Materials and Methods to study relationships between woodpigeon (Columba palumbus ) autumn migrations’ flight’s heights and meteorological-orographical factors : preliminary report-experience 2019 on a single “crossing site-region”( Liguria) in Italy. Cavina Enrico(*),Bucchi Rinaldo,Bianchi Denis,Giovanetti Graziano,Feligetti Vasco Club Italiano del Colombaccio (*)–


The present paper is according to
and it is built to be read – preferably – on P.C.







The definition of “dynamic flight habitat”puts strongly the need to investigate the flight-height of birds during the migration and inside/over the “grounds’ habitat”.
We investigated these items in a special region –Liguria- in North Italy that is crossed in autumn by thousands of flocks of woodpigeons ( Columba palumbus) : they meet various meteo-orographic conditions and over them the birds realize flight’s strategies to overcome a complex orography and connected temporal atmosphere ( Air Pressure,isobaric corridors,winds,humidity,visibility,extreme events).
The present paper –online- offers a detailed material examined by original methods ,comparing flight’s height and some meteo-factors.

The overview is based on  this Map and depending on the recording-work of 9 hunters developed by special accuracy on three sectors ( special Project MSM) East,Central,West.

Our results show a clear relationship between high Air Pressure ,tail winds,and best temporal flight’s height conditions.variously recorded from East to West .

The Authors underline the need of better statistical insights and better research on the anatomy-phisiology of the Para-Tympanic Organo (PTO) of Vitali as supposed barometer-altimeter.

Key-words : flight’s height,Air Pressure,winds,orography,migration routes/paths,isobaric corridors

INTRODUCTION and preliminary discussion

The study of the birds’ flight-height’s dynamics is a special research-section involving the research on the migration phenology of many birds species also as woodpigeons ( Columba palumbus) (A) ,birds migrating on long-medium-short distance ,and usually by medium-large flocks.

Very recently (2019) it was written “ Interactions between landscape and atmoshere result in a dynamic flight habitat wich birds may use opportunistically to save energy during flight.However ,their ability to utilize these dynamic landscapes and its influence on shapping movement paths is not well understood    ( Sage E. & co-Authors :Orographic lift shapes flight routes of gulls in virtually flat landscapes” Scientific Reports (2019) 9:9659 on-line at

-This paper ( and connected references) must be considered basic for our report.

Recently our research efforts developed many items concerning autumn migration phenology in Italy , and some of them are focused on the relationships between Air Pressure and “Paratympanic Organ of Vitali” ‘s ( PTO)   supposed function as barometer/altimeter : references (2017-18-19-20) of our investigations are available on the Italian Journal Woodpigeon Research (IJWR)- journal on-line ( ( AC,AD,AE,AG)

During the last two seasons 2018-19 –inside the spirit of Citizen Science – some papers have been published on IJWR : Rinaldo Bucchi –, // Denis Bianchi also these papers are basic for the present report together with the fundamental paper :  Cavina E., Bucchi R.,Busse P. – 2018 – THE GENERAL PATTERN OF SEASONAL DYNAMICS OF THE AUTUMN MIGRATION OF THE WOOD PIGEON (COLUMBA PALUMBUS) IN ITALY-  THE RING 40 (2018) 10.1515/ring-2018-0001 (AA)

Other recent analysis is related to the autumn migration 2018 : “Woodpigeon’s (Columba palumbus) autumn 2018 migration : a new method to study dynamic patterns along a single crossing route in Central Italy .Focus on “flocking” , “hunting pressure” , “ woddpigeons’ ages “ in a preliminary report .” 

Italian Journal Woodpigeon Research – 19 August.2019

On these basic papers (main reference :MSM )(25) we have the opportunity to do a special focus on a special transit area ( tyrrenian coast and inlands of Liguria Region ) wich presents various and special orographic characters as following .

  • coastal  inlands with many valleys and hills until the mountain barrier of Marittime Alps
  • large sea-gulf that is first on the Mediterranean coast just before the second Lion Gulf (France) at west.










The geography of the Liguria area realizes various metereological conditions (10) ( sometimes suddenly changing) (    and and connected atmosheric movements ( interactions between landscape and atmoshere ) through those the birds must choice virtual air corridors possibly “opportunistically  useful to save energy during the flight “.

The winds (3,6,18,22bis,29,40,44,47,50,58,76,91,99,105,113,123,131,142,163,169,185,197,205,212,215,220,223,AH) play a fundamental role as by seasonal prevalence in various territories and locations

Sometimes –as in November-the Genoa gulf is offended by extreme meteorological events ,testimonials of the frequent variability of Air Pressure conditions   between inlands and the open sea.

