat, shot, etc. If a bird is subsequently recaptured by other ringers, resighted repeated measurements make the study of different aspects of the annual life cycle, such as body mass changes prior to migration or the seasonal progression of moult possible. The National Ringing Centres collect all the information provided by ringers and members of the public who find ringed birds. The Hungarian Ringing Centre of BirdLife Hungary is located in Budapest. When birds overfly national boundaries information needs to be exchanged between Ringing Centres and the European Union for Bird Ringing (EURING) ensures that this happens efficiently (KESTENHOLZ 2007). Bird ringing has been carried out in Hungary throughout whole twentieth century. As the result, the long-term recovery and biometrical data for birds became one of the most valuable sets of information available for any group of animals. Despite all these ringing activities, many problems still remain unsolved, moreover, many more new ones recently emerged. For instance, the current global climate change – the past decade has been the warmest for 1000 years – is confronting us with so

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many rapid changes in the phenomenology of bird migration that we can hardly keep up with recording and analyzing them (BERTHOLD et al. 2003). In this publication the autumn migration of birds captured in the Bird Ringing Project of Tömörd Bird Observatory between 1998 and 2007 is characterized. A total of 49619 individuals of the 118 studied species were marked. The main aspects of the analysis were:

1. To register annual change of the migrating population size and trends found in migratory birds.

2. To describe the progress in time (dynamics) of the post-breeding movement by analysis of daily capture-recapture data.

3. To describe the habitat use of the migratory population by analysis of sites of captures.

4. To reveal the origin, migration direction and wintering sites of the migratory populations of Tömörd area by evaluating recoveries.

Study area and methods

Study site and field work

The birds were captured and ringed at the Tömörd Bird

Observatory (47°21'23"N 16°40'04"E) (Photo 1) in the west of Hungary. Tömörd is a small village in Vas County. It is located 15 kilometres from Szombathely (Fig. 1).

Bird ringing took place during the autumn migration, from the end of July or early August to the end of October or early November (Table 1).

Fig. 1. Location of the Tömörd ringing site in Hungary (Figure is from the website of SEEN)

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Table 1. Ringing activity periods

Year Date 1998 10 Aug. – 20 Sep. and 16 – 31 Oct. 1999 22 Aug. – 31 Oct. 2000 20 Aug. – 19 Nov. 2001 5 Aug. – 18 Nov. 2002 28 Jul. – 10 Nov. 2003 27 Jul. – 9 Nov. 2004 1 Aug. – 9 Nov. 2005 1 Aug. – 6 Nov. 2006 29 Jul. – 11 Nov. 2007 30 Jul. – 11 Nov.

For catching the birds 29 numbered mist-nets (12 meter long and

2.5 meter high with 5 shelves and a mesh size of 16 mm) were used. The nets were positioned in four different habitat types around the ringing station (Photo 2). These were characterised below according to the SE European Bird Migration Network (BUSSE 200) (See below).

Forest (W.LB6N0.2EUPR.N.). The 6-9 m high trees and bushes made up a compact, dense edge vegetation. It had an ecoton community. Its characteristic plant species were: Quercus cerris, Prunus spinosa, Crataegus monogyna, Pyrus pyraster. There were 4 nets in this habitat (Photo 3).

Bushy (S.BH2N0.2PRCR.O.). The 2-3 m high bushes made up a compact, dense vegetation, which was dissected by small grass patches. Its characteristic plant species were: Prunus spinosa, Crataegus monogyna, Pyrus pyraster, Rosa canina. 11 nets were standing in this biotope (Photo 4).

Grassland with scrubs (S.BG2N9.1ROCA.O.). It made a transition between the wet biotopes of the swamp and the steppe communities that used to cover the croplands around. There were a few bushes in the grassland. Its characteristic plant species were: Calamagrostris epigeios, Cirsium arvense, Arrhenatherum elatius, Verbascum thapsus. There were two small patches of Sambucus ebuli. There were 8 nets in this biotope (Photo 5).

Marsh (S.TC1S0.0.R.). Characteristic plants were Glyceria maxima, Typha latifolia, Salix cinerea. It was dried out in 2000 and 2001.

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There was 0.5-1 ha open water in the other years. There was a line of 6 nets here (Photo 6).

Data processing and statistical analysis

All birds were ringed and aged according to SVENSSON (1994) and Actio Hungarica (SZENTENDREY et al. 1979). Biometric data were taken using the methods of Actio Hungarica. All birds were ringed, sexed and weighed to the nearest 0.1 g (using a spring balance). The wing and tail lengths were measured to the nearest 1 mm. The fat reserves were estimated visually according to BUSSE (2000) ranging from 0 (no fat) to 8 (bulging fat).

Linear regression analysis and Pearson’s correlations were used to determine trends in the number of yearly captures. The average daily captures of the years were compared using Kruskal-Wallis test. Migration curves and cumulative graphs for the age groups or sexes were calculated based on the average daily captures of the years smoothed by a 3-day moving average. Based on this information the peak migration period (when we captured the most birds in that season) and the month with the highest captures were identified. By using the minimum stopover time of the recaptured birds the migration dynamics curves were plotted. The beginning of migration was determined based the analysis of the fat score of the captured birds and the dynamics of daily captures. Birds on migration accumulate much larger fat reserves than in other seasons. This fact made it possible to distinguish residents from migrants, as well as the discrimination of the beginning of migration from the post breeding dispersal. In this study it was assumed that the beginning of migration was the day when the first passage of very fat birds i.e. in which the fat covered completely the intestines (fat score 3 or higher) was noted. The average date of the beginning of migration is marked with arrow ( ) on the migration curves. Linear regression analysis and Pearson’s correlations were used to determine the trend observed in the change of the beginning of migration. The median dates of the migration were calculated with the exclusion of, the number of birds caught before the day starting the migration. The end date of migration was considered as the day after which only single individuals were captured in few-day intervals. The date of the end of migration is marked with arrow ( ) on the migration curves. The distribution of the captures in the different biotope types was checked by χ2-test for the year with the most captures.

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Sites of ringing and sites of recovery were delineated on a map for all the birds ringed abroad or at other Hungarian ringing sites and recovered in Tömörd and also those ringed in Tömörd and recovered abroad or at other Hungarian ringing sites (Photo 6).

Results

ANSERIFORMES

Anatidae 1. Teal (Anas crecca)

One juvenile bird of this species was captured on 5 September 2006 in the marsh.

2. Garganey (Anas querquedula)

One juvenile bird of this species was captured on the 10th of August, 2004 in the marsh.

ACCIPITIFORMES

Accipitridae 3. Marsh Harrier (Circus aeruginosus) In total, 3 individuals of Marsh Harriers were marked between 1998 and 2007.One adult female of this species was captured on the 20th of August, 2000, one adult male and one juvenile female were captured on the 25th of August,. 2002. 4. Hawk (Accipiter gentilis)

In total, 4 individuals of Hawks were marked between 1998 and 2007. One bird of this species was captured on the 15th of August, 2001, one adult female was captured on the 28th of July, 2002, one juvenile bird was captured on the 22nd of September, 2007 and one juvenile male was captured on the 6th of October, 2007. 5. Sparrowhawk (Accipiter nisus) In total, 12 individuals of Sparrowhawks were marked between 1998 and 2007. There were not captures in 1998, 1999 and 2000. One bird

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Photo 1. The Tömörd Bird Observatory

Photo 2. A view of the study area. Photos by T. Markovics

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Photo 3. A net site in the forest

Photo 4. A net site in the bushy

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Photo 5. A net site in the grassland with scrub

Photo 6. A net site in the marsh

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Photo 7. Recovery of ringed birds. Yellow triangles: Birds ringed abroad or at other Hungarian ringing sites which located further than 5 km from Tömörd and recovered in Tömörd. Red circles: Birds ringed in Tömörd and recovered abroad or at other Hungarian ringing sites which located further than 5 km from Tömörd. The map was made by Zs. Karcza, Hungarian Ringing Centre

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was captured each year in 2001, 2002, 2004 or 2006, 2 birds were captured in 2007 and 3 birds were captured in 2003 and in 2005. 6. Buzzard (Buteo buteo) The Buzzard was very common species in the study area. In total, 11 individuals of Buzzards were marked between 1998 and 2007. There were no captures in 1998, 1999, 2004, 2005 and 2007. One bird was captured in each year of 2000, 2003 and 2006, 4 birds were captured in 2001 and in 2002.

FALCONIFROMES

Falconidae 7. Kestrel (Falco tinnunculus)

One bird of this species was captured on the 24th of August, 2000.

GALLIFORMES Phasianidae

8. Quail (Coturnix coturnix) In total, 31 individuals of Quails were marked between 1998 and 2007. The most birds were captured in 2003 (11 indiv.) and 2004 (10 indiv.). There were no captures in 2002 and 2007. The date of the last capture was 3 October. 9. Pheasant (Phasianus colchicus) In total, 6 individuals of Pheasants were marked between 1998 and 2007. One bird was captured in 2005 and in 2007. Two birds were captured in 2000 and in 2002.

GRUIFORMES Rallidae

10. Water Rail (Rallus aquaticus) In total, 7 individuals of Water Rails were marked between 1998 and 2007. Four birds were captured in 2004, 1 bird was captured in 2005 and 2 birds were captured in 2006. The date of the last capture was 2 October.

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11. Spotted Crake (Porzana porzana) In total, 13 individuals of Spotted Crakes were marked between 1998 and 2007. The most birds (10 indiv.) were captured in 2004. One bird was captured in 1998, in 1999 and in 2003. The date of the last capture was 15 September. 12. Moorhen (Gallinula chloropus) Two birds were captured in 1999 and in 2004. The dates of last captures were 19 September.

CHARADRIIFORMES Scolopacidae

13. Jack Snipe (Lymnocryptes minimus) One bird was captured on the 18th and on the 25th of October, 1999, also on the 2nd of October, 2003 and on the 22nd of October, 2006 in the marsh. 14. Snipe (Gallinago gallinago) In total, 6 individuals of Snipes were marked between 1998 and 2007. One bird was captured in 2003 and in 2005, 2 birds were captured in 2004 and in 2006. The date of the last capture was 22 October. 15. Woodcock (Scolopax rusticola)

One juvenile bird of this species was captured on the 5th of November, 2004 in the bushy. 16. Green Sandpiper (Tringa ochropus) In total, 10 individuals of Green Sandpipers were marked between 1998 and 2007. The most birds were captured in 2005, 2 individuals were on the 22nd of September and 3 individuals were on the 11th of October. One bird was captured in each year of 1998, 1999, 2004, 2006 and 2007.

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17. Wood Sandpiper (Tringa glareola) One bird was captured of this species in August, also in years 1998, 2003, 2006 and 2007.

COLUMBIFORMES

Columbidae

18. Turtle Dove (Streptopelia turtur) In total, 9 individuals of Turtle Doves were marked between 1998 and 2007. The most birds (4 indiv.) were captured in 1998, 3 birds were captured in 2005 and 1 bird was captured both in 2002 and in 2007.

STRIGIFORMES

Strigidae

19. Long-eared Owl (Asio otus)

In total, 14 individuals of Long-eared Owls were marked between 1998 and 2007. The most birds (6 indiv.) were captured from August to early November in 2007. The date of the last capture was 4 November.

20. Short-eared Owl (Asio flammeus)

One juvenile bird of this species was captured on the 8th of September, 2004.

CAPRIMULGIFORMES

Caprimulgidae

21. Nightjar (Caprimulgus europaeus)

One juvenile bird of this species was captured on the 27th of August, 2001 in the bushy.

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APODIFORMES

Apodidae 22. Swift (Apus apus)

One bird of this species was captured on the 28th of August, 1998 in the marsh.

CORACIIFORMES Alcedinidae

23. Kingfisher (Alcedo atthis) One bird was captured on the 10th of August, 1998, 1 bird was captured on the 2nd of September, 2005 and 1 bird was captured on the 25th of September, 2007.

Meropidae

24. Bee-eater (Merops apiaster)

One bird of this species was captured on the 28th of August, 1998 in the marsh.

Upopidae 25. Hoopoe (Upopa epops) One bird of this species was captured on the 10th of September, 2006 in the grassland.

PICIFROMES Picidae

26. Wryneck (Jynx torquilla)

The Wryneck was accessory bird in the study area. In total, 114 individuals of Wrynecks were marked between 1998 and 2007. The most birds were captured in 2006, the least were in 1999. The average daily captures did not change significantly, Kruskal-Wallis test, H=14.21, p>0.05, but the yearly captures showed a significant increasing trend from 1998 to 2007 (Fig. 2). The migration started in the first half of August. The date of the last capture was 23 September. August was the month with the most captures (Fig. 3). 75 per cent of the Wrynecks were captured in the bushy in 2006, χ2=2, p>0.05 (Fig. 4).

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0

5

10

15

20

25

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 2. Yearly capture numbers of Wrynecks.

Linear regression of yearly captures, r=0.85, p<0.01

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 3. Average daily captures and migration dynamics

(smoothed moving average) of Wrynecks

Forest

Bushy

Marsh

Fig. 4. Capture rate (%) of Wrynecks in the four habitats, 2006

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27. Grey-headed Woodpecker (Picus canus)

One juvenile bird of this species was captured on the 14th of August, 2002 in the bushy. One adult female was captured on the 6th August, 2005 in the bushy. 28. Green Woodpecker (Picus viridis)

The Green Woodpecker was rare bird in the study area. In total, 24 individuals of Green Woodpeckers were marked between 1998 and 2007. The most birds (7 indiv., 6 in the bushy and 1 in the forest) were captured in 2007. There was no capture in 1998.

29. Black Woodpecker (Dryocopus martius)

One male bird was captured on the 18th of October, 2001 in the bushy. One juvenile male was captured on the 12th of August, 2004 in the bushy and one juvenile bird was captured on the 26th of October, 2004 in the grassland. 30. Great Spotted Woodpecker (Dendrocopos major) The Great Spotted Woodpecker was rare bird in the study area. In total, 93 individuals of Great Spotted Woodpeckers were marked between 1998 and 2007. The most birds (17 indiv.) were captured from July to November in 2007. From them 6 birds were captured in the bushy 6 birds were captured in the grassland and 5 birds were captured in the forest. 31. Syrian Woodpecker (Dendrocopos syriacus)

The Syrian Woodpecker was very rare bird in the study area. One bird was captured in 2001 and in 2006. 32. Middle Spotted Woodpecker (Dendrocopos medius)

The Middle Spotted Woodpecker was very rare bird in the study area. In total, 12 individuals of Middle Spotted Woodpeckers were marked between 1998 and 2007. The most birds (3 indiv.) were captured in 2000. There were no captures in 1998, 2001 and 2005.

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33. Lesser Spotted Woodpecker (Dendrocopos minor) The Lesser Spotted Woodpecker was rare bird in the study area. In total, 54 individuals of Lesser Spotted Woodpeckers were marked between 1998 and 2007. The most birds (12 indiv.) were captured from the early August to the end of October in 2004. From them 6 birds were captured in the grassland, 4 birds were captured in the bushy and 2 birds were captured in the forest.

PASSERIFORMES Alaudidae

34. Skylark (Alauda arvensis) The Skylark was rare songbird in the study area. One bird was captured on the 14th of August, 2004.

Hirundinidae

35. Sand Martin (Riparia riparia) The Sand Martin was rare songbird in the study area. In total, 13 individuals of Sand Martins were marked between 1998 and 2007. The most birds (6 indiv.) were captured in 2004. 36. Swallow (Hirundo rustica)

The Swallow was dominant songbird in the study area, but it was rare when there was small water in the lake or the lake dried out. In total, 1103 individuals of Swallows were marked between 1998 between 2007. The most birds were captured in 2002, the least in 1999. The average daily captures of the years changed significantly, Kruskal-Wallis test, H=20.57, p<0.01 (Fig. 5).

On average, migration of the Swallow started on the 9th of August (the earliest 31 July, the last 26 August). The average median date of the passage was 24 August. The average date of the end of migration was 2 October. The date of the last capture was also 2 October. The peak period of migration fell on the early August in the years of 1998-2007. August was the month with the most captures (Fig. 6). 96 per cent of the Swallows were captured in the marsh in 2002, χ2=6, p<0.05 (Fig. 7). The main migration direction is presumed SSW from Tömörd (Table 2).

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0

50

100

150

200

250

300

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 5. Yearly capture numbers of Swallows

0

5

10

15

20

25

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

No.

of b

irds

Fig. 6. Average daily captures and migration dynamics

(smoothed moving average) of Swallows

Forest

Bushy

Grassland

Marsh

Fig. 7. Capture rate (%) of Swallows in the four habitats, 2002

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Table 2. Data for all the Swallows ringed in Tömörd and recovered at the other Hungarian ringing sites or abroad

Capture Type Age/Sex Date Place Coordinates Text Distance

km Direction

degree Elapsed

days

Ringing adult female 24.08.2003. Tömörd

(AH) 47°21'23"N 16°40'04"E

1. Recovery adult 27.03.2005.

Potenza Picena,

Italy 43°23'N 13°41'E 500 208 581

Ringing juvenile 15.09.2005. Tömörd (AH)

47°21'23"N 16°40'04"E

2. Recovery adult

male 20.09.2006. Sumony (AH) 45°58'N 17°56'E 182 148 370

37. House Martin (Delichon urbica)

The House Martin was subdominant songbird in the study area, but it was rare when there was small water in the lake or the lake was dried out. In total, 598 individuals of House Martins were marked between 1998 and 2007. The most birds were captured in 2003, the least in 1999. The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=6.94, p>0.05 (Fig. 8).