On this geographic  area and its climatology and meteorology , we have migration routes followed by woodpigeons at the end of their journey just before the final jump and arrivals to the wintering territories in France,Iberian peninsula,Corse and Sardinia,North Africa .These routes are enclosed in the main Mediterranean Flyway on the way from North-East /East Europe passing through Adriatic coast ,East Italy,and mostly through  Padania valley beneath Alpes chain :  it happens by variable stop-over timing.








The last main route on North Italy is from the flat Padania valley ( rich of agriculture-feeding-fields ) ,crossing Appenine mountains between Emilia and Liguria regions,arriving  at west in front of the Marittime Alps ,then crossing them or avoiding at south on the sea , continuing along mediterranean French coast.

An important south-branch route is on the East “door” of the Liguria gulf where after crossed first mountains and passes of Liguria Appennine ,many flocks fly south along the Tyrrenian coast to SanRossore pine-forest(Pise) and other Tuscany stop-over areas or directly to Baratti’s bay and Elba island or Argentario-Punta Ala ,crossing then the sea to Corse (France) and Sardinia big islands.All the Tuscany Achipelago area is crucial to overcome open sea to west /south-west as by data in the paper of Denis Bianchi .   

All the area are subjected to sudden movements of air sacs in atmosphere as by complex meteorological maps of the Region(  ).                                                               

                                                                             ***      ***     ***

We’ll try  to  expose our “reading” of “Liguria” migration’s  air paths by a scientific (Citizen Science) lingo and method , but we must underline that the best knowledge of the various migration’s movements is prerogative of local hunters ,valley by valley,mountain by mountain,village by village.They say that the variables of the migration’s patterns are many,incostant,impredictable through soft or strong winds on the valleys and green canyons,where the flocks choise offhanded fly-routes and altimetric air-corridors  .

If the most important records are connected with flocks started from Padania valley and Adriatic coast flying at low or medium heights over the grounds , many times the hunters observe –only by binoculars – many big flocks at very high atmospheric corridors ( 800-1500 m ) : these flocks very probably- specially during “furor” waves- arrive directly –starting at night or very early hours of the day – from long distance East Europe mass take-offs’ sites .

On the base of so complex local (Liguria) migration’s patterns on the recording sites (nine), we can only present some –also unusual- models of investigation on the relationships between flight-dynamics and meteorological factors. 

We are well conscious that the present our preliminary report (Citizen Science) will need criticism,revisions and better statistical processing ,approches,algoritmic methods by official scientific Researchers and Institutions ,if available to analyze these problems. We ( Citizen science) need help of statistics’ ornithologists specialists :



Our materials (A) are extracted from the basic work MSM producted 2020  Author :Rinaldo Bucchi  where on 3 “combs” in Liguria as filters on nine migration’s routes crossing Italian peninsula, the total count of the monitored woodpigeons –and related flight’s heights – is 168.090 recorded birds / total Italy MSM 751.600 . ( A )

Other data and informations arrived from “Forum” ( ) and Regional contributes . 

Our focus was on three combs ( 1 EAST -1 CENTRAL- 1 WEST ) along 240 km of the Liguria-coast  and three stations for each “comb” have collected ( 1st October-15th November 2020) 168.090 woodpigeons wich represents 22,35 % of the total MSM ( differentiated peaks in MSM ) recorded in Italy as :

  • Liguria (East-Central-West ) 168.090
  • Emilia-Romagna  518.000
  • Marche 65.610





The counts of Flocks :






The total area Liguria was monitored by 9 stations as by represented area Latitudes-Longitudes between Long. 10,30 E – 7,30 E / Lat. 43,30 N – $5 N  , mostly focused between Long.10,30 E – 7,30 E / Lat.44 N – 44,30 N 


Stations located ( by Municipality-area ) as :

  • EAST  : Fosdinovo 44,07 North/10,01 East – Camogli 44,20 / 9.09 – Recco 44,21 / 9,08
  • CENTRAL : Genoa Coronata 44,43 / 8,88 – Pegli 44,42 / 8,80 – Albisola Marina 44,19 / 8,30
  • WEST: Noli 44,12 / 08,24 – Pietrabruna 43,88 / 7,90 – Garlenda 44,01 / 8,05

managed by Bruzzone Antonio,Bruzzone Sergio,Canepa Stefano,Capurro Mauro,Celsi Federico,Mamberto Nicolò,Meistro Marco,Pelazza Ivo,Quarantella Andrea ( alphabetical  list ). 

All the expert hunters have recorded and collected detailed data about the flight’s heights ( high,medium,low) of the observed flocks –day by day,hour by hour – on their note-books ,then revised in our analysis .

The hunters (nine) have recorded  ( 1st October-15th November ) 168.090 woodpigeons in 2057 flocks ( average 82 woodpigeons / 1 flock ) and the flight height was :

  • < 100 m.  : 744 flocks ( 36,17 %) for 69.097 birds
  • 100-300 m. : 733 flocks ( 35,63 %) for 50.446 birds 
  • > 300 m. :580 flocks ( 28,2 %) for 48.547 birds as by timetabled graphics

The flight-height data “Liguria” seem quite in contrast respect the data of transit in Central Italy as by   .

In this paper  ( monitored  607.749 woodpigeons in 5.805 flocks) the analysis of flight-altitude is related on a path from Adriatic coast to Tyrrenina coas along a single corridor of 300 Km (large 15-25 km) over hills and Appennine mountains  ( “continental” orography quite different respect the Liguria’s orography ) mostly dominated by stable Air Pressure isobares  : most important data extracted from this analysis ( Central Italy 2018) are as following 

  • flocking : the sizes of flocks changes during the fly crossing Central Italy in the corridor “Mesola forest – Elba island” as by a sequence fom average values of 306 birds (Mesola) to 46 ( Central Tuscany) and then 81 and finally 156 ( increasing 239%) in Elba island .
  • Height of flying: low 24,52% ,medium 26.66% ,high 51,87%
  • Meteo : all the peaks/waves are connected with jumps of Air Pressure more than 10 hPa.

It is evident the contrast high “ flight-height “average percentages between  Liguria  28,2 % –and Central Italy  51,87%- 

It confirms the complexity of the Liguria orography between inlands and sea where the migration developes itself mostly arriving from the flat Padania valley and then along the coastal borders . 

We must also consider that in 2018 the migratory season happened with a prevalent stable high A.P. all over Europe and Mediterranea area ,contrary of the season 2019 when the Air Pressure was unstable and weather’s conditions very variable ,also in Liguria during the season.

About the character of the size-flock during migration across Italy ( A ),specifically for 2019 a detailed analysis is reported in MSM 2020 (, ) but we must underline again that  a depth detailed analysis on a specific migration corridor crossing Central Italy (2018) has been reported as by : big flocks arrive after the crossing of Adriatic sea ,the size of the flocks reduces itself over the continental Italy , and is growing up again in front of the flight over Tyrrenian sea . The same happens along the secod route from Liguria “East door” until Elba island along Tuscany coasts , ready to continue flying over the sea to  Corse and Sardinia. Biggest flocks fly over the sea at higher flight’s height . 

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The related analysis(77) of meteorological factors and relationships with migration phenology concerns all the area crossed by two main migration paths ,as by preliminary discussion in present “Introduction” and related maps and complex orography  where         the winds can channel and force the flocks at various height .



















The factor “winds” ( as force and direction) is most import factor as well known by all hunters , important to determine the flight’s height .( 3,6,7)

A schematic syntesis of the winds evolution over the migrations’routes is very difficult specially if considering the complex orography of the Liguria region as well represented in 3D Googlearth maps.Inside the single valleys or canyons or crests and passes of the mountains the normal or falling winds can change suddenly in direction and/or force influencing the flight decision making preferences of the birds during their journey .

However – as overview of prevalent winds – we have extracted the directions of winds as aforementioned above in present “Introduction” . Concerning our study ,transit area and related dates-timing and peaks , we have:














All these winds’data are extracted by detailed reports in Archives of……….._

selecting the prevalent winds in the first hours of every day .

What emerges ?

EAST area (between Padania valley and sea-coast ) : N-NE-E for 21 days / NE for 10 days/ SE for 12 days/no winds for 2 days

CENTRAL area : N-NE for 33 days / NW for 3 days / SE for 9 days

WEST area : NW 42 days /NE for 1 day/ SE for 2 days

It is evident that different local orographies (85) are basic for the prevalence of winds during the migratory season. All the peaks-days have prevalence of tail-winds from North (N-NE-NW ). A special mention must be done for the Gulf of Genoa (Central area) where the prevalence of North winds (36/45) is very evident , and for the “west exit-door” the North prevalence is NW 42/45 days and NE 1/45 . 

A part the local orography we must consider the isobaric & Air Pressure maps on larger European areas or total  Europe .Indeed , if the local movements are influenced by local meteo-orographic conditions ,these conditions are inside larger seasonal meteo-climatological-status of the atmosphere in Western Paleartic continental area . Consulting many Meteo-websites (10)we can take detailed knowledge on the isobaric general Map of Europe day by day .