On average, migration of the House Martin started on the 24th of August (the earliest 9 August, the last 13 September). The average median date of the passage was 5 September. The average date of the end of migration was 26 September.

020406080

100120140160180

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 8. Yearly capture numbers of House Martins

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The date of the last capture was also 26 September. The peak period of migration fell on the end of August in the years 1998-2007. The most captures occurred in August (Fig. 9). 97 per cent of the House Martins were captured in the marsh in 2002, χ2=6, p<0.05 (Fig. 10).

0123456789

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 9. Average daily captures and migration dynamics

(smoothed moving average) of House Martins

Forest

Bushy

Grassland

Marsh

Fig. 10. Capture rate (%) of House Martins in the four habitats, 2002

Motacillidae

38. Tree Pipit (Anthus trivialis) The Tree Pipit was subdominant songbird in the study area. In total, 656 individuals of Tree Pipits were marked between 1998 and 2007. The most birds were captured in 2002, the least in 2007. The average daily

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captures of the years changed significantly, Kruskal-Wallis test, H=77.41, p<0.0001 (Fig. 11).

0

50

100

150

200

250

300

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 11. Yearly capture numbers of Tree Pipits

On average, migration of the Tree Pipit started on the 26th of

August (the earliest 1 August, the last 13 September). The average median date of the passage was 4 September. The average date of the end of migration was 26 September. The date of the last capture was 17 October. The peak period of migration fell on the early September in the years 1998-2007. The most captures occurred in August (Fig. 12). 63 per cent of the Tree Pipits were captured in the grassland in 2002, χ2=0, p>0.05 (Fig. 13). Some Tree Pipits migrating above Tömörd originated from Finland (Table 3).

0123456789

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 12. Average daily captures and migration dynamics

(smoothed moving average) of Tree Pipits

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Forest

Bushy

Grassland

Marsh

Fig. 13. Capture rate (%) of Tree Pipits in the four habitats, 2002

Table 3. Data for the Tree Pipit ringed in Tömörd and recovered abroad

Capture Type AgeCode Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile 05.09.2002. Tömörd (AH)

47°21'23"N 16°40'04"E

Recovery - 28.05.2004.Lerviken, Seiplax, Finland

63°20'N 22°23'E 1814 11 631

39. Meadow Pipit (Anthus pratensis) The Meadow Pipit was rare songbird in the study area. In total, 40 individuals of Meadow Pipits were marked between 1998 and 2007. The most birds were captured in 2006, the least in 2004 and 2005. The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=7.54, p>0.05 (Fig. 14). Before 20 August Meadow Pipits were caught only in 2006. On average, the migration of the Meadow Pipit started on the 5th of October (the earliest 30 September, the last 23 October). The average median date of the passage was 2 November. The average date of the end of migration was 11 November. The date of the last capture was also 11 November. The peak period of migration fell on the end of October and the early November in the years 1998-2007. The most capture occurred in September-October (Fig. 15). 82 per cent of the Meadow Pipits were captured in the marsh in 2006, χ2=6, p<0.05 (Fig. 16).

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0

2

4

6

8

10

12

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 14. Yearly capture numbers Meadow Pipits

0

1

2

3

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 15. Average daily captures and migration dynamics (smoothed moving average) of Meadow Pipits

Forest

Bushy

Grassland

Marsh

Fig. 16. Capture rate (%) of Meadow Pipits in the four habitats, 2006

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40. Water Pipit (Anthus spinolenta) One adult bird of this species was captured on the 1st of October, 2003 in the marsh.

41. Yellow Wagtail (Motacilla flava) The Yellow Wagtail was rare songbird in the study area. In total, 15 individuals of Yellow Wagtails were marked between 1998 and 2007. The most birds (7 indiv.) were captured at the end of August and September 2003. 42. Grey Wagtail (Motacilla cinerea) The Grey Wagtail was rare songbird in the study area. In total, 18 individuals of Grey Wagtails were marked between 1998 and 2007. The most birds (14 indiv.) were captured in the second half of August and early September 1998. 43. White Wagtail (Motacilla alba) The White Wagtail was rare songbird in the study area. In total, 67 individuals of White Wagtails were marked between 1998 and 2007. The most birds (20 indiv.) were captured in 2003.

Troglodytidae 44. Wren (Troglodytes troglodytes) The Wren was rare songbird in the post-breeding dispersal period, but it was subdominant passerine during autumn migration in the study area. In total, 539 individuals of Wrens were marked between 1998 and 2007. The most birds were captured in 2000, the least in 2003. The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=7.88, p>0.05 (Fig. 17).

On average, migration of the Wren started on the 29th of September (the earliest 17 August, the last 18 October). The average median date of the passage was 22 October. The average date of the end of migration was 11 November. The date of the last capture was also 11 November. The peak period of migration fell on the mid-October in the years 1998-2007.

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The most captures occurred in October (Fig. 18). According to the distribution of recaptures, birds arrived at the peak period of migration left the area quickly by early November.

0102030405060708090

100

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 17. Yearly capture numbers of Wrens

0

1

2

3

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 18. Average daily captures and migration dynamics

(smoothed moving average) of Wrens

1-Aug 15-Aug 29-Aug 12-Sep 26-Sep 10-Oct 24-Oct 7-Nov

Fig. 19. Individual capture histories of Wrens captured more than once within a season during the study period, 2004 – 2005. Each point indicates a capture

of this bird and lines join the captures of the same individual.

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The quick migration through the area was also suggested by lower proportion of recaptures (8-18%). Wrens spent short time in the area, the maximum was 11 days (Fig. 19). The ratio of overwintering birds was probably very low in the area. 38 per cent of the Wrens were captured in the bushy in 2000, χ2=0, p>0.05 (Fig. 20).

Forest

Bushy

Grassland

Marsh

Fig. 20. Capture rate (%) of Wrens in the four habitats, 2000

Prunellidae

45. Dunnock (Prunella modularis)

The Dunnock was rare songbird in the post-breeding dispersal period, but it was subdominant passerine during autumn migration in the study area. In total, 797 individuals of Dunnoks were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1998. The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=3.81, p>0.05, but the yearly captures showed significant increasing trend between 1999 and 2007 (Fig. 21). On average, migration of the Dunnock started on the 27th of September (the earliest 21 September, the last 17 October). The average median date of the passage was 9 October. The average date of the end of migration was 28 October. The date of the last capture was 8 November. The peak period of migration fell on the first half of October in the years 1998-2007. The most captures occurred in October (Fig. 22). According to the distribution of recaptures, birds arrived at the peak period of migration left the area quickly by early November. The quick migration through the area was also suggested by lower proportion of recaptures (4-16%). Dunnocks spent short time in the area, the maximum was 26 days (Fig. 23). The ratio of overwintering birds was probably very low in the area. 51 per cent of the Dunnocks were captured in the grassland in 2005, χ2=0, p>0.05 (Fig. 24).

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0

20

40

60

80

100

120

140

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 21. Yearly capture numbers of Dunnocks.

Linear regression of yearly captures, r=0.69, p<0.05

0

1

2

3

4

5

6

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 22. Average daily captures and migration dynamics

(smoothed moving average) of Dunnocks

29-Jul

12-Aug

26-Aug

9-Sep

23-Sep

7-Oct

21-Oct

4-Nov

18-Nov

Fig. 23. Individual capture histories of Dunnocks captured more than once within a season during the study period, 2006. Each point indicates a capture of this bird

and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 24. Capture rate (%) of Dunnocks in the four habitats, 2005

Turdidae

46. Robin (Erithacus rubecula)

The Robin was most dominant songbird in the study area. In total, 6375 individuals of Robins were marked between 1998 and 2007. During the autumn migration the most birds were captured in 2007, the least in 1998 (Fig. 25). The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=9.16, p>0.05, but the yearly captures showed a significant increasing trend from 1999 to 2007 (Fig. 25).

0100200300400500600700800900

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 25. Yearly capture numbers of Robins.

Linear regression of yearly captures, r=0.67, p<0.05

The course of the catching dynamics was very similar for years and age groups, Kruskal-Wallis test, p>0.05. On average, migration of the

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Robin started on the 24th of August (the earliest 27 July, the last 22 September). The average median date of the passage was 3 October. The average date of the end of migration was 15 November. The date of the last capture was 18 November. The peak period of migration fell on the end of September and mid-October in the years 1998-2007. The most captures occurred in October (Fig. 26). The apparent shift of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.34, p>0.05 (Fig. 27).

The proportion of recaptured birds was 16-39%, however, it significantly varied from year to year. Majority of Robins spent only a short time, whereas some Robins spent a longer time in the area. The mean minimum stopover time of recaptured birds was 4.2 ± 4.94 days, the maximum was 64 days. Local breeding birds ringed in August left the study area by the end of September (Fig. 28).

The Robins were grouping in the bushy areas, the width of their habitat was small compared to other species (GYURÁCZ et al. 2005). 49 per cent of the Robins were captured in the bushy in 2007, χ2=2, p>0.05 (Fig. 29). Individuals captured in the bushy were fatter than those captured in the forest, grassland and marsh (ANOVA, p<0.01) (Table 4). The most of the Robins ringed in Tömörd probably wintering in Italy (Table 5).

0

5

10

15

20

25

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 26. Average daily captures and migration dynamics

(smoothed moving average) of Robins

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26-Jul

5-Aug

15-Aug

25-Aug

4-Sep

14-Sep

24-Sep

4-Oct

1998 2000 2002 2004 2006 Fig. 27. Changes of date of the beginning of autumn migration of Robins

1-Aug 15-Aug 29-Aug 12-Sep 26-Sep 10-Oct 24-Oct 7-Nov

Fig. 28. Individual capture histories of Robins captured more than once within a season during the study period, 2004. Each point indicates a capture of this bird

and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 29. Capture rate (%) of Robins in the four habitats, 2005

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Table 3. Average fat score of Robins captured id different habitats 1 Oct – 8 Nov

One-way ANOVA, Tukey HSD test

Forest Bushy Grassland and

marsh

N 313 269 261

Mean 1.98 2.53 1.67

S.D. 1.41 1.28 1.26

Forest - p<10-5 P<0.01

Bushy - p<0.001

Table 4. Data for all the Robins ringed in Tömörd and recovered abroad

Capture Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile 20.09.1999. Tömörd (AH) 47°22'N 16°41'E

1. Recovery adult 22.01.2002.

Arbus (Campagna di),

Italy

39°32'N 08°36'E 1089 217 855

Ringing adult 12.09.1999. Tömörd (AH) 47°22'N 16°41'E

2. Recovery - 03.05.2000. Anterselva,

Italy 46°52'N 12°05'E 353 261 234

Ringing juvenile 10.09.2004. Tömörd (AH) 47°21'23"N 16°40'04"E

3. Recovery adult 05.05.2006.

Klostermarienberg, Oberpullendorf,

Austria

47°25'N 16°34'E 10 313 602

Ringing adult 14.03.2001. Tömörd (AH) 47°22'N 16°41'E

4. Recovery adult 17.03.2003. Ladná, Czech

Republic 48°48'N 16°52'E 160 5 733

47. Thrush Nightingale (Luscinia luscinia) The Thrush Nightingale was rare songbird in the study area. In total, 28 individuals of Thrush Nightingales were marked between 1998

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and 2007. The most birds (4 indiv.) were captured in 2003, 2004 and in 2006. All birds were captured in August. 48. Nightingale (Luscinia megarhynchos) The Nightingale was rare songbird in the study area. In total, 71 individuals of Nightingales were marked between 1998 and 2007. The most birds (12 indiv.) were captured in 2003. The most birds were captured in August. The date of the last capture was 21 September. 49. Bluethroat (Luscinia luscinia) The Bluethroat was very rare songbird in the study area. In total, 4 individuals of Bluethroats were marked between 1998 and 2007. The date of the captures: 16 September 1999, 3 October 2000, 3 September 2003, 15 September 2007. 50. Black Redstart (Phoenicurus ochruros)

The Black Redstart was accessory songbird in the study area. In total, 275 individuals of Black Redstarts were marked between 1998 and 2007. The most birds were captured in 2001, the least were in 1998. The average daily captures did not change significantly, Kruskal-Wallis test, H=3.92, p>0.05 (Fig. 30).

05

1015202530354045

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 30. Yearly capture numbers of Black Redstarts

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On average, migration of the Black Redstart started on the 14th of October (the earliest 4 October, the last 23 October). The average median date of the passage was 19 October. The average date of the end of migration was 6 November. The date of the last capture was 10 November. The peak period of migration fell in the second half of October in the years 1998-2007. The most captures occurred in October (Fig. 31). There were no recaptures. 50 per cent of the Black Redstarts were captured in the grassland in 2001, χ2=0, p>0.05 (Fig. 32).

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 31. Average daily captures and migration dynamics

(smoothed moving average) of Black Redstarts

ForestBushyGrasslandMarsh

Fig. 32. Capture rate (%) of Black Redstarts in the four habitats, 2001

51. Redstart (Phoenicurus phoenicurus)

The Redstart was accessory songbird in the study area. In total, 161 individuals of Redstarts were marked between 1998 and 2007. The most

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birds were captured in 2001, the least in 1998. The average daily captures of the years did not change significantly, Kruskal-Wallis test, H=1.19, p>0.05 (Fig. 33).

0

5

10

15

20

25

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 33. Yearly capture numbers of Redstarts

On average, migration of the Redstart started on the 9th of

September (the earliest 11 August, the last 20 September). The average median date of the passage was 22 September. The average date of the end of migration was 3 October. The date of the last capture was 20 October. The peak period of migration fell on the mid-September in the years 1998-2007. The most captures occurred in September (Fig. 34). There were no recaptures. 40 per cent of the Redstarts were captured in the grassland in 2001, χ2=0, p>0.05 (Fig. 35).

0

1

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 34. Average daily captures and migration dynamics

(smoothed moving average) of Redstarts

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ForestBushyGrasslandMarsh

Fig. 35. Capture rate (%) of Redstarts in the four habitats, 2001

52. Whinchat (Saxicola rubetra) The Whinchat was accessory songbird in the study area. In total, 183 individuals of Whinchats were marked between 1998 and 2007. The most birds were captured in 2002, there was no capture in 1999. The average daily captures changed significantly, Kruskal-Wallis test, H=29.53, p<0.001 (Fig. 36).

010203040

50607080

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 36. Yearly capture numbers of Whinchats

On average, migration of the Whinchats started on the 25th of

August (the earliest 4 August, the last 12 September). The average median date of the passage was 6 September. The average date of the end of

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migration was 22 September. The date of the last capture was 5 October. The peak period of migration fell in the end of August and early September in the years 1998-2007. The most captures occurred in August (Fig. 37). 92 per cent of the Whinchats were captured in the grassland in 2002, χ2=6, p<0.05 (Fig. 38).

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 37. Average daily captures and migration dynamics (smoothed moving average) of Whinchats

ForestBushyGrasslandMarsh

Fig. 38. Capture rate (%) of Whinchats in the four habitats, 2002

53. Stonechat (Saxicola torquata) The Stonechat was accessory songbird in the study area. In total, 274 individuals of Stonechats were marked between 1998 and 2007. The most birds were captured in 2002, there was no capture in 1998. The

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average daily captures changed significantly, Kruskal-Wallis test, H=20.95, p<0.01 (Fig. 39).

0

10

20

30

40

50

60

70

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 39. Yearly capture numbers of Stonechats

On average, migration of the Stonechats started on the 13th of September (the earliest 13 August, the last 5 October). The average median date of the passage was 3 October. The average date of the end of migration was 18 October. The date of the last capture was 23 October. The peak period of migration fell in the second half of September in the years 1998-2007. The most captures occurred in September (Fig. 40). 60 per cent of the Stonechats were captured in the marsh in 2002, χ2=2, p>0.05. There were no captures in the forest and bushy (Fig. 41).

0123456789

10

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 40. Average daily captures and migration dynamics

(smoothed moving average) of Stonechats

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ForestBushyGrasslandMarsh

Fig. 41. Capture rate (%) of Stonechats in the four habitats, 2002

Table 60. Data for the Stonechats ringed at the other Hungarian ringing sites and

recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing adult male 27.09.2000.

Keszthely-Fenékpuszta,

Balaton

46°42'55"N 17°14'48"E

Recovery adult male 04.06.2004. Tömörd 47°22'N

16°41'E 85 330 1346

54. Wheatear (Oenanthe oenanthe) The Wheatear was very rare songbird in the study area. In total, 10 individuals of Wheatears were marked between 1998 and 2007. The most birds (4 indiv.) were captured in 2002. The date of the last capture was 1 October. 55. Blackbird (Turdus merula) The Blackbird was dominant songbird in the study area. In total, 1997 individuals of Blackbirds were marked between 1998 and 2007. The most birds were captured in 2007, the least in 2002. The average daily captures changed significantly, Kruskal-Wallis test, H=49.64, p<0.0001 (Fig. 42). On average, migration of the Blackbirds started on the 1st of October (the earliest 6 September, the last 21 October). The date of the earliest beginning of the migration was 6 September in 2004. The average median date of the passage was 17 October. The average date of the end of

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migration was 18 November. The date of the last capture was 18 November. The peak period of migration fell in the mid-October in the years 1998-2007. The most captures occurred in October (Fig. 43).

0

100

200

300

400

500

600

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 42. Yearly capture numbers of Blackbirds

012345678

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 43. Average daily captures and migration dynamics

(smoothed moving average) of Blackbirds

Proportion of recaptured birds were 13-33%, however, it varied significantly from year to year. According to the distribution of recaptures, some of the local breeding birds captured in August stayed in the study area until early November and probably overwintered in the study area. The most of birds arriving at the peak period of migration leave the area quickly by early November. Some of them probably also overwintered in

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the study area (Fig. 45) 46 per cent of the Blackbirds were captured in the bushy in 2007, χ2=0, p>0.05 (Fig. 44).

Forest

Bushy

Grassland

Marsh

Fig. 44. Capture rate (%) of Blackbirds in the four habitats, 2007

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

Fig. 45. Individual capture histories of Blackbirds captured more than once within a season during the study period, 2006. Each point indicates a capture of this bird

and lines join the captures of the same individual.

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The number of recoveries abroad was 11. The direction of migration averaged 225.17° (SW) and the distance of migration averaged 667.17km for Blackbirds. The most Blackbirds ringed in Tömörd wintered at the Apennine peninsula from mid-October to the end of January (Table 7).

Table 7. Data for all the Blackbirds ringed in Tömörd and recovered abroad Capture

Type Age/Sex Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing juvenile male 21.08.2003. Tömörd (AH) 47°21'23"N

16°40'04"E 1.

Recovery adult female 12.10.2003. M. Proace,

Idro, Italy 45°43'N 10°28'E 509 249 52

Ringing juvenile male 03.10.1998. Tömörd (AH) 47°22'N

16°41'E 2.

Recovery juvenile male 01.11.2001. Portovenere,

Italy 44°03'N 09°51'E 647 235 1125

Ringing adult female 20.08.1998. Tömörd (AH) 47°21'23"N

16°40'04"E 3.

Recovery adult female 22.12.2002. Les Aires,

France 43°35'N 03°04'E 1141 248 1585

Ringing juvenile male 30.10.1998. Tömörd (AH) 47°22'N

16°41'E 4.

Recovery adult 08.11.1998. Caino, Italy 45°37'N 10°19'E 525 248 9

Ringing juvenile male 27.09.1999. Tömörd (AH) 47°21'23"N

16°40'04"E

5. Recovery adult 28.12.2000.

Cimini (Monti),

Viterbo, Italy

42°25'N 12°09'E 655 213 458

Ringing juvenile female 13.10.1999. Tömörd (AH) 47°21'23"N

16°40'04"E

6. Recovery adult 14.12.1999.

Montegaudio, Monteciccardo,

Italy

43°48'N 12°47'E 498 217 62

Ringing juvenile male 13.10.1999. Tömörd (AH) 47°21'23"N

16°40'04"E

7. Recovery adult 23.12.2000.

Mandela (Comune di),

Italy

42°02'N 12°55'E 663 207 437

Ringing juvenile male 14.10.1999. Tömörd (AH) 47°22'N

16°41'E

8. Recovery adult

male 31.01.2003.Santa-Lucia-di-Mercurio,

France

42°19'N 09°13'E 814 226 1205

Ringing juvenile male 18.09.2000. Tömörd (AH) 47°21'23"N

16°40'04"E 9.

Recovery adult male 18.10.2000. Capena, Italy 42°09'N

12°33'E 664 209 40

10. Ringing juvenile male 27.10.2000. Tömörd (AH) 47°21'23"N

16°40'04"E

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Recovery adult 15.11.2000. Monte Sant Angelo, Italy

41°42'N 15°57'E 632 185 19

Ringing juvenile female 22.10.2003. Tömörd (AH) 47°21'23"N

16°40'04"E

11. Recovery adult 15.11.2003.

P.Te Del Diavolo,

Tarquinia, Italy

42°15'N 11°45'E 688 214 24

Ringing juvenile male 30.10.2003. Tömörd

(AH) 47°21'23"N 16°40'04"E

12. Recovery adult 21.10.2004.

Colognola Del Piano,

Bergamo, Italy

45°40'N 09°39'E 570 251 357

56. Fieldfare (Turdus pilaris) The Fieldfare was rare songbird in the study area. In total, 14 individuals of Fieldfares were marked between 1998 and 2007. The most birds (5 indiv.) were captured after 25 October in 2007. There were no captures in 1998, 2001, 2003 and 2006. 57. Song Thrush (Turdus philomelos) The Song Thrush was subdominant songbird in the study area. In total, 833 individuals of Song Thrushes were marked between 1998 and 2007. The most birds were captured in 2007, the least in 1988. The average daily captures changed significantly, Kruskal-Wallis test, H=24.52, p<0.001, and the yearly captures showed a significant increasing trend from 1999 to 2007 (Fig. 46).

020

406080

100120

140160

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 46. Yearly capture numbers of Song Thrushes. Linear regression of yearly captures, r=0.76, p<0.01

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On average, migration of the Song Thrushes started on the 4th of October (the earliest 1 September, the last 21 October). The date of the earliest beginning of the migration was 1 September in 2007. The average median date of the passage was 16 October. The average date of the end of migration was 5 November. The date of the last capture was 9 November. The peak period of migration fell in the early October in the years 1998-2007. The most captures occurred in October (Fig. 47). Proportion of recaptured birds were 5-10%, however, it varied from year to year. According to the distribution of recaptures, the local breeding birds captured in August left the study area by mid-September and the most of birds arrived in October left the area quickly by early November (Fig. 48). 51 per cent of the Song Thrushes were captured in the bushy in 2002, χ2=1, p>0.05 (Fig. 49).

0

1

2

3

4

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 47. Average daily captures and migration dynamics

(smoothed moving average) of Song Thrushes

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov Fig. 48. Individual capture histories of Song Thrushes captured more than once within a season during the study period, 2006. Each point indicates a capture

of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 49. Capture rate (%) of Song Thrushes in the four habitats, 2007

The number of recoveries abroad was 6. The direction of migration

averaged 220.33° (SSW) and distance of migration averaged 681.5km for Song Thrushes. The most Song Thrushes ringed in Tömörd wintered at the Apennine Peninsula from mid-October to the end of January (Table 8).

Table 8. Data for all the Song Thrushes ringed in Tömörd and recovered abroad Capture

Type Age/Sex Date Place Coordinates Text

Distancekm

Direction degree

Elapsed days

Ringing juvenile 03.11.2002. Tömörd (AH)

47°21'23"N 16°40'04"E

1.

Recovery adult 17.10.2005. Marsaz,

France 45°07'N 04°56'E 937 255 1079

Ringing juvenile 08.10.2000. Tömörd (AH)

47°21'23"N 16°40'04"E

2.

Recovery adult 11.11.2000.

Castel Rinaldi, Massa

Martana, Italy

42°48'N 12°31'E 603 213 34

Ringing juvenile 17.10.2000. Tömörd (AH)

47°21'23"N 16°40'04"E

3.

Recovery adult 12.11.2000. Guidonia, Italy

42°01'N 12°45'E 670 208 26

4. Ringing juvenile 06.11.2000. Tömörd (AH)

47°21'23"N 16°40'04"E

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Recovery adult 04.11.2001.

Bosco Signura, Maglie,

Italy

40°08'N 18°17'E 814 171 59

Ringing juvenile 26.10.2003. Tömörd (AH)

47°21'23"N 16°40'04"E

5. Recovery adult 24.10.2005.

Peglio, Firenzuola,

Italy

44°09'N 11°22'E 545 229 729

Ringing juvenile 28.10.2006. Tömörd (AH)

47°21'23"N 16°40'04"E

6.

Recovery adult 05.10.2007.

Cendella, Lonato, Bresica,

Italy

45°27'N 10°29'E 520 246 342

58. Redwing (Turdus iliacus) The Redwing was accessory songbird in the study area. In total, 356 individuals of Redwings were marked between 1998 and 2007. The most birds were captured in 2001, there were no captures in 1988. The average daily captures did not change significantly, Kruskal-Wallis test, H=10.7, p>0.05 (Fig. 50). On average, migration of the Redwings started on the 22nd of October (the earliest 20 October, the last 24 October).

0

20

40

60

80

100

120

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 50. Yearly capture numbers of Redwings

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The date of the earliest capture was 14 October. The average median date of the passage was 3 November. The date of the last capture was 19 November. The peak period of migration fell in the early November in the years 1998-2007. The most captures occurred in November (Fig. 51).

Proportion of recaptured birds were 0 and 19 %, however, it varied from year to year. According to the distribution of recaptures, the most of birds arrived in October-November left the area quickly by mid-November, but some birds overwintered in Tömörd (Fig. 52). 51 per cent of the Redwings were captured in the bushy in 2001, χ2=, p>0.05 (Fig. 53). The main migration direction was presumed SE from Tömörd (Table 9).

0

1

2

3

4

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 51. Average daily captures and migration dynamics

(smoothed moving average) of Redwings

28-Jul

11-Aug

25-Aug

8-Sep

22-Sep

6-Oct

20-Oct

3-Nov

17-Nov

Fig. 52. Individual capture histories of Song Thrushes captured more than once within a season during the study period, 2006. Each point indicates a capture

of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Fig. 53. Capture rate (%) of Redwings in the four habitats, 2001

Table 9. Data for all the Redwings ringed in Tömörd and recovered abroad Capture

Type Age Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing adult 12.10.1999. Tömörd (AH)

47°22'N 16°41'E

Recovery

adult

08.01.2000. Attiki, Greece

38°13'N 23°56'E 1178 150 88

59. Mistle Thrush (Turdus viscivorus) One bird of this species was captured on the 1st of November 2002 in the bushy.

Sylviidae 60. Grasshopper Warbler (Locustella naevia) The Grasshopper Warbler was accessory songbird in the study area. In total, 129 individuals of Grasshopper Warblers were marked between 1998 and 2007. The most birds were captured in 2003, the least in 2002 (Fig. 54). The average daily captures did not change significantly, Kruskal-Wallis test, H=12.28, p>0.05, but the yearly captures showed a significant decreasing trend from 2003 to 2007 (Fig. 55).

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54

0

5

10

15

20

25

30

35

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 54. Yearly capture numbers Grasshopper Warblers

0

5

10

15

20

25

30

2003 2004 2005 2006 2007

No.

of b

irds

Fig 55. Linear regression of yearly captures, r=-0.94, p<0.02

On average, migration of the Grasshopper Warblers started on the

20th of August (the earliest 2 August, the last 10 September) . The average median date of the passage was 12 September. The average date of the end of migration was 20 September. The date of the last capture was 3 October. The most captures occurred in August in the post-breeding dispersal (Fig. 56). 45 per cent of the Grasshopper Warblers were captured in the grassland in 2003, χ2=0, p>0.05. There were no captures in the forest (Fig. 57).

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0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 56. Average daily captures and migration dynamics (smoothed moving average) of Grasshopper Warblers

Bushy

Grassland

Marsh

Fig. 57. Capture rate (%) of Grasshopper Warblers in the four habitats, 2003

61. River Warbler (Locustella fluviatilis) The River Warbler was rare songbird in the study area. In total, 28 individuals of River Warblers were marked between 1998 and 2007. The most birds (9 indiv.) were captured in August and first half of September, 2004. The date of the last capture was 13 September. 62. Savi’s Warbler (Locistella luscinioides) The Savi’s Warbler was very rare songbird in the study area. In total, 8 individuals of Savi’s Warblers were marked between 1998 and

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2007. The first capture was in 2003. The most birds (3-3 indiv.) were captured from the end of July to mid-September in 2005 and 2007. The date of the last capture was 11 September. 63. Moustached Warbler (Acrocephalus melanopogon) One bird of this species was captured on 24 August 2004 in the marsh. 64. Sedge Warbler (Acrocephalus schoenobaenus) The Sedge Warbler was accessory songbird in the study area. In total, 229 individuals of Sedge Warblers were marked between 1998 and 2007. The most birds were captured in 2004, the least in 2001 when the lake was dried out. The average daily captures changed significantly, Kruskal-Wallis test, H=36.43, p<0.0001 (Fig. 58).

0

20

40

60

80

100

120

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 58. Yearly capture numbers of Sedge Warblers

On average, migration of the Sedge Warblers started on the 19th of August (the earliest 31 July, the last 14 September). The average median date of the passage was 26 August. The average date of the end of migration was 29 September. The date of the last capture was 28 September. The most captures occurred in August in the post-breeding dispersal (Fig. 59).

Proportion of recaptured birds were 0 and 30 %, however, it varied from year to year. According to the distribution of recaptures, the local

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birds ringed in the post-breeding dispersal left the study area by mid-August, the most of birds arrived in August left the area quickly by early September (Fig. 60). 84 per cent of the Sedge Warblers were captured in the marsh in 2004, χ2=4, p<0.05 (Fig. 61). Some Sedge Warblers migrated above Tömörd originated from Sweden (Table 10).

0

1

2

3

4

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 59. Average daily captures and migration dynamics

(smoothed moving average) of Sedge Warblers

31-Jul 14-Aug 28-Aug 11-Sep 25-Sep 9-Oct 23-Oct 6-Nov

Fig. 60. Individual capture histories of Sedge Warblers captured more than once within a season during the study period, 2004. Each point indicates a capture

of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 61. Capture rate (%) of Sedge Warblers in the four habitats, 2004

Table 10. Data for all the Sedge Warblers ringed in Tömörd and recovered abroad and also those ringed abroad and recovered in Tömörd

Capture Type Age Date Place Coordinates

Text Distance

km Direction Degree

Elapsed days

Ringing juvenile 13.08.2004. Tömörd (AH)

47°21'23"N 16°40'04"E

1. Recovery adult 15.05.2005.

Rynningeviken, Venan, Sweden

59°17'N 15°15'E 1332 356 275

Ringing juvenile 26.08.2005.

Vnanje Gorice, Ljubjansko

barje Slovenia

46°00'N 14°25'E

2.

Recovery juvenile 04.09.2005. Tömörd (AH)

47°21'23"N 16°40'04"E 229 49 9

65. Marsh Warbler (Acrocephalus palustris)

The Marsh Warbler was accessory songbird in the study area. In total, 242 individuals of Marsh Warblers were marked between 1998 and 2007. The most birds were captured in 2004, the least in 2000 and 2001 when the lake dried out. The average daily captures of the years changed significantly, Kruskal-Wallis test, H=20.93, p<0.01 (Fig. 62).

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0102030405060708090

100

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 62. Yearly capture numbers of Marsh Warblers

On average, migration of the Marsh Warblers started on the 7th of

August (the earliest 1 August, the last 20 August). The average median date of the passage was 26 August. The average date of the end of migration was 7 September. The date of the last capture was 30 September. The most captures occurred in August in the post-breeding dispersal (Fig. 63).

The recapture ratio was 18 % in 2004. The minimum stopover time of the recaptured individuals was min. 2, max. 9 days in 2004 (Fig. 65). 64 per cent of the Marsh Warblers were captured in the marsh in 2004, χ2=2, p>0.05 (Fig. 66).

0

1

2

3

4

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

No.

of b

irds

Fig. 64. Average daily captures and migration dynamics (smoothed moving average) of Marsh Warblers

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29-Jul 18-Aug 7-Sep 27-Sep 17-Oct 6-Nov

Fig. 65. Individual capture histories of Marsh Warblers captured more than once within a season during the study period, 2004. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 66. Capture rate (%) of Marsh Warblers in the four habitats, 2004

66. Reed Warbler (Acrocephalus scirpaceus) The Reed Warbler was rare songbird in the study area. In total, 93 individuals of Reed Warblers were marked between 1998 and 2007. The most birds were captured in 2004, the least in 2000 and 2001 when the lake dried out. The average daily captures did not change significantly, Kruskal-Wallis test, H=4.28, p>0.05 (Fig. 67).

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0

5

10

15

20

25

30

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 67. Yearly capture numbers of Reed Warblers

On average, migration of the Reed Warblers started on the 24th of

August (the earliest 9 August, the last 20 September). The average median date of the passage was 5 September. The average date of the end of migration was 25 September. The date of the last capture was 7 October. The most captures occurred August in the post-breeding dispersal (Fig. 68).

The recapture ratio was 12 % in 2004. The minimum stopover time of the recaptured individuals was min. 2, max. 4 days in 2004. 84 per cent of the Reed Warblers were captured in the marsh in 2004, χ2=6, p<0.05 (Fig. 69).

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 68. Average daily captures and migration dynamics (smoothed moving average) of Reed Warblers

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Grassland

Marsh

Fig. 69. Capture rate (%) of Reed Warblers in the four habitats, 2004

67. Great Reed Warbler (Acrocephalus arundinaceus) The Great Reed Warbler was very rare songbird in the study area. In total, 9 individuals of Great Reed Warblers were marked between 1998 and 2007. The first capture was in 2003. The most birds (4 indiv.) were captured in the early August 2004. The date of the last capture was 19 August. 68. Icterine Warbler (Hippolais icterina)

The Icterine Warbler was accessory songbird in the study area. In total, 151 individuals of Icterine Warblers were marked between 1998 and 2007. The most birds were captured in 2005, the least in 2000 when the lake dried out. The average daily captures did not change significantly, Kruskal-Wallis test, H=7.6, p>0.05 (Fig. 70).