Extracted from these general conditions and local meteo-data we know that altimetric air corridors –as “interactions between landscape and atmoshere” ( “dynamic flight habitat”) –(72,91)  are realized and  offer themselves to the flight choices of the birds (83) who utilize all the coordinated forces of their admirable sensitive-eco-system (PTO , mechanoreceptors, central nervous system).

Regarding the relationship between migration and Air Pressure changes , we have collected all the A.P. changes ( day by day  – East-Central-West ) between 24h before and the recording day , and marked peaks’dates. 



On the same investigative target we have collected the “Air Pressure tendency Index”  (APTI) (,  as above along all the 45 days of the migration season and related peaks as by the graphics ( blue –East/ red-Central/green-West ——- black Average ).                                                                                                                         


Most important aim of the present study is to identify possible correlations between flight-height of woodbigeons and meteorological factors –as above “winds”- remembering cited “dynamic flight habitat”.

The hunters-observers have signed every day the height of flocks monitored as low(< 100m)-medium ( 100-300m.) – high ( > 300 m.)  on the total numbers of birds and flocks . Here basic raw numbers total Liguria : 

The total results –all over Liguria- of “ heights’ monitoring” are represented as above by the following graphics 

and AIR PRESSURE TENDENCY index (Appendix -1- )

Legenda : it appears quite evident that we have some differences related the “AP tendency indexwich is higher(red line) in Central area ( open sea –Gulf) respect the other two areas (East- West )  particularly during the Peaks  as confirmed by the “changing AP 24h before the migration day monitored “ higher .

                                  Peaks  >                         ^                       ^                            ^                     ^

To better investigate the daily preferential height of the birds we have calculated every day – disrecpectful of numbers-values’ quantity of woodpigeons – the percentage of birds at high,medium,low height in the same day  . The “East,Central,West” transits are represented in the following graphics ,also considering the peaks’days more representative of the migration’s phenology of the day and local meteo-conditions .





























































A syntesis of all these graphics – all these graphics are the results of long detailed collecting of data,numbers,percentages – a syntesis by uniform looks of confront appears really difficult in the field of Citizen Science where we have poor confidence with statistical methods .

We underline as above “We are well conscious that the present our preliminary report (Citizen Science) will need criticism,revisions and better statistical processing ,approches,algoritmic methods by official scientific Researchers and Institutions ,if available to analyze these problems. We ( Citizen science) need help of statistics’ ornithologists specialists”

However we see that higher flights’ heights are documented in some areas and waves/ peacks’timing when higher isobaric corridors run at more than 100 m. ,and when the Air Pressure is increasing and persists ,and when the “migration furor” is stronger well represented by big flocks in very high altitude ,and when Northern winds are prevalent .   


The above presentation of  “materials and methods” very probably could be read as a “dead letter” if projected to do answers as requested about “interactions between landscape and atmosphere “ at present time “ not well understood”. We tried to present “materials and methods” , as title of the present paper becouse we are not able to present much more that is in the traditional experience of hunters. 

Apart from the rigid reading of the recorded numbers ,it seems important considering the empirical portray of “Liguria transit” as by historical knowledge of some traditional hunters ( Walter Porcile in Forum,4223.0.html)

< The preferential routes are from East to West ( France coast ) ,but we must consider the populations entering on extreme East Riviera and preferring turning the direction to South  along Tyrrhenian coast until Elba area ,and from here crossing the sea  to Corse ( France) . It seems difficult to believe that also big flocks cross the open-sea directly to Corse ( Walter Porcile ) . On other hand many flocks arriving from Piacenza area ( Padania valley) ,depending on winds ,can cross the Gulf of Liguria directly to Caprazoppa Point – Alassio . Departing from Padania valley they have to choices ,influenced by winds/weather : crossing hills and  mountains of inlands through Val Trebbia or directly to Arenzano Varazze .Many flocks outflank the mountains of the Liguria inland until the barrier of Marittime Alps ,some flocks crossing them on the Cuneo area ,many flocks lowering close to the coast crossing the Melogno Pass or directly to Albenga and then following the coast until France coast .> 