On average, migration of the Icterine Warblers started on the 19th of August (the earliest 6 August, the last 31 August). The average median date of the passage was 27 August. The average date of the end of migration was 19 September. The date of the last capture was 28 September. The peak period of migration fell on the second half of August in the years 1998-2007. The most captures occurred in August (Fig. 71).

Proportion of recaptured birds was 30% in 2005. The minimum stopover time of the recaptured individuals was min. 2, max. 10 days in 2005 (Fig. 72). 41 - 41 per cent of the Icterine Warblers were captured in the bushy and grassland in 2005, χ2=4, p<0.05 (Fig. 73).

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0

5

10

15

20

25

30

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 70. Yearly capture numbers of Icterine Warblers

0

1

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 71. Average daily captures and migration dynamics

(smoothed moving average) of Icterine Warblers

29-Jul 18-Aug 7-Sep 27-Sep 17-Oct 6-Nov Fig. 72. Individual capture histories of Icterine Warblers captured more than once

within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 73. Capture rate (%) of Icterine Warblers in the four habitats, 2005

69. Barred Warbler (Sylvia nisoria) The Barred Warbler was rare songbird in the study area. In total, 30 individuals of Barred Warblers were marked between 1998 and 2007. The most birds (8-8 indiv.) were captured in 2002 and 2003. The date of the last capture was 27 September. 70. Lesser Whitethroat (Sylvia curruca)

The Lesser Whitethroat was dominant songbird in the study area. In total, 1045 individuals of Lesser Whitethroats were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=18.08, p<0.05 (Fig. 1).

On average, migration of the Lesser Whitethroats started on the 7th of August (the earliest 27 July, the last 8 September). The average median date of the passage was 29 August. The average date of the end of migration was 1 October. The date of the last capture was 12 October. The peak period of migration fell on the end of August and the early September in the years 1998-2007. The most captures occurred in August (Fig. 75). The apparent shift of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.47, p<0.05 (Fig. 76). Proportion of recaptured birds was 25% in 2005. The minimum stopover time of the recaptured individuals was min. 2, max. 16 days in 2005 (Fig. 77). 88 per cent of the Lesser Whitethroats were captured in the grassland in 2005, χ2=4, p<0.05 (Fig. 78). Some Lesser Whitethroats migrated above Tömörd originated from Germany and Finland (Table 11).

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020406080

100120140160180

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 74. Yearly capture numbers of Lesser Whitethroat

0

1

2

3

4

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 75. Average daily captures and migration dynamics

(smoothed moving average) of Lesser Whitethroats

21-Jul

31-Jul

10-Aug

20-Aug

30-Aug

9-Sep

19-Sep

1998 2000 2002 2004 2006

Fig. 76. Changes of date of the beginning of autumn migration of Lesser Whitethroats

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31-Jul 14-Aug 28-Aug 11-Sep 25-Sep 9-Oct 23-Oct 6-Nov Fig. 77. Individual capture histories of Lesser Whitethroats captured more than

once within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 78. Capture rate (%) of Lesser Whitethroats in the four habitats, 2005

Table 11. Data for all the Lesser Whitethroats ringed in Tömörd and recovered

abroad and also those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing juvenile 06.08.2003. Tömörd (AH) 47°21'23"N 16°40'04"E

1. Recovery juvenile 23.08.2003.

Bukovac, Sombor,

Jugoslavia

45°47'N 19°05'E 255 133 17

Ringing juvenile 02.09.2004. Tömörd (AH) 47°21'23"N 16°40'04"E

2. Recovery adult 05.06.2005.

Lágskár, Lemland, Finland

59°50'N 19°56'E

1406 9 276

Ringing juvenile 26.08.2003.Numburg,

Kyffhauserkreis,Germany

51°25'N 10°59'E

3.

Recovery adult 17.09.2003. Tömörd (AH) 47°21'23"N 16°40'04"E 612 138 22

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71. Whitethroat (Sylvia communis) The Whitethroat was dominant songbird in the study area. In total, 1033 individuals of Whitethroats were marked between 1998 and 2007. The most birds were captured in 2005, the least in 2000. The average daily captures changed significantly, Kruskal-Wallis test, H=27.51, p<0.001 (Fig. 79). On average, migration of the Whitethroats started on the 10th of August (the earliest 1 August, the last 26 August). The average median date of the passage was 26 August. The average date of the end of migration was 27 September.

020406080

100120140160180200

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 79. Yearly capture numbers of Whitethroats

The date of the last capture was 29 September. The peak period of

migration fell on the mid-August in the years 1998-2007. The most captures occurred in August (Fig. 80). The apparent shift of the beginning of autumn migration towards earlier date was significant in the years 1998-2007, Pearson’s r=-0.68, p<0.05 (Fig. 81).

Proportion of recaptured birds was 13-14% in 2004-2006. The minimum stopover time of the individuals recaptured was min. 2, max. 20 days in 2005 (Fig. 83). 88 per cent of the Whitethroats were captured in the grassland in 2005, χ2=6, p<0.05 (Fig. 82). One bird, which was ringed in Czech Republic, recovered in Tömörd (Table 12).

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0

1

2

3

4

5

6

7

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 80. Average daily captures and migration dynamics

(smoothed moving average) of Whitethroats

26-Jul

31-Jul

5-Aug

10-Aug

15-Aug

20-Aug

25-Aug

30-Aug

1998 2000 2002 2004 2006 Fig. 81. Changes of date of the beginning of autumn migration of Whitethroats

Bushy

Grassland

Marsh

Fig. 82. Capture rate (%) of Whitethroats in the four habitats, 2005

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27-Jul 16-Aug 5-Sep 25-Sep 15-Oct 4-Nov

Fig. 83. Individual capture histories of Whitethroats captured more than once within a season during the study period, 2005. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Table 12. Data for all the Whitethroats ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates Text

Distancekm

Direction degree

Elapsed days

Ringing adult male 06.06.2002.

Studénki Czech

Republic

50°27'N 14°59'E

Recovery adult male 03.09.2002. Tömörd (AH) 47°21'23"N

16°40'04"E 366 160 89

72. Garden Warbler (Sylvia borin) The Garden Warbler was accessory songbird in the study area. In total, 425 individuals of Garden Warblers were marked between 1998 and 2007. The most birds were captured in 2006, the least in 2000 (Fig. 1). The average daily captures changed significantly, Kruskal-Wallis test, H=20.72, p<0.01 and the yearly captures showed a significant increasing trend from 1998 to 2007 (Fig. 84). On average, migration of the Garden Warblers started on the 19th of August (the earliest 7 August, the last 2 September). The average median date of the passage was 5 September. The average date of the end of migration was 23 September. The date of the last capture was 3 October. The peak period of migration fell on the early September in the years 1998-2007. The most captures occurred in August (Fig. 85).

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0102030405060708090

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 84. Yearly capture numbers of Garden Warbler. Linear regression of yearly captures, r=0.80, p<0.01

The apparent shift of the beginning of autumn migration towards

earlier date was significant in the years 1998-2007, Pearson’s r=-0.66, p<0.05 (Fig. 86). There were no recaptures. 62 per cent of the Garden Warblers were captured in the grassland in 2006, χ2=2, p>0.05 (Fig. 87).

0

1

2

3

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 85. Average daily captures and migration dynamics

(smoothed moving average) of Garden Warblers

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26-Jul

5-Aug

15-Aug

25-Aug

4-Sep

14-Sep

24-Sep

1998 2000 2002 2004 2006

Fig. 86. Changes of date of the beginning of autumn migration of Garden Warbler in Tömörd

Forest

Bushy

Grassland

Marsh

Fig. 87. Capture rate (%) of Garden Warblers in the four habitats, 2006

73. Blackcap (Sylvia atricapilla) The Blackcap was dominant songbird in the study area. In total, 4551 individuals of Blackcaps were marked between 1998 and 2007. The most birds were captured in 2007, the least in 2000. The average daily captures changed significantly, Kruskal-Wallis test, H=50.27, p<0.0001 and the yearly captures showed a significant increasing trend from 1998 to 2007 (Fig. 88).

On average, migration of the Blackcaps started on the 20th of August (the earliest 31 July, the last 7 September). The average median date of the passage was 14 September. The average date of the end of

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migration was 31 October. The date of the last capture was 7 November. The peak period of migration fell on the early September and end of September in the years 1998-2007. The most captures occurred in September (Fig. 89). The significant shift of the beginning of autumn migration towards earlier date was not revealed in the years 1998-2007, Pearson’s r=-0.46, p>0.05 (Fig. 90.). Proportion of recaptured birds was 5-10% in 2004-2006. The minimum stopover time of the recaptured individuals was min. 2, max. 57 days in 2005 (Fig. 91). 36 per cent of the Blackcaps were captured in the grassland in 2007, χ2=0, p>0.05 (Fig. 92). Some Blackcaps migrated above Tömörd originated from Czech Republic and Finland (Table 13).

0

100

200

300

400

500

600

700

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 88. Yearly capture numbers Blackcaps.

Linear regression of yearly captures, r=0.95, p<0.001

02468

10121416

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

No.

of b

irds

Fig. 89. Average daily captures and migration dynamics

(smoothed moving average) of Blackcaps

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26-Jul31-Jul5-Aug

10-Aug15-Aug20-Aug25-Aug30-Aug4-Sep9-Sep

14-Sep

1998 2000 2002 2004 2006 Fig. 90. Changes of date of the beginning of autumn migration of Blackcaps

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

Fig. 91. Individual capture histories of Blackcaps captured more than once within a season during the study period. Each point indicates a capture of this bird and

lines join the captures of the same individual.

2005

2006

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Forest

Bushy

Grassland

Marsh

Fig. 92. Capture rate (%) of Blackcaps in the four habitats, 2007

Table 13. Data for all the Blackcaps ringed in Tömörd and recovered abroad and

also those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place CoordinatesText

Distance km

Direction degree

Elapsed days

Ringing juvenile male 11.09.1998. Tömörd

(AH) 47°21'23"N 16°40'04"E

1. Recovery adult

male 28.04.2001.

Lanzhot, Breclav, Czech

Republic

48°43'N 16°58'E 153 8 960

Ringing juvenile female 29.08.1999. Tömörd

(AH) 47°22'N 16°41'E

2. Recovery juvenile

female 28.09.1999. Vrhnika, Slovenia

45°57'N 14°18'E 241 229 30

Ringing juvenile female 14.09.2007. Tömörd

(AH) 47°22'N 16°41'E

3. Recovery juvenile

female 17.06.2008.

Stetteldorf am

Wgram, Korneburg,

Austria

48°24'N 16°01'E 126 337 277

Ringing juvenile female 28.08.2007.

Praha 5 – Hlubocepy,

Czech Republic

50°03'N 14°23'E

4.

Recovery juvenile female 23.09.2007. Tömörd

(AH) 47°21'23"N 16°40'04"E 344 151 26

Ringing juvenile female 16.09.2000.

Hrase, Smlednik, Slovenia

46°10'N 14°27'E

Recovery adult female 30.04.2001. Tömörd

(AH) 47°21'23"N 16°40'04"E 215 52 226

5.

Recovery adult female 02.05.2001. Tömörd

(AH) 47°21'23"N 16°40'04"E 215 52 228

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Recovery adult female 05.05.2001. Tömörd

(AH) 47°21'23"N 16°40'04"E 215 52 231

Ringing adult female 12.17.1999.

Kvismaren, Ängfallet, Sweden

59°11'N 15°24'E

6.

Recovery adult female 25.09.2000. Tömörd 47°22'N

16°41'E 1318 176 441

74. Pallas’s Warbler (Phylloscopus proregulus) The Pallas’s Warbler was very rare songbird in the study area. In total, 2 individuals of Pallas’s Warblers were marked between 1998 and 2007. The dates of the captures were 23 October 2003 and November 2004. 75. Yellow-browed Warbler (Phylloscopus inornatus) The Yellow-browed Warbler was very rare songbird in the study area. In total, 2 individuals of Yellow-browed Warblers were marked between 1998 and 2007. The dates of the captures were 12 October 2002 and 25 September 2007. 76. Wood Warbler (Phylloscopus sibilatrix) The Wood Warbler was accessory songbird in the study area. In total, 163 individuals of Wood Warblers were marked between 1998 and 2007. The most birds were captured in 2002, the least in 1998. The average daily captures did not changed significantly, Kruskal-Wallis test, H=20.47, p>0.05 (Fig. 93).

On average, migration of the Wood Warblers started on the 16th of August (the earliest 29 July, the last 21 September). The average median date of the passage was 26 August. The average date of the end of migration was 16 September. The date of the last capture was 24 September. The peak period of migration fell in the first half of August in the years 1998-2007. The most captures occurred in August (Fig. 94). The significant shift of the beginning of autumn migration towards earlier date was not revealed in the years 2002-2007, Pearson’s r=-0.41, p>0.05 (Fig 95).

One recaptured bird was in 2002, there were no recaptures in the other years. 45 per cent of the Wood Warblers were captured in the forest in 2002, χ2=0, p>0.05 (Fig. 96).

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05

101520

2530

3540

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 93. Yearly capture numbers of Wood Warblers

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 94. Average daily captures and migration dynamics

(smoothed moving average) of Wood Warblers

0

10

20

30

40

50

60

2002 2003 2004 2005 2006 2007

No.

of b

irds

Fig. 95. Changes of date of the beginning of autumn migration of Wood Warblers

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Forest

Bushy

Grassland

Marsh

Fig. 96. Capture rate (%) of Wood Warblers in the four habitats, 2002

77. Chiffchaff (Phylloscopus collybita) The Chiffchaff was dominant songbird in the study area. In total, 3359 individuals of Chiffchaffs were marked between 1998 and 2007. The most birds were captured in 2004, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=36.65, p<0.0001 and the yearly captures showed a significant increasing trend from 1998 to 2007 (Fig. 97).

0

100

200

300

400

500

600

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 97. Yearly capture numbers of Chiffchaffs.

Linear regression of yearly captures, r=0.79, p<0.01

On average, migration of the Chiffchaffs started on the 2nd of September (the earliest 1 August, the last 24 September). The average

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median date of the passage was 5 October. The average date of the end of migration was 6 November.

The date of the last capture was 11 November. The peak period of migration fell on the early October in the years 1998-2007. The most captures occurred in October (Fig. 98). The apparent shift of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.45, p>0.05 (Fig. 99).

Proportion of recaptured birds was 13-22% in 2004-2006. The minimum stopover time of the recaptured individuals was min. 2, max. 26 days in 2005 (Fig. 100). The body mass and fat load of the recaptured birds did not changed significantly during stopover period (t probe, p>0.05). 36 per cent of the Chiffchaffs were captured in the grassland in 2004, χ2=2, p>0.05 (Fig. 101). Some Chiffchaffs migrated above Tömörd originated from the Czech Republic and Austria (Table 14).

0

2

4

6

8

10

12

14

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

Fig. 98. Average daily captures and migration dynamics

(smoothed moving average) of Chiffchaffs

1-Jul11-Jul21-Jul31-Jul

10-Aug20-Aug30-Aug9-Sep

19-Sep29-Sep

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007

Fig. 99. Changes of date of the beginning of autumn migration of Chiffchaffs

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27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

Fig. 100. Individual capture histories of Chiffchaffs captured more than once within a season during the study period, 2005. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 101. Capture rate (%) of Chiffchaffs in the four habitats, 2004

Table 14. Data for all the Chiffchaffs ringed in Tömörd and recovered abroad and

also those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing

juvenile

05.10.2004. Tömörd (AH)

47°21'23"N 16°40'04"E

1.

Recovery adult male 05.05.2007.

Bulhary, Breclav Czech

Republic

48°50'N 16°45'E 165 2 942

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Ringing juvenile 11.10.2003.

Hohenau, Ringelsdorf,

March, Austria

48°35'N 16°55'E

2.

Recovery adult 22.10.2006. Tömörd (AH)

47°21'23"N 16°40'04"E 138 188 1107

78. Willow Warbler (Phylloscopus trochilus) The Willow Warbler was subdominant songbird in the study area. In total, 858 individuals of Willow Warblers were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1999. The average daily captures changed significantly, Kruskal-Wallis test, H=21.05, p<0.01 (Fig. 102).

0

50

100

150

200

250

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 102. Yearly capture numbers of Willow Warblers

On average, migration of the Willow Warblers started on the 19th of

August (the earliest 5 August, the last 3 September). The average median date of the passage was 9 September. The average date of the end of migration was 12 October. The date of the last capture was 14 October. The peak period of migration fell on the early September in the years 1998-2007. The most captures occurred in September (Fig 103). The apparent shift of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.53, p>0.05 (Fig. 104).

Proportion of recaptured birds was 20% in 2005. The minimum stopover time of the recaptured individuals was min. 2, max. 20 days in 2005 (Fig. 105). 80 per cent of the Willow Warblers were captured in the grassland in 2005, χ2=6, p<0.05 (Fig. 106). Some Willow Warblers probably migrated probably to SW from Tömörd (Table 15).

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0

1

2

3

4

5

6

7

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

No.

of b

irds

Fig. 103. Average daily captures and migration dynamics (smoothed moving average) of Willow Warblers

27-Jul1-Aug6-Aug

11-Aug16-Aug

21-Aug26-Aug31-Aug5-Sep

1998 2000 2002 2004 2006

Fig. 104. Changes of date of the beginning of autumn migration of Willow Warblers

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov Fig. 105. Individual capture histories of Willow Warblers captured more than

once within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 106. Capture rate (%) of Willow Warblers in the four habitats, 2005

Table 15. Data for all the Willow Warblers ringed in Tömörd and recovered

abroad and also those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile 29.04.2003. Tömörd (AH)

47°21'23"N 16°40'04"E

1. Recovery adult 22.08.2003.

Lagoa de Santo André,

Portugal

38°05'N 08°47'W 2318 244 115

Ringing

juvenile

26.08.2001. Gbelce, Slovakia

47°51'N 18°29'E

2.

Recovery adult male

31.08.2001. Tömörd

(AH) 47°21'23"N 16°40'04"E 147 248 5

79. Goldcrest (Regulus regulus) The Goldcrest was dominant songbird in the study area. In total, 2064 individuals of Goldcrests were marked between 1998 and 2007. The most birds were captured in 2000, the least in 2002. The average daily captures changed significantly, Kruskal-Wallis test, H=54.64, p<0.0001 (Fig. 107). On average, migration of the Goldcrests started on the 4th of October (the earliest 2 September, the last 22 October). The average median date of the passage was 24 October. The average date of the end of migration was 10 November. The date of the last capture was 18

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November. The peak period of migration fell on the end of October in the years 1998-2007. The most captures occurred in October (Fig. 108).

0

100

200

300

400

500

600

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 107. Yearly capture numbers of Goldcrests

The seasonal dynamics of sexes was somewhat different in 2000

and 2001. It seemed that the share of males increased during peak period of migration (Fig. 109) (GYURÁCZ et al. 2003a). The apparent shift of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.45, p>0.05 (Fig. 110). Proportion of recaptured birds was 5-8% in the years 2000-2006. The minimum stopover time of the recaptured individuals was min. 2, max. 26 days in 2004 (Fig. 111). 64 per cent of the Goldcrests were captured in the bushy, χ2=2, p>0.05 (Fig. 112). The most Goldcrests migrating above Tömörd probably originated probably from the Baltic region and NW Russia (Table 16).

02468

1012141618

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 108. Average daily captures and migration dynamics

(smoothed moving average) of Goldcrests

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Fig. 109. Migration dynamics of the male and female Goldcrests in 2000

11-Aug

21-Aug

31-Aug

10-Sep

20-Sep

30-Sep

10-Oct

20-Oct

30-Oct

1998 1999 2000 2001 2002 2003 2004 2005 2006 2007

Fig. 110. Changes of date of the beginning of autumn migration of Goldcrest

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

Fig. 111. Individual capture histories of Willow Warblers captured more than once within a season during the study period, 2004. Each point indicates a

capture of this bird and lines join the captures of the same individual.

0

20

40

60

80

100

24-Aug

7- Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

%

male

female

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Forest

Bushy

Grassland

Marsh

Fig. 112. Capture rate (%) of the Goldcrests in the four habitats, 2000

Table 16. Data for all the Goldcrests ringed abroad and recovered in Tömörd

Capture Type

Age/Sex Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing adult female 02.10.2005. Gumbaritsy,

Russia 60°41'N 32°57'E

Recovery adult female 06.04.2006. Tömörd (AH) 47°21'23"N

16°40'04"E 1820 215 186 1.

Recovery adult female 07.04.2006. Tömörd (AH) 47°21'23"N

16°40'04"E 1820 215 187

Ringing juvenile 05.10.2001.

Rybachiy, Zelenogradskiy

distr. Russia

55°09'N 20°51'E

2.

Recovery juvenile female 15.11.2001. Tömörd (AH) 47°21'23"N

16°40'04"E 915 199 41

80. Firecrest (Regulus ignicapillus) The Firecrest was accessory songbird in the study area. In total, 311 individuals of Firecrests were marked between 1998 and 2007. The most birds were captured in 1998, the least in 2002. The average daily captures did not change significantly, Kruskal-Wallis test, H=3.93, p>0.05 (Fig. 113).

On average, migration of the Firecrests started on the 16th of September (the earliest 17 August, the last 7 October). The average median date of the passage was 6 October. The average date of the end of migration was 5 November. The date of the last capture was 7 November. The peak period of migration fell on the end of October in the years 1998-2007. The most captures occurred in October (Fig. 114). The apparent shift

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of the beginning of autumn migration towards earlier date was not significant in the years 1998-2007, Pearson’s r=-0.53, p>0.05 (Fig. 115).

0

10

20

30

40

50

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 113. Yearly capture numbers of Firecrests

Proportion of recaptured birds was 2 and 12 % in the years 1998 and 2007. The minimum stopover time of the recaptured individuals was min. 2, max. 4 days. 42 per cent of the Firecrests were captured in the bushy, χ2=2, p>0.05 (Fig. 116).

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 114. Average daily captures and migration dynamics

(smoothed moving average) of Firecrests

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26-Jul5-Aug

15-Aug25-Aug4-Sep

14-Sep24-Sep

4-Oct14-Oct

1998 2000 2002 2004 2006

Fig. 115. Changes of date of the beginning of autumn migration of Firecrests

Forest

Bushy

Grassland

Fig. 116. Capture rate (%) of Firecrests in the four habitats, 1998

Muscicapidae 81. Spotted Flycatcher (Muscicapa striata) The Spotted Flycatcher was accessory songbird in the study area. In total, 282 individuals of Spotted Flycatchers were marked between 1998 and 2007. The most birds were captured in 2002, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=14.86, p<0.05 (Fig. 117). On average, migration of the Spotted Flycatchers started on 25 August (the earliest 1 August, the last 9 September). The average median date of the passage was 12 September.

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88

0

10

20

30

40

50

60

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 117. Yearly capture numbers of Spotted Flycatchers

The average date of the end of migration was 26 September. The date of the last capture was 15 October. The peak period of migration fell on the mid-September in the years 1998-2007. The most captures occurred in September (Fig. 118). The significant shift of the beginning of autumn migration towards earlier date was not revealed in the years 1998-2007, Pearson’s r=-0.61, p>0.05 (Fig. 119). Proportion of recaptured birds was 3 % in 2002. The minimum stopover time of the recaptured individuals was min. 13, max. 17 days. 51 per cent of the Spotted Flycatchers were captured in the forest, χ2=0, p>0.05 (Fig. 120).

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 118. Average daily captures and migration dynamics

(smoothed moving average) of Spotted Flycatchers

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26-Jul31-Jul5-Aug

10-Aug15-Aug20-Aug25-Aug30-Aug4-Sep9-Sep

14-Sep

1998 2000 2001 2002 2003 2004 2005 2006 2007

Fig. 119. Changes of date of the beginning of autumn migration of Spotted Flycatchers

Forest

Bushy

Grassland

Marsh

Fig. 120. Capture rate (%) of Spotted Flycatchers in the four habitats, 2002

82. Red-breasted Flycatcher (Ficedula parva) The Red-breasted Flycatcher was rare songbird in the study area. In total, 15 individuals of Red-breasted Flycatchers were marked between 1998 and 2007. The most birds (3-3 indiv.) were captured in 2001, 2002 and 2003. There were no captures in 2004, 2005 and 2006. The date of first capture was 16 August; the date of the last capture was 7 October.

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83. Collared Flycatcher (Ficedula albicollis) The Collared Flycatcher was rare songbird in the study area. In total, 36 individuals of Collared Flycatchers were marked between 1998 and 2007. The most birds (8 indiv.) were captured in 2003. There were no captures in 1998, 1999 and 2000. The most birds were captured in August. The date of first captures was 30 July; the date of the last capture was 31 October. 84. Pied Flycatcher (Ficedula hypoleuca) The Pied Flycatcher was subdominant songbird in the study area. In total, 843 individuals of Pied Flycatchers were marked between 1998 and 2007. The most birds were captured in 2001, the least in 2005. The average daily captures changed significantly, Kruskal-Wallis test, H=19.13, p<0.01 (Fig. 121).

0102030405060708090

100110120130140

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 121. Yearly capture numbers of Pied Flycatchers

On average, migration of the Pied Flycatchers started on 27 August

(the earliest 3 August, the last 6 September). The average median date of the passage was 6 September. The average date of the end of migration was 26 September. The date of the last capture was 7 October. The peak period of migration fell on the first half of September in the years 1998-2007. The most captures occurred in September (Fig 122). The significant shift of the beginning of autumn migration towards earlier date was not revealed in the years 1998-2007, Pearson’s r=-0.41, p>0.05 (Fig. 123).

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91

0

1

2

3

4

5

6

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 122. Average daily captures and migration dynamics

(smoothed moving average) of Pied Flycatchers

26-Jul31-Jul5-Aug

10-Aug15-Aug20-Aug25-Aug30-Aug4-Sep9-Sep

1998 1999 2000 2001 2002 2003 2005 2006 2007

Fig. 123. Changes of date of the beginning of autumn migration of Pied Flycatchers

30-Jul 13-Aug27-Aug 10-Sep24-Sep 8-Oct 22-Oct 5-Nov 19-Nov

Fig. 124. Individual capture histories of Pied Flycatchers captured more than once within a season during the study period, 2006. Each point indicates a capture

of this bird and lines join the captures of the same individual.

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Proportion of recaptured birds was 7 % in 2006 and 2007. The minimum stopover time of the recaptured individuals was min. 2, max. 23 days (Fig. 124). 52 per cent of the Pied Flycatchers were captured in the forest, χ2=0, p>0.05 (Fig. 125).

Forest

Bushy

Grassland

Fig. 125. Capture rate (%) of Pied Flycatchers in the four habitats, 2007

Aegithalidae

85. Long-tailed Tit (Aegithalos caudatus) The Long-tailed Tit was subdominant songbird in the study area. In total, 740 individuals of Long-tailed Tits were marked between 1998 and 2007. The most birds were captured in 2000, the least in 1998. The average daily captures did not change significantly, Kruskal-Wallis test, H=6.37, p>0.05 (Fig. 126).

0102030405060708090

100110120130

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 126. Yearly capture numbers of Long-tailed Tits

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On average, migration of the Long-tailed Tits started on 28

September (the earliest 15 August, the last 5 November). The average median date of the passage was 22 October. The average date of the end of migration and the last capture was 10 November. The peak period of migration fell on the early November in the years 1998-2007. The most captures occurred in October (Fig. 127). Proportion of recaptured birds was 24-34 % in 2004 and 2006. The minimum stopover time of the recaptured individuals was min. 2, max. 23 days (Fig. 128). Some birds probably overwintered in the study area. 51 per cent of the Long-tailed Tits were captured in the grassland, χ2=0, p>0.05 (Fig. 129).

0

1

2

3

4

5

6

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 127. Average daily captures and migration dynamics

(smoothed moving average) of Long-tailed Tits

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov Fig. 128. Individual capture histories of Long-tailed Tits captured more than once within a season during the study period, 2006. Each point indicates a capture of

this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 129. Capture rate (%) of Long-tailed Tits in the four habitats, 2000

Paridae

86. Marsh Tit (Parus palustris)

The Marsh Tit was accessory songbird in the study area. In total, 217 individuals of Marsh Tits were marked between 1998 and 2007. The most birds were captured in 2000, the least in 1999. The average daily captures changed significantly, Kruskal-Wallis test, H=31.08, p<0.05 (Fig. 130).

0

10

20

30

40

50

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 130. Yearly capture numbers of Marsh Tits

The average median date of the passage was 10 September. The

average date of the end of movement was 4 November. The date of the last capture was 6 November. The peak period of passage fell on the early October in the years 1998-2007. The most captures occurred in October (Fig 131). Proportion of recaptured birds was 14% in 2000, 42-54% in 2005 and 2007 (Fig. 132). 46 per cent of the Marsh Tits were captured in the grassland, χ2=0, p>0.05 (Fig. 133).

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95

0

1

2

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 131. Average daily captures and migration dynamics

(smoothed moving average) of Marsh Tits

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

Fig. 132. Individual capture histories of Marsh Tits captured more than once within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Fig. 133. Capture rate (%) of Marsh Tits in the four habitats, 2000

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96

Table 17. Data for the Marsh Tit ringed at the other Hungarian ringing site and recovered in Tömörd

Capture Type

Age/Sex Date Place Coordinates Text

Distancekm

Direction degree

Elapsed days

Ringing

juvenile

10.08.2000. Szőce, Őrség Vas County

46°53'N 16°34'E

Recovery adult 22.08.2000. Tömörd (AH) 47°22'N 16°41'E 55 9 12

87. Willow Tit (Parus montanus) The Willow Tit was very rare songbird in the study area. In total, 5 individuals of Willow Tits were marked between 1998 and 2007. The date of the captures: 26 September 2000, 2 and 26 October 2004, 3 October 2007 (2 indiv.). 88. Crested Tit (Parus cristatus) One juvenile bird of this species was captured on 17 September 2007 in the bushy. 89. Coal Tit (Parus ater) The Coal Tit was accessory songbird in the study area. In total, 185 individuals of Coal Tits were marked between 1998 and 2007. The most birds (86 indiv.) were captured in 2005. There were no captures in 1998 and 2003. The most birds were captured at the end of September and October. The date of the earliest capture was 23 August; the date of the last capture was 9 November. One bird ringed in Slovenia recovered in Tömörd (Table 18).

Table 18. Data for the Coal Tit ringed abroad and recovered in Tömörd Capture

Type Age Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing pullus 10.06.2005.

Mrzli studenec, Pokljuka (Triglav

National Park), Slovenia

46°21'N 13°59'E

Recovery adult 01.10.2005. Tömörd (AH) 47°21'23"N 16°40'04"E 233 61 113

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90. Blue Tit (Parus caeruleus) The Blue Tit was dominant songbird in the study area. In total, 3429 individuals of Blue Tits were marked between 1998 and 2007. The most birds were captured in 2004, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=78.84, p<0.0001 (Fig 134). More than 80% of ringed birds were juvenile in 2004-2006 (Fig. 136).

0100200300400500600700800900

1000

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 134. Yearly capture numbers of Blue Tits

On average, migration of the Blue Tits started on 25 August (the earliest 9 August, the last 1 September). The average median date of the passage was 2 October. The average date of the end of migration was 8 November. The date of the last capture was 11 November. The peak period of migration fell on the end of September and early October in the years 1998-2007. The most captures occurred in October (Fig. 136). Proportion of recaptured birds was 5% in 2004, 13% in 2005 and 15% in 2006. 62 per cent of the recaptured birds were male (Fig. 137). According to the distribution of recaptures, the local breeding birds captured in August stayed until early November and probably overwintered at the study area. The most of birds arrived in the peak migration period left the area quickly by the end of October (Fig. 138). 68 per cent of the Blue Tits were captured in the grassland, χ2=2, p>0.05 (Fig. 139). The migratory Blue Tits probably followed SSW direction from Tömörd and wintered at the Adriatic region (Table 19).

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98

0102030405060708090

2004 2005 2006

%

juvenile

adult

Fig. 135. Ratio of juvenile and adult Blue Tits

02468

101214161820

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 136. Average daily captures and migration dynamics

(smoothed moving average) of Blue Tits

Fig. 137. Ratio of the recaptured males and females of Blue Tits

38%

62%

male female

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29-Jul 12-Aug 26-Aug 9-Sep 23-Sep 7-Oct 21-Oct 4-Nov 18-Nov

Fig. 138. Individual capture histories of Blue Tits captured more than once within a season during the study period, 2004. Each point indicates a capture of this bird

and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 139. Capture rate (%) of Blue Tits in the four habitats, 2004

Table 19. Data for all the Blue Tits ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing adult 28.10.2002. Tömörd

(AH) 47°21'23"N 16°40'04"E

1. Recovery adult 01.01.2003.

Völcsej Győr-

Moson-Sopron County

47°30'N 16°46'E 18 25 65

Ringing juvenile male 05.10.2005. Tömörd

(AH) 47°21'23"N 16°40'04"E

2. Recovery juvenile

male 11.10.2005. Ócsa (AH) Pest County

47°17'50"N 19°12'40"E 192 92 6

Ringing juvenile 20.07.2002. Szombathely 47°14'N 16°38'E

3. Recovery juvenile 18.08.2002. Tömörd

(AH) 47°21'23"N 16°40'04"E 14 11 29

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100

Ringing juvenile female 21.10.2004. Tömörd

(AH) 47°21'23"N 16°40'04"E

4. Recovery adult 07.01.2006.

Vransko Jezero,

Pakoštane, Croatia

43°56'57"N 15°30'57"E 390 193 443

Ringing juvenile 16.10.2000. Tömörd (AH)

47°21'23"N 16°40'04"E

5. Recovery adult

male 04.03.2001. Sombor Yugoslavia

45°46'N 19°07'E 258 133 139

91. Great Tit (Parus major) The Great Tit was dominant songbird in the study area. In total, 2843 individuals of Great Tits were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=67.99, p<0.0001 and the yearly captures showed significant increasing trend from 1998 to 2007 (Fig 140). Significant positive correlation was found between the yearly numbers of males and females (Fig. 141). Significant negative correlation was found between the ratio of first-year birds (production) and the number of adults in a specific year (Fig. 142).

On average, migration of the Great Tits started on 26 September (the earliest 1 August, the last October). The average median date of the passage was 13 October. The average date of the end of migration was 8 November. The date of the last capture was 16 November. The peak period of migration fell on mid-October in the years 1998-2007. The most captures occurred in October (Fig. 144).

0

100

200

300

400

500

600

700

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 140. Yearly capture numbers of Great Tits

Linear regression of yearly captures, r=0.65, p<0.05

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101

100 200 300No. of females

100

200

300

No.

of m

ales

Fig. 141. Linear regression of yearly captures of females and males,

r=0.95, p<0.001

100 200No. of adults

2

3

4

5

6

7

ratio

juve

milis

/adu

lts

Fig. 142. Linear regression of yearly captures of adults and juveniles,

r=-0.68, p<0.05 64

19 143 0

010203040506070

1 2 3 4 5year from ringing

%

Fig. 143. Ratio of recaptured Great Tits

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Proportion of recaptured birds was 9% in 2004, 15% in 2005 and 22% in 2006. According to the distribution of recaptures, the local breeding birds captured in May and August stayed until early November and probably overwintered at the study area (Fig. 145). 65 per cent of birds recaptured in November were male (Fig. 146). The most of birds arrived in the peak migration period left the area quickly by the end of October. 45 per cent of the Great Tits were captured in the grassland, χ2=0, p>0.05 (Fig. 147). Some migratory Great Tits probably wintered in Slovenia and Croatia (Table 20).

02468

10121416

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 144. Average daily captures and migration dynamics

(smoothed moving average) of Great Tits

1-May

15-May

29-May

12-Jun

26-Jun

10-Jul

24-Jul

7-Aug

21-Aug

4-Sep

18-Sep

2-Oct

16-Oct

30-Oct

13-Nov

Fig. 145. Individual capture histories of Great Tits captured more than once within a season during the study period, 2004. Each point indicates a capture of this bird

and lines join the captures of the same individual.

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103

11

31

24

34

05

10152025303540

female male female male

adult juvenile

ratio

of r

ecap

ture

s %

Fig. 146. Ratio of sex and age groups of recaptured Great Tits in November

Forest

Bushy

Grassland

Marsh

Fig. 147. Capture rate (%) of Great Tits in the four habitats, 2005

Table 20. Data for all the Great Tits ringed at the other Hungarian ringing sites

and recovered in Tömörd and also those ringed in Tömörd and recovered abroad Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing

juvenile

10.06.2004.SzelcepusztaHajdu-Bihar

County

48°31'N 20°36'E

1.

Recovery juvenile female 17.10.2004. Tömörd

(AH) 47°21'23"N 16°40'04"E 321 246 129

Ringing juvenile female 23.10.2005. Tömörd

(AH) 47°21'23"N 16°40'04"E

2. Recovery adult 16.12.2007. Zidani Most,

Slovenia 46°05'N 15°10'E 182 219 784

Ringing juvenile 26.09.2007. Tömörd (AH)

47°21'23"N 16°40'04"E

3. Recovery juvenile 11.10.2007.

Brezno Donje, Croatia

46°11'N 15°39'E 152 211 15

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Sittidae 92. Nuthatch (Sitta europaea)

The Nuthatch was accessory songbird in the study area. In total, 131 individuals of Nuthatchs were marked between 1998 and 2007. The most birds were captured in 2001, the least in 1998. The average daily captures did not change significantly, Kruskal-Wallis test, H=2.08, p>0.05 (Fig. 148). According to distribution of daily captures and recaptures there were no migratory birds from other areas and the local birds stayed at the study area from August to November (Fig. 149, 150). 69 per cent of the Nuthatchs were captured in the forest, χ2=6, p<0.05 (Fig. 151).

0

5

10

15

20

25

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 148. Yearly capture numbers of Nuthatchs

0

1

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 149. Average daily captures of Nuthatchs

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105

29-Jul

12-Aug

26-Aug

9-Sep

23-Sep

7-Oct

21-Oct

4-Nov

18-Nov

Fig. 150. Individual capture histories of Nuthatchs captured more than once within a season during the study period, 2006. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 151. Capture rate (%) of Nuthatchs in the four habitats, 2001

Certhiidae 93. Treecreeper (Certhia familiaris) The Treecreeper was accessory songbird in the study area. In total, 110 individuals of Treecreeper were marked between 1998 and 2007. The most birds were captured in 2000, the least in 1998. The average daily captures did not change significantly, Kruskal-Wallis test, H=7.99, p>0.05 (Fig. 152).

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106

0

5

10

15

20

25

30

35

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 152. Yearly capture numbers of Treecreepers

On average, migration of the Treecreepers started on 17 September (the earliest 5 August, the last 25 October). The average median date of the passage was 13 October. The date of the last capture was 6 November. The most captures occurred in October (Fig. 153).

0

1

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 153. Average daily captures and migration dynamics

(smoothed moving average) of Treecreepers

Proportions of recaptured birds were 20% in 2000 and 50% in 2006. According to the distribution of recaptures, the local breeding birds captured in August left the study area by mid-September. The most of birds arrived in the end of September and October stayed until mid-November at the study area (Fig. 154). 64 per cent of the Treecreepers were captured in the bushy, χ2=2, p>0.05 (Fig. 155).

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107

29-Jul

12-Aug

26-Aug

9-Sep

23-Sep

7-Oct

21-Oct

4-Nov

18-Nov

Fig. 154. Individual capture histories of Treecreepers captured more than once within a season during the study period, 2006. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Fig. 155. Capture rate (%) of Treecreepers in the four habitats, 2000

94. Short-toed Treecreeper (Certhia brachydactyla)

The Short-toed Treecreeper was accessory songbird in the study area. In total, 117 individuals of Short-toed Treecreeper were marked between 1998 and 2007. The most birds were captured in 2007, the least in 1998 and 1999. The average daily captures did not change significantly,

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Kruskal-Wallis test, H=2.87, p>0.05, but the yearly captures showed significant increasing trend from 1998 to 2007 (Fig. 156).

0

5

10

15

20

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 156. Yearly capture numbers of Short-toed Treecreepers.

Linear regression of yearly captures, r=0.65, p<0.05

On average, migration of the Short-toed Treecreepers started on 28 September (the earliest 11 September, the last 18 October). The average median date of the passage was 12 October. The date of the last capture was 10 November. The most captures occurred in September (Fig. 157). Proportions of recaptured birds were 26% in 2004 and 50% in 2007. According to the distribution of recaptures, the local breeding birds captured in July and August left the study area by mid-September. The most of birds arrived in September stayed until mid-October at the study area (Fig. 158). 70 per cent of the Short-toed Treecreepers were captured in the grassland, χ2=2, p>0.05 (Fig. 159).

0

1

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 157. Average daily captures and migration dynamics (smoothed moving average) of Short-toed Treecreepers

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27-Jul 16-Aug 5-Sep 25-Sep 15-Oct 4-Nov

Fig. 158. Individual capture histories of Short-toed Treecreepers captured more than once within a season during the study period, 2004. Each point indicates

a capture of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassaland

Fig. 159. Capture rate (%) of Short-toed Treecreepers in the four habitats, 2007

Remizidae

95. Penduline Tit (Remiz pendulinus) The Penduline Tit was rare songbird in the study area. In total, 22 individuals of Penduline Tits were marked between 1998 and 2007. The most birds (11 indiv.) were captured in 2000. There were no captures between 2002 and 2006. The date of the earliest capture was 2 August 2007; the date of the last capture was 11 November. Some Penduline Tits migrated above Tömörd originate from the Baltic region (Table 21).

Table 21. Data for the Penduline Tit ringed in Tömörd and recovered abroad

Capture Type

Age/Sex Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing adult female 15.10.1999. Tömörd (AH) 47°21'23"N

16°40'04"E

Recovery adult 07.04.2001. Kalnciems, Litvania

56°49'N 23°38'E 1155 24 540

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Oriolidae 96. Golden Oriole (Oriolus oriolus) The Golden Oriole was rare songbird among the captured species but it was not rare in the study area. In total, 2 individuals of Golden Orioles were marked between 1998 and 2007. The date of captures: 18 August 1998, 2 September 2004.

Laniidae 97. Red-backed Shrike (Lanius collurio)

The Red-backed Shrike was subdominant songbird in the study area. In total, 693 individuals of Red-backed Shrikes were marked between 1998 and 2007. The most birds were captured in 2004, the least in 1999. The average daily captures changed significantly, Kruskal-Wallis test, H=17.54, p<0.05, and the yearly captures showed a significant increasing trend from 1998 to 2007 (Fig. 160).

On average, migration of the Red-backed Shrikes started on the 20th of August (the earliest 1 August, the last 31 August). The average median date of the passage was 4 August for females, 6 August for males and 19 August for first year birds (Fig. 161). The average date of the end of migration was 22 September. The date of the last capture was 6 October. The peak period of migration fell on the first half of August in the years 1998-2007. The most captures occurred in August (Fig. 162).

0

20

40

60

80

100

120

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 160. Yearly capture numbers of Red-backed Shrikes

Linear regression of yearly captures, r=0.67, p<0.05

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111

0102030405060708090

100

27-Jul

3-Aug

10-Aug

17-Aug

24-Aug

31-Aug

7-Sep

14-Sep

21-Sep

28-Sep

5-Oct

daily

cap

ture

s %

malefemalejuvenile

Fig. 161. Cumulative migration curves, 1998-2004

0

1

2

3

4

5

6

7

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 162. Average daily captures and migration dynamics

(smoothed moving average) of Red-backed Shrikes

Proportion of recaptured birds was 27% in 2004 and 8% in 2005 and 32 % in 2006. According to the distribution of recaptures, the local breeding birds captured in July and early August left the study area by mid-August (Fig 163). The average minimum stopover period was 9 days, the minimum was 2, and the maximum was 44 days. The majority of the Red-backed Shrikes captured at Tömörd had a small (<3) migrating fat supply before continuing their migration. 74 per cent of the Red-backed Shrikes were captured in the grassland, χ2=2, p<0.05 (Fig. 164).

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29-Jul 12-Aug

26-Aug

9-Sep 23-Sep

7-Oct 21-Oct

4-Nov 18-Nov

29-Jul 12-Aug

26-Aug

9-Sep 23-Sep

7-Oct 21-Oct

4-Nov 18-Nov

Fig. 163. Individual capture histories of Red-backed Shrikes captured more than

once within a season during the study period. Each point indicates a capture of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 164. Capture rate (%) of Red-backed Shrikes in the four habitats, 2004

2004

2005

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98. Great Grey Shrike (Lanius excubitor) The Great Grey Shrike was rare songbird in the study area. In total, 18 individuals of Great Grey Shrikes were marked between 1998 and 2007. The most birds (4 indiv.) were captured in 2004. The most birds were captured in October. The date of earliest capture was 20 September 2006; the date of the last capture was 4 November.

Corvidae

99. Jay (Garrulus glandarius)

The Jay was accessory songbird in the study area. In total, 124 individuals Jays were marked between 1998 and 2007. The most birds were captured in 2002, there was no capture in 1998 (Fig. 165). The peak period of migration fell on the end of September in the years 1998-2007. The most captures occurred in September (Fig. 166).

0

10

20

30

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 165. Yearly capture numbers of Jays

0

1

2

31-Jul

14-Aug

28-Aug

11-Sep

25-Sep

9-Oct

23-Oct

6-Nov

No.

of b

irds

Fig. 166. Average daily captures and migration dynamics

(smoothed moving average) of Jays

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Sturnidae 100. Starling (Sturnus vulgaris) The Starling was rare songbird among the captured species but it was not rare in the study area. In total, 90 individuals of Starlings were marked between 1998 and 2007. The most birds (44 indiv.) were captured in 2007. The most birds were captured at the end of July and early August. Some Starlings migrated above Tömörd wintered in Italy (Table 22).

Table 22. Data for the Starling ringed in Tömörd and recovered abroad Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile female 27.10.2005. Tömörd (AH) 47°21'23"N

16°40'04"E

Recovery - 15.12.2005. Portazzolo,

Sale Marasino, Italy

45°42'N 10°07'E 535 250 49

Passeridae 101. House Sparrow (Passer domesticus) The House Sparrow was rare songbird in the study area. In total, 14 individuals of House Sparrows were marked between 1998 and 2007. The most birds (8 indiv.) were captured in 2003. 102. Tree Sparrow (Passer montanus)

The Sparrow was dominant songbird in the study area. In total, 3301 individuals of Sparrows were marked between 1998 and 2007. The most birds were captured in 2003, the least in 2007. The average daily captures changed significantly, Kruskal-Wallis test, H=32.05, p<0.001 (Fig. 167). The most daily captures were in the post-breeding dispersal. The most captures occurred in August (Fig. 168). Proportions of recaptured birds were 9% in 2004 and 5% in 2005 and 4 % in 2006. Some birds originated from Austria and wintered at the study area (Fig. 169, Table 23). 53 per cent of the Sparrows were captured in the grassland, χ2=2, p<0.05 (Fig. 170).

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115

0100200300400

500600700800

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 167. Yearly capture numbers of Sparrows

02468

101214161820

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 168. Average daily captures (smoothed moving average) of Sparrows

31-Jul 20-Aug 9-Sep 29-Sep 19-Oct 8-Nov

Fig. 169. Individual capture histories of Sparrows captured more than once within a season during the study period, 2005. Each point indicates a capture of this bird

and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 170. Capture rate (%) of Sparrows in the four habitats, 2005

Table 23. Data for all the Sparrows ringed in Tömörd at the other Hungarian

ringing site and those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile 04.08.2003. Tömörd (AH)

47°21'23"N 16°40'04"E

1. Recovery adult 23.04.2004. Csepreg 47°24'N

16°43'E 6 36 263

Ringing

juvenile

06.09.2002.

Hohenau, Ringelsdorf,

March, Austria

48°35'N 16°55'E

2.

Recovery adult 28.10.2002. Tömörd (AH)

47°21'23"N 16°40'04"E 138 188 52

Fringillidae

103. Chaffinch (Fringilla coelebs)

The Chaffinch was dominant songbird in the study area. In total, 1108 individuals of Chaffinchs were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=138.7, p<0.0001 (Fig. 171).

On average, migration of the Chaffinch started on the 4th of October (the earliest 1 September, the last 23 October). The average median date of the passage was 14 October. The average date of the end of migration was 7 November. The date of the last capture was 18 November. The peak period of migration fell on first half of October in the years 1998-2007. The most captures occurred in October (Fig. 172).

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0100200300400

500600700800

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 171. Yearly capture numbers of Chaffinchs

0123456789

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 172. Average daily captures and migration dynamics

(smoothed moving average) of Chaffinchs

29-Jul 18-Aug 7-Sep 27-Sep 17-Oct 6-Nov

Fig. 173. Individual capture histories of Chaffinchs captured more than once within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

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Proportion of recaptured birds was 1.5% in 2005. According to the distribution of recaptures, the local breeding birds captured in July and early August left the study area by mid-October (Fig. 173). The longest minimum stopover period was 57 days. 79 per cent of the Chaffinchs were captured in the bushy, χ2=6, p<0.05 (Fig. 174). Most of Chaffinchs migrated above Tömörd probably wintered in Italy (Table 24).

Forest

Bushy

Grassland

Marsh

Fig. 174. Capture rate (%) of Chaffinchs in the four habitats, 2005

Table 24. Data for all the Chaffinchs ringed in Tömörd recovered abroad

Capture Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing adult female 04.11.2001. Tömörd

(AH) 47°21'23"N 16°40'04"E

1.

Recovery adult 11.12.2003.

Monaciano, Castelnuovo Berardenga,

Italy

43°21'N 11°21'E 610 223 767

Ringing adult male 23.09.2005. Tömörd

(AH) 47°21'23"N 16°40'04"E

2.

Recovery adult 29.10.2005. Amandola, Italy

42°59'N 13°21'E 552 208 36

Ringing juvenile female 08.10.2005. Tömörd

(AH) 47°21'23"N 16°40'04"E

3.

Recovery adult female 29.10.2005.

Pranovi, Trissino,

Italy

45°33'N 11°22'E 453 244 21

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Ringing juvenile male 06.11.2005. Tömörd

(AH) 47°21'23"N 16°40'04"E

4. Recovery adult 07.12.2005.

Corcironda, Zermeghedo,

Italy

45°29'N 11°23'E 456 243 31

104. Brambling (Fringilla montifringilla) The Brambling was subdominant and irruption songbird in the study area. In total, 766 individuals of Bramblings were marked between 1998 and 2007. The most birds (708 indiv.) were captured in 2005. There was no capture in 2003 (Fig. 175). The migration of the Brambling started on the 8th of October in 2005. The peak period of migration fell in the mid-October in 2005 (Fig. 176). The date of earliest capture was 25 September 2005 and there were 11 (1.5%) recaptures after 23 October 2005. 92 per cent of the Bramblings were captured in the bushy, χ2=6, p<0.05 (Fig. 177).

0100

200300400

500600

700800

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 175. Yearly capture numbers of Bramblings

01020304050607080

1-Aug

15-Aug

29-Aug

12-Sep

26-Sep

10-Oct

24-Oct

7-Nov

No.

of b

irds

Fig. 176. Average daily captures and migration dynamics

(smoothed moving average) of Bramblings

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Forest

Bushy

Grassland

Marsh

Fig. 177. Capture rate (%) of Bramblings in the four habitats, 2005

105. Serin (Serinus serinus) The Serin was rare songbird in the study area. In total, 50 individuals of Serins were marked between 1998 and 2007. The most birds (13 indiv.) were captured in 2005. There was no capture in 2000. The date of earliest capture was 29 July; the date of the last capture was 31 October.

106. Greenfinch (Carduelis chloris) The Greenfinch was dominant songbird in the study area. In total, 1467 individuals of Greenfinchs were marked between 1998 and 2007. The most birds were captured in 2005, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=156.3, p<0.0001 (Fig. 178).

0100200300400500600700800

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 178. Yearly capture numbers of Greenfinchs

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On average, migration of the Greenfinch started on the 4th of September (the earliest 31 July, the last 18 October). The average median date of the passage was 2 October. The average date of the end of migration was 1 November. The date of the last capture was 19 November. The peak period of migration fell on the end of August and early September in the years 1998-2007. The most captures occurred in September (Fig. 179). Proportion of recaptured birds was 2% in 2005. According to the distribution of recaptures, the local breeding birds captured in July and early August left the study area by mid-October (Fig. 180). The longest minimum stopover period was 60 days. 68 per cent of the Greenfinchs were captured in the bushy, χ2=2, p>0.05 (Fig. 181). Some Greenfinchs migrated above Tömörd probably wintered in Croatia (Table 25).

0

2

4

6

8

10

12

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 179. Average daily captures and migration dynamics

(smoothed moving average) of Greenfinchs

Table 25. Data for all the Greenfinchs ringed in Tömörd and recovered at the other Hungarian ringing site or abroad

Capture Type Age Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing juvenile 03.09.2004. Tömörd (AH) 47°21'23"N 16°40'04"E

1. Recovery adult 17.04.2005. Völcsej

Hungary 47°30'N 16°46'E 18 25 226

Ringing juvenile 27.08.2005. Tömörd (AH) 47°21'23"N 16°40'04"E

2. Recovery adult 18.10.2005.

Draževitići, Vrgorac, Croatia

43°09'N 17°24'E 472 173 52

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122

27-Jul 16-Aug 5-Sep 25-Sep 15-Oct 4-Nov 24-Nov

Fig. 180. Individual capture histories of Greenfinchs captured more than once within a season during the study period, 2005. Each point indicates a capture

of this bird and lines join the captures of the same individual.

Forest

Bushy

Grassland

Marsh

Fig. 181. Capture rate (%) of Greenfinchs in the four habitats, 2005

107. Goldfinch (Carduelis carduelis) The Goldfinch was rare songbird in the study area. In total, 52 individuals of Goldfinchs were marked between 1998 and 2007. The most birds (15 indiv.) were captured in 2001. There was no capture in 1999. The date of earliest capture was 12 August; the date of the last capture was 11 November. 108. Siskin (Carduelis spinus) The Siskin was rare songbird in the study area. In total, 40 individuals of Siskins were marked between 1998 and 2007. The most

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birds (23 indiv.) were captured in 2004. There was no capture in 1998, 1999, 2001, 2003 and 2006. The date of earliest capture was 6 September; the date of the last capture was 8 November. 109. Linnet (Carduelis cannabina) The Linnet was accessory songbird in the study area. In total, 116 individuals of Linnets were marked between 1998 and 2007. The most birds (53 indiv.) were captured in 2002. There was 1-1 bird captured in 1998, 1999 and 2000. The date of earliest capture was 28 July; the date of the last capture was 5 November. 110. Redpoll (Carduelis flammea) One juvenile bird of this species was captured on the 20th of September, 2000 in the grassland. 111. Crossbill (Loxia curvirostra) One juvenile female bird of this species was captured on the 6th of September, 2002 in the marsh. 112. Scarlet Rosefinch (Carpodacus erythrinus) One juvenile bird of this species was captured on the 17th of August, 2003 in the grassland. 113. Bullfinch (Pyrrhula pyrrhula)

The Bullfinch was accessory songbird in the study area. In total, 225 individuals of Bullfinchs were marked between 1998 and 2007. The most birds were captured in 2004, there was no capture in 1998. The average daily captures did not change significantly, Kruskal-Wallis test, H=8.49, p>0.05 (Fig. 182). On average, migration of the Bullfinch started on the 28th of October (the earliest 24 October, the last 6 November). The average median date of the passage was 6 November. The date of earliest capture was 6 August; the date of the last capture was 19 November. The most captures occurred in November (Fig. 183). There were 11 (6%) recaptures after 21 October 2004 (Fig. 184). 66 per cent of the Hawfinchs were captured in the bushy, χ2=2, p>0.05 (Fig. 185).

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0

20

40

60

80

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 182. Yearly capture numbers of Bullfinchs

0

1

2

3

4

5

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 183. Average daily captures and migration dynamics

(smoothed moving average) of Bullfinchs

1-Aug 21-Aug 10-Sep 30-Sep 20-Oct 9-Nov

Fig. 184. Individual capture histories of Bullfinchs captured more than once within a season during the study period, 2004. Each point indicates a capture of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 185. Capture rate (%) of Bullfinchs in the four habitats, 2004

114. Hawfinch (Coccothrausthes coccothrausthes)

The Hawfinch was accessory songbird in the study area. In total, 403 individuals of Hawfinchs were marked between 1998 and 2007. The most birds were captured in 2004, there was no capture in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=240.4, p<0.0001 (Fig. 186).

0

50

100

150

200

250

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 186. Yearly capture numbers of Hawfinchs

On average, migration of the Hawfinch started on the 22nd of

September (the earliest 3 August, the last 29 October). The average median date of the passage was 23 October. The date of the last capture was 16 November. The peak period of migration fell on the end of October the

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years 1998-2007. The most captures occurred in October (Fig. 187). Proportion of recaptured birds was 3% in 2005. According to the distribution of recaptures, the local breeding birds captured in July and early August left the study area until the mid-October (Fig. 188). The longest minimum stopover period was 81 days. 77 per cent of the Hawfinchs were captured in the bushy, χ2=2, p>0.05 (Fig. 189). Some Hawfinchs migrated above Tömörd originated from Poland (Table 26).

0

1

2

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov 16-Nov

No.

of b

irds

Fig. 187. Average daily captures and migration dynamics

(smoothed moving average) of Hawfinchs

31-Jul 20-Aug 9-Sep 29-Sep 19-Oct 8-Nov

Fig. 188. Individual capture histories of Hawfinchs captured more than once within a season during the study period, 2005. Each point indicates a capture of this bird and lines join the captures of the same individual.

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Forest

Bushy

Grassland

Marsh

Fig. 189. Capture rate (%) of Hawfinchs in the four habitats, 2005

Table 26. Data for all the Hawfinchs ringed in Tömörd and recovered in abroad

Capture Type Age/Sex Date Place Coordinates

Text Distance

km Direction

degree Elapsed

days

Ringing adult female 26.10.2005. Tömörd (AH) 47°21'23"N

16°40'04"E

Recovery adult female 17.02.2008.

Poreba Wielika, Malopolskie,

Poland

49°37'N 20°04'E 356 45 844

Emberizidae

115. Yellowhammer (Emberiza citrinella)

The Yellowhammer was dominant songbird in the study area. In total, 1623 individuals of Yellowhammers were marked between 1998 and 2007. The most birds were captured in 2003, the least in 1998. The average daily captures changed significantly, Kruskal-Wallis test, H=73.35, p<0.001 (Fig. 190).

On average, migration of the Yellowhammer started on the 16th of September (the earliest 25 August, the last 15 October). The average median date of the passage was 15 October. The average date of the end of migration was 10 November. The date of the last capture was 19 November. The peak period of migration fell on the second half of September the years 1998-2007. The most captures occurred in September (Fig. 191).

Proportion of recaptured birds was 3% in 2005. According to the distribution of recaptures, some local breeding birds captured in April and May probably were wintering at the study area (Fig. 193). The longest minimum stopover period was 89 days. 41 per cent of the Yellowhammers were captured in the grassland, χ2=2, p>0.05 (Fig. 192).

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0

100

200

300

400

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 190. Yearly capture numbers of Yellowhammers

0123456789

10

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 191. Average daily captures and migration dynamics

(smoothed moving average) of Yellowhammers

Forest

Bushy

Grassland

Marsh

Fig. 193. Capture rate (%) of Yellowhammers in the four habitats, 2003

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26-Apr

10-May

24-May

7-Jun

21-Jun

5-Jul

19-Jul

2-Aug

16-Aug

30-Aug

13-Sep

27-Sep

11-Oct

25-Oct

8-Nov

29-Jul 18-Aug 7-Sep 27-Sep 17-Oct 6-Nov Fig. 193. Individual capture histories of Yellowhammers captured more than once

within a season during the study period. Each point indicates a capture of this bird and lines join the captures of the same individual.

116. Ortolan Bunting (Emberiza hortulana) One juvenile bird of this species was captured on the 20th of September, 2000 in the grassland. 117. Reed Bunting (Emberiza schoeniclus)

The Reed Bunting was subdominant songbird in the study area. In total, 826 individuals of Reed Buntings were marked between 1998 and 2007. The most birds were captured in 2004, the least in 2002. The average daily captures changed significantly, Kruskal-Wallis test, H=40.18, p<0.01 (Fig. 194).

2004

2006

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On average, migration of the Reed Bunting started on 10 October (the earliest 25 September, the last 18 October). The average median date of the passage was 24 October. The average date of the end of migration was 18 November. The date of the last capture was 18 November. The peak period of migration fell on the mid-October the years 1998-2007. The most captures occurred in October (Fig. 195).

There were only few recaptured birds per year. There was no capture in the forest and the 49 per cent of the Reed Buntings were captured in the marsh, χ2=4, p<0.05 (Fig. 196). Most of Reed Buntings migrated above Tömörd probably wintered in Italy (Table 27).

020

406080

100120

140160

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 194. Yearly capture numbers of Reed Buntings

0

1

2

3

4

5

6

7

27-Jul

10-Aug

24-Aug

7-Sep

21-Sep

5-Oct

19-Oct

2-Nov

16-Nov

No.

of b

irds

Fig. 195. Average daily captures and migration dynamics

(smoothed moving average) of Reed Buntings

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Bushy

Grassland

Marsh

Fig. 196. Capture rate (%) of Reed Buntings in the four habitats, 2004

Table 27. Data for all the Reed Buntings ringed in Tömörd and recovered in

abroad and those ringed abroad and recovered in Tömörd Capture

Type Age/Sex Date Place CoordinatesText

Distancekm

Direction degree

Elapsed days

Ringing juvenile 05.10.2001. Tömörd (AH)

47°21'23"N 16°40'04"E

1. Recovery adult 08.12.2001.

Motta, Arcisate,

Italy

45°51'N 09°52'E 547 252 64

Ringing juvenile female 03.11.2004. Tömörd

(AH) 47°21'23"N 16°40'04"E

2. Recovery adult

female 19.12.2004.

Lago di Vico,

Caprarola, Italy

42°19'12"N 12°10'42"E 663 212 46

Ringing juvenile 09.07.2003.

Smrzov – Koclírov,

Czech Republic

49°05'N 14°41'E

3.

Recovery juvenile 10.10.2003. Tömörd (AH)

47°21'23"N 16°40'04"E 242 143 93

118. Corn Bunting (Emberiza calandra)

The Corn Bunting was accessory songbird in the study area. In total, 102 individuals of Corn Buntings were marked between 1998 and 2007. The most birds (51 indiv.) were captured in 2003. There was no capture in 2005 and 2007. The average daily captures changed significantly, Kruskal-Wallis test, H=91.51, p<0.0001(Fig. 197). The date of the last capture was 3 October. The most captures occurred in August in 2003 (Fig. 198).

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0

20

40

60

1998 2000 2002 2004 2006

No.

of b

irds

Fig. 197. Yearly capture numbers of Corn Buntings

0

1

2

3

4

5

27-Jul 10-Aug 24-Aug 7-Sep 21-Sep 5-Oct 19-Oct 2-Nov

No.

of b

irds

Fig. 198. Average daily captures and migration dynamics

(smoothed moving average) of Corn Buntings

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Summary and discussion From the 118 species ringed in Tömörd between 1998 and 2007 there were 14 dominant species (Fig. 199). These species, listed in order of decreasing occurrence were: Robin, Blackcap, Blue Tit, Chiffchaff, Tree Sparrow, Great Tit, Goldcrest, Blackbird, Yellowhammer, Greenfinch, Chaffinch, Swallow, Lesser Whitethroat, Whitethroat. Only the last three species migrated over the Sahara (CRAMP 1998). 71 per cent of all of the individuals ringed during the work were dominant passerine birds.

010203040

50607080

dominant subdominant accessory rare

No.

of s

peci

es

Fig. 199. Number of species ringed according to dominance categories

The number of birds ringed in the post-breeding dispersal and migration seasons in the study area around Tömörd showed considerable yearly fluctuations between 1998 and 2007. The yearly numbers showed an increasing trend for Wryneck, Dunnock, Robin, Song Thrush, Garden Warbler, Blackcap, Chiffchaff, Short-toed Treecreeper, Red-backed Shrike and Great Tit between 1998 and 2007. The yearly number for Grasshopper Warbler showed a negative trend between 2003 and 2007. There was a Blue Tit and Bullfinch invasion in 2004. The number of Chaffinchs, Bramblings, Hawfinchs and Greenfinchs ringed in 2005 was the highest because of the high amount of helianthus crop on the fields around the study area in that year. The number of Sparrows ringed in 2003 was the highest because of the plenty of grain of wheat around Tömörd in that period. The marsh dried out in 2000 and 2001. For this reason the smallest number of Sedge Warblers, Marsh Warblers and Reed Warblers were ringed in those years.

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The migration curves based on the average number of individual species ringed daily between 1998 and 2007 showed that the temporal pattern of the migration of long-distance migratory species was different from that of the median- and short-distance migratory species. Birds could be classified into three distinct groups based on the migration dynamics characteristics of migration passage through the study area. The first group began the autumn migration as early as August and terminated it in early October. On average, the migration of five species, Swallow, Marsh Warbler, Whitethroat, Lesser Whitethroat and Thrush Nightingale started earliest in the first half of August. Most of trans-Sahara migratory birds left the study area by October. The migration of the second group commenced in the last part of August and September and might last until the end of October and early November (e.g. Wren, Dunnock, Robin, Redstart, Chiffchaff and Great Tit). The third group included the short-distance and vagrant species with migration commencement in October and migration termination by the end of October and early November (e.g. Blackbird, Song Thrush, Goldcrest and Reed Bunting). Based on the monthly data, October was the migration peak period for the three dominant and short-distance migratory species, Robin, Chiffchaff and Blue Tit in Tömörd. In the case of the long-distance migrants, the earlier and more intensive migration meant not only an adaptive migration strategy but resulted in the avoidance of the competition with the short-distance migrating species (BERTHOLD et al. 2003).. Sedge and Reed Warblers and other long-distance migrants migrated over Hungary in August and September taking advantage of the anticyclones which occur most intensively to the West of Carpathian basin or over the Carpathian basin. The most migration peak days of these species observed during the anticyclones. These weather situations ensured good conditions to them, because there were only a few clouds and a light northern wind, which facilitated the orientation and movement of the birds. The cold front of the cyclones which appeared in the Baltic region or over Ukraine also took a great effect on the autumn migration of some short-distance migrants, e.g. Robin, Blackcap, Chiffchaff, Goldcrest and Blue Tit (GYURÁCZ et al. 2003b). One day after the cold fronts great numbers of individuals were caught in Tömörd. During the cold front the strong wind was favorable only when its direction was suitable for the migration (back wind), because in that case flying did not needed much energy. Moreover, birds could reach higher migration speed and they could minimize the time of migration. The long-distance and short-distance partial migrants preferred anticyclones during their autumn migration, but there were some

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smaller differences in their migration patterns caused by the different behavior of the birds, because they followed different migration strategies to minimize the time and energy of the migration (BERTHOLD 1993). Among the four habitats, marsh habitat held species (e.g. Swallow, Sedge Warbler, Marsh Warbler and Reed Warbler) which were differed from those of the other habitats. The other species were located along a gradient from the forest to the grassland, matching to the pattern as the closeness and height of the vegetation was decreasing. At the end of the gradient we could find the species aggregating in the dense, high trees and bushes (e.g. Redwing, Nuthatch, Treeccreeper, Chaffinch, Brambling) and at the other end we could find the species grouping in the heterogeneous, bushy grassland (e.g. Whinchat, Whitethroat, Lesser Whitethroat, Willow Warbler, Red-backed Shrike). The species situated between the two ends of the gradient were distributed in a relatively even distribution (e.g. Wren, Chiffchaff, Garden Warbler and Blackcap). During the migration the habitat selection of birds had two main goals: to assure the energy resource required for the continuation of the migration and to avoid the predators. Both goals were served by the strategy to reach the next stopover site as well as the wintering place as soon as possible. The selection of the most appropriate stopover site was done by a third-level decision process including inherited and learned elements as well. The most important key-stimulus was the vegetation structure of the site, the quality and density of the nutrient resources (CODY 1985). During their stopover, some insectivorous species, e.g. the Whitethroat and Lesser Whitethroat chose the scrubs with high sugar content for rest and feeding. Therefore, these species grouped predominantly on more open, shrubby, grasslands that resembled to the savannah wintering areas. Other insectivorous species e.g. Sedge Warbler, Reed Warbler which fed on insects and spiders settled on plants standing in water. Therefore, these birds accumulated on watery, swampy habitats during their migration period. Among the short-distance migratory species, those wintering in the Mediterranean (e.g. Robin) as well as those wintering in Hungary (e.g. Brambling) were continuously present on the study area. The majority of the individuals were staying on the more closed bushy habitats. Their migration was slower, they were spending a longer time on the area than the trans-Sahara migrates (e.g. Willow Warbler) therefore avoidance of predators might play a greater role in the selection of rest and feeding sites. The forest and bushy habitats having a denser, closed vegetation structure provided a greater safety against the raptors.

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The migratory birds were maximizing the availability of the next stopover site by minimizing the predation i.e. by choosing the safety. The Bramblings for example, were choosing the feeding sites (fields) with bushy surroundings versus the open spaces because this choice offered more safety against the Sparrowhawk. According to these results, the migration dynamics and habitat selection of the long-distance migratory species exhibited essential differences compared to the short-distance migratory species. These differences were part of an adaptive strategy which was preserved from evolutionary changes. The results of bird ringing project of Tömörd Bird Observatory gave new information for population dynamics, migration route and habitat preference of some bird species migrating over Hungary. Information about connectivity of breeding, resting and wintering areas as well as yearly and daily numbers of populations is crucial information for any conservation (e.g. survey of biotope) and economic (e.g. planning of wind farm) action or widespread transport of disease (e.g. avian influenza). As migration pattern change over time, particularly in relation to factors such as climate change, continued bird ringing is important even for common species (KESTENHOLZ 2007). For this reason it would be extremely important to reveal further details about the migration strategy of birds and to get to know the breeding and wintering sites of birds migrating over Tömörd. Thus, we would like to continue our research.

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Acknowledgements We wish to express our gratitude to all those members of BirdLife Hungary who helped us in our field works. Staff: SZILÁRD SOMLAI (1998-99), JÁNOS EICHARDT (2000-01), HUBA HUSZÁR (2002-03), ALEX SZABOLCS (2004), NORBERT MÁTRAI (2005), HENRIETTA GABNAI (2006), ÉVA TARR (2007). Ringers: CSABA NÉMETH, GERGELY KIRÁLY, GÉZA SZENTENDREY, JÁNOS KISS, JÁNOS WISZTERCILL, JÓZSEF GÓCZÁN, LÁSZLÓ JÁNOSA, LÁSZLÓ VARGA, MIHÁLY LENCZL, PÉTER ILLÉS, PÉTER KOSZORÚS, PÉTER POLOVITZER, SÁNDOR TATAI. HELPERS: ANDREA HAFFNER, ANDREA RUFF, ÁGNES LEPOLD, BALÁZS HALGAS, BALÁZS KOVÁCS, CSILLA LŐRINCZ, IMRE LENDVAI, JÓZSEF HAFFNER, MÁRTON BÁNHIDI, TIBOR KELEMEN, TIBOR VIZSLÁN, TÍMEA NÉMETH, ZOLTÁN LEPOLD, ZOLTÁN SOMOGYI. The ringing work were supported by Environmental Ministry (KAC, KÖVICE), Scientific Board of Savaria Campus of University of West Hungary and National Civil Foundation of Hungary. J. GYURÁCZ was supported by the Békésy Fellowship and Hungarian Academy of Science (Regional Committee in Veszprém, MTA-VEAB). The publishing of this book was supported by Scientific Board of Savaria Campus of University of West Hungary and Hungarian Academy of Science (Regional Board in Szombathely, MTA-VATT). This book is part of the South-East Bird Migration Network and Actio Hungarica publications.

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References ALERSTAM, T. 1990. Bird Migration. Cambridge University Press.

Cambridge. BAIRLEIN, F. 1997. The European-African songbird migration network:

new chllenges fro large-scale study of bird migration. The 1st Conference of EOU, Bologna. Abstracts: 1.

BERTHOLD, P. 1993. Bird Migration. A general survey. Oxford University Press, Oxford.

BERTHOLD, P. GWINNER, E., SONNENSCHEIN, E. (eds.). 2003. Avian Migration. Springer-Verlag, Berlin.

BIRDLIFE INTERNATIONAL (2004): Birds in Europe: population estimates, trends and conservation status. BirdLife Conservation series 12. BirdLife International, Cambridge, UK.

BUSSE, P. 2000. Bird Station Manual. University of Gdansk. Gdansk. CODY, L. M. (ed.). 1985. Habitat Selection in Birds. Academic Press.

London. CRAMP, P. 1998. The Birds of the Western Palearctic. Oxford CD-ROM Oxford University Press, Oxford. GYURÁCZ J., GÓCZÁN J., BÁNHIDI P., LEPOLD Á. 2003a. Autumn Migration of the Goldcrest Regulus regulus in western Hungary. The RING 25, 1-2: 38-46. GYURÁCZ J., HORVÁTH G., CSÖRGŐ T., BANK L., PALKÓ S. 2003b.

Influence of the macrosynoptical weather situations on the autumn migration of birds. The RING 25, 1-2: 18-36.

GYURÁCZ J., BÁNHIDI P., GYIMÓTHY ZS. 2005. Habitat selection and migration dynamics of the migrating populations os Robins Erithacus rubecula in the autumn migration period. Alauda 73(3): 296-297. The 5th Conference of EOU, Strasbourg.

KESTENHOLZ, M. (comp.). 2007. Bird ringing in science and conservation. EURING, Heteren.

RAPPOLE, J. H., JONES, P. 2002. Evolution of old and new world migration system. Ardea 90. (3), Special Issue: 525-537. SVENSSON, L. 1992. Identification Guide to European Paserines.

Stockholm. SZENTENDREY G., LÖVEI G., KÁLLAY GY. 1979. Az "Actio Hungarica"

madárgyűrűző tábor mérési módszerei. Állattani Közlemények 66:161-166.

TIELEMAN, I. 2007. Physiological dynamics of birds in desert, tropical and temperate environments. The 6th Conference of EOU, Vienna. Abstract voulme: 9-10.

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Appendix Index and numbers of birds caught in different years at the Tömörd Bird Observatory

No. Page Species 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 Total 1. 12 Accipiter gentilis 0 0 0 1 1 0 0 0 0 2 4 2. 12 Accipiter nisus 0 0 0 1 1 3 1 3 1 2 12 3. 62 Acrocephalus arundinaceus 0 0 0 0 0 1 4 1 2 1 9 4. 56 Acrocephalus melanopogon 0 0 0 0 0 0 1 0 0 0 1 5. 58 Acrocephalus palustris 5 11 5 5 10 28 89 29 50 10 242 6. 56 Acrocephalus schoenobaenus 15 28 4 1 5 13 109 20 27 7 229 7. 60 Acrocephalus scirpaceus 4 5 2 2 3 8 25 17 23 4 93 8. 92 Aegithalos caudatus 27 64 124 109 70 51 58 67 86 84 740 9. 23 Alauda arvensis 0 0 0 1 0 0 1 0 0 0 2

10. 20 Alcedo atthis 1 0 0 0 0 0 0 1 0 1 3 11. 12 Anas crecca 0 0 0 0 0 0 0 0 1 0 1 12. 12 Anas querquedula 0 0 0 0 0 0 1 0 0 0 1 13. 27 Anthus pratensis 5 2 3 3 2 6 1 1 11 6 40 14. 28 Anthus spinoletta 0 0 0 0 0 1 0 0 0 0 1 15. 26 Anthus trivialis 55 39 17 31 251 144 51 33 24 11 656 16. 20 Apus apus 1 0 0 0 0 0 0 0 0 0 1 17. 19 Asio flammeus 0 0 0 0 0 0 1 0 0 0 1 18. 19 Asio otus 0 0 2 0 2 0 3 1 0 6 14 19. 17 Buteo buteo 0 0 1 4 4 1 0 0 1 0 11

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20. 21 Caprimulgus europaeus 0 0 0 1 0 0 0 0 0 0 1 21. 123 Carduelis cannabina 1 1 1 3 53 13 2 37 2 3 116 22. 122 Carduelis carduelis 3 0 3 15 3 3 4 8 7 6 52 23. 120 Carduelis chloris 19 47 60 67 88 43 113 834 109 87 1467 24. 123 Carduelis flammea 0 0 1 0 0 0 0 0 0 0 1 25. 122 Carduelis spinus 0 0 1 0 10 0 23 1 0 5 40 26. 123 Carpodacus erythrinus 0 0 0 0 0 1 0 0 0 0 1 27. 107 Certhia brachydactyla 4 4 8 17 17 9 15 13 10 20 117 28. 105 Certhia familiaris 3 8 31 11 8 6 5 8 14 16 110 29. 12 Circus aeruginosus 0 0 1 0 2 0 0 0 0 0 3 30. 125 Coccothraustes coccothraustes 0 10 49 11 29 4 12 232 12 44 403 31. 17 Coturnix coturnix 1 1 2 3 0 11 10 1 2 0 31 32. 25 Delichon urbica 132 1 5 7 160 161 36 44 41 11 598 33. 22 Dendrocopos medius 0 1 3 0 2 1 1 0 2 2 12 34. 23 Dendrocopos minor 2 4 5 10 6 8 12 4 0 3 54 35. 22 Dendrocopos syriacus 0 0 0 1 0 0 0 0 1 0 2 36. 22 Denrocopos major 2 5 7 6 16 7 10 12 11 17 93 37. 22 Dryocopus martius 0 0 0 1 0 0 2 0 0 0 3 38. 131 Emberiza calandra 29 9 1 3 1 51 6 0 2 0 102 39. 127 Emberiza citrinella 8 53 103 209 242 382 269 79 167 111 1623 40. 129 Emberiza hortulana 1 0 0 0 1 0 0 0 0 0 2 41. 129 Emberiza schoeniclus 46 94 79 49 26 44 159 84 151 94 826

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42. 34 Erithacus rubecula 229 530 656 753 649 566 667 673 737 915 6375 43. 17 Falco tinnunculus 0 0 1 0 0 0 0 0 0 0 1 44. 90 Ficedula albicollis 0 0 0 3 6 8 3 3 7 6 36 45. 90 Ficedula hypoleuca 74 52 66 137 99 118 65 22 95 115 843 46. 89 Ficedula parva 1 1 2 3 3 3 0 0 0 2 15 47. 116 Fringilla coelebs 12 26 40 25 67 32 29 767 54 56 1108 48. 118 Fringilla montifringilla 7 2 18 2 1 0 14 708 11 3 766 49. 18 Gallinago gallinago 0 0 0 0 0 1 2 1 2 0 6 50. 18 Gallinula chloropus 0 2 0 0 0 0 2 0 0 0 4 51. 113 Garrulus glandarius 0 18 11 5 32 2 28 4 7 17 124 52. 62 Hippolais icterina 18 6 4 7 24 25 18 27 14 8 151 53. 23 Hirundo rustica 235 5 19 69 271 197 131 36 92 48 1103 54. 20 Jynx torquilla 5 3 5 9 7 15 15 20 22 13 114 55. 110 Lanius collurio 41 25 37 77 72 92 110 63 109 67 693 56. 113 Lanius excubitor 0 2 2 1 2 1 4 2 3 1 18 57. 55 Locustella fluviatilis 4 2 0 2 0 3 9 4 2 2 28 58. 55 Locustella luscinioides 0 0 0 0 0 1 0 3 1 3 8 59. 53 Locustella naevia 7 9 5 22 4 29 25 12 8 8 129 60. 123 Loxia curvirostra 0 0 0 0 1 0 0 0 0 0 1 61. 37 Luscinia luscinia 3 3 2 0 3 4 4 3 4 2 28 62. 38 Luscinia megarhynchos 3 2 1 10 6 12 11 11 10 5 71 63. 38 Luscinia svecica 0 1 1 0 0 1 0 0 0 1 4

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64. 18 Lymnocryptes minimus 0 2 0 0 0 1 0 0 1 0 4 65. 20 Merops apiaster 1 0 0 0 0 0 0 0 0 0 1 66. 28 Motacilla alba 9 2 0 6 13 20 1 0 4 12 67 67. 28 Motacilla cinerea 14 1 0 0 0 3 0 0 0 0 18 68. 28 Motacilla flava 0 0 0 0 4 7 2 1 1 0 15 69. 87 Muscicapa striata 10 16 22 47 57 27 37 17 22 27 282 70. 44 Oenanthe oenanthe 0 0 0 1 4 3 0 0 2 0 10 71. 110 Oriolus oriolus 1 0 0 0 0 0 1 0 0 0 2 72. 96 Parus ater 0 12 41 2 9 0 27 86 1 7 185 73. 97 Parus caeruleus 69 142 484 188 364 132 1045 396 179 430 3429 74. 96 Parus cristatus 0 0 0 0 0 0 0 0 0 1 1 75. 100 Parus major 92 171 357 153 339 176 540 635 139 241 2843 76. 96 Parus montanus 0 0 1 0 0 0 2 0 0 2 5 77. 94 Parus paustris 9 8 50 16 38 16 14 22 12 32 217 78. 114 Passer domesticus 1 0 0 0 1 8 1 2 1 0 14 79. 114 Passer monatnus 237 364 391 182 349 784 253 350 228 163 3301 80. 17 Phasianus colchicus 0 0 2 2 0 0 0 1 0 1 6 81. 38 Phoenicurus ochruros 5 9 38 43 27 39 39 19 29 27 275 82. 39 Phoenicurus phoenicurus 4 16 20 23 15 15 20 18 13 17 161 83. 77 Phylloscopus collybita 100 221 309 332 239 360 556 415 347 480 3359 84. 75 Phylloscopus inornatus 0 0 0 0 1 0 0 0 0 1 2 85. 75 Phylloscopus proregulus 0 0 0 0 0 1 1 0 0 0 2

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86. 75 Phylloscopus sibilatrix 3 4 8 11 38 26 9 13 15 36 163 87. 80 Phylloscopus trochilus 50 33 86 123 39 136 57 233 39 62 858 88. 22 Picus canus 0 0 0 0 1 0 0 1 0 0 2 89. 22 Picus viridis 0 3 3 2 3 1 3 1 1 7 24 90. 18 Porzana porzana 1 1 0 0 0 1 10 0 0 0 13 91. 32 Prunella modularis 10 46 59 59 90 79 123 137 119 75 797 92. 123 Pyrrhula pyrrhula 0 15 27 27 43 1 71 28 3 10 225 93. 17 Rallus aquaticus 0 0 0 0 0 0 4 1 2 0 7 94. 85 Regulus ignicapillus 48 33 30 38 15 25 37 20 25 40 311 95. 82 Regulus regulus 128 112 521 451 42 85 270 108 146 201 2064 96. 109 Remiz pendulinus 0 3 11 1 0 0 0 0 0 7 22 97. 23 Riparia riparia 4 1 0 0 2 0 6 0 0 0 13 98. 41 Saxicola rubetra 5 0 6 16 74 27 45 3 5 2 183 99. 42 Saxicola torquata 0 19 10 46 61 35 37 23 12 31 274

100. 18 Scolopax rusticola 0 0 0 0 0 0 1 0 0 0 1 101. 120 Serinus serinus 1 2 0 4 5 6 4 13 6 9 50 102. 104 Sitta europaea 5 6 14 22 19 7 15 15 14 14 131 103. 19 Streptopelia turtur 4 0 0 0 1 0 0 3 0 1 9 104. 114 Sturnus vulgaris 0 2 0 1 2 1 21 12 7 44 90 105. 71 Sylvia atricapilla 248 364 236 319 394 489 535 630 627 709 4551 106. 69 Sylvia borin 27 24 16 20 36 46 60 70 81 45 425 107. 67 Sylvia communis 47 34 30 81 147 129 181 184 118 82 1033

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108. 64 Sylvia curruca 65 72 68 71 161 116 123 163 114 92 1045 109. 64 Sylvia nisoria 1 0 0 0 8 8 3 3 4 3 30 110. 17 Tringa glareola 1 0 0 0 0 1 0 0 1 1 4 111. 18 Tringa ochropus 1 1 0 0 0 0 1 5 1 1 10 112. 28 Troglodytes troglodytes 56 88 89 55 30 27 70 34 43 47 539 113. 51 Turdus iliacus 0 37 33 102 6 3 5 53 36 81 356 114. 44 Turdus merula 147 193 138 162 119 138 169 172 189 570 1997 115. 48 Turdus philomelos 21 50 89 64 51 62 105 146 95 150 833 116. 48 Turdus pilaris 0 2 2 0 1 0 3 1 0 5 14 117. 53 Turdus viscivorus 0 0 0 0 1 0 0 0 0 0 1 118. 20 Upopa epops 0 0 0 0 0 0 0 0 1 0 1 Total

2428 3185 4579 4367 5109 5151 6668 7923 4618 5591 49619