The observations of many big flocks ( more than thousands of woodpigeons’ size) visible only by binoculars and the increasing populations crossing Liguria during the last 2-3 years , according with the observations on Tagliamento valley ( Friuli region at East ) ( personal  communication Piero Barbieri -2019 ) and according to the results of the investigation concerning the woodpigeons’ migration ( recent seasons) on the Polish Carpathians (  Rafael Bobrek, Tomasz Wilk, Aleksandra Pępkowska-Król  – Migration of the Woodpigeon Columba palumbus in the Polish Carpathians – migration parameters and birds’ selectivity for meteorological variables Ornis Polonica 2017, 58: 160–177 ) and according with the arrivals’census in Mesola forest area ( Denis Bianchi  2020 ) , all these factors leave to speculate that during last 3-4 years many northern populations ( North East Russia ,Belarus,Baltic Countties) have abandoned the Central Europe Flyway ( north of Alps Mountains ) preferring to alight at South on the Mediterranean Flyway through the Morava Port ( Sudeti mountains- Polish Carpathians )  . The observations  ,specially during peaks/waves , of many big flocks at very high altitudes ( High Pressure altimetric corridors) could be explained by very long distance take-offs of populations flying over Padania valley in a single step ,directly overcoming the Gulf of Liguria . This item is an acceptable  hypothesis to be confirmed , probably only by satellite monitoring . 

Concerning  the numbers of woodpigeons crossing Liguria – as for all Italian peninsula- are growing in order 200-300% during last 3 seasons : the local observations can differ very much between themselves depending on weather and winds’ condition.

As a first step of our investigation we have focused the study on a difficult geographic area characterized by a complex orography that influences drammatically the variables of air corridors useful to optimize the flight’s height to save energies .

Concluding  ,some elements can be underlined : 

  • All our data confirm that the main behaviour’s character of Columba palumbus is the flexibilty (224),wich is very probably the best weapon  to defend the optimal excellent status of the species in Western Paleartic –Europe

We have underlined the special characteristic of Orographic factors present on the Liguria Region , so much rich of variables able to influence strongly the local ,and also detailed local atmospheric conditions where the Woodpigeons’ flocks need to perform instant decision-making useful for the best altimetric corridors over the open sea or along the coasts or over the inlands hills and mountains or mostly Inside valleys and their changing of winds , all factors influencing their choices to continue to West along the French coasts and inlands or divert to South along the Tuscany coasts – until the Tuscany Archipelago ( isle of Elba) – before crossing the sea to West to Corse and Sardinia .

It seems to us that this crucial area ( Liguria Region) can be considered as a natural geographic transit “ laboratory” able to try the most important character of Columba palumbus Species : flexibility .

It seems to us that our results are according with the deep sense of analysis ,very recently ( Front. Ecol. Evol., 26 March 2020 | reported in a Review Artcle ( Endogenous Programs and Flexibility in Bird Migration by Susanne Åkesson and Barbara Helm) .Some important concepts can be extracted from this Review Article ,all of them pertinent with many items of our study .

“….. Central questions in this research field are concerned with inheritance of migration programs, their integration of environmental cues, and their physiological and genetic mechanisms (van Noordwijk et al., 2006; Åkesson et al., 2017; Merlin and Liedvogel, 2019). Here, we review key insights, and apply them to address sources of variation in spatiotemporal migratory traits within and among individuals, as well as among populations. ……”

“….. , we still need to understand exactly how the endogenous migration program interacts with external information, and how birds keep track of space during long migrations throughout the annual cycle…………..

….. Variation within the population can reveal interesting characteristics, where interactions of the inherited migration phenotype with different environmental factors can lead to the evolution of diverse migration patterns. Phenotypic plasticity may for example lead to advancement of migration timing in response to environmental conditions, in particular in flock-migrating birds…….”

“ ………..Programmed Flexibility in Response to Environmental

Within the time window set by the program for migration, decisions about its implementation are sensitive to a range of environmental factors that determine successful migration . Departure time may be adjusted in response to level of fuel reserves, and relative to the expected onward migration route, including distance of barrier crossings (Müller et al., 2018). Responses to these environmental factors are partly inherited, and therefore, we here expand on their effects. We consider some other aspects of flexibility, for example learning, social behavior and responses to weather, to represent residual flexibility……”

“……For some aspects of migration, we begin to understand the birds’ flexibility on the basis of inherited reaction norms that provide solutions which were effective over evolutionary time. “ ( 224- Susanne Åkesson , Barbara Helm -2020 –   )

  • Positive : the collecting ad registering methods to outline data, have been developed by particular accuracy.
  • Negative : our ( Citizen Science)  ability to develop right scientific statistical analysis is poor.
  • Evidence based analysis seemed almost representative about a right reading of many items,based also on  raw numbers.
  • It appears almost impressive the coincidence of the peaks’dates and the average line(black) of the Air Pressure jumps on the total area  as in

The growing up of the “A.P.tendency index” is significative of a trend of atmosphere’s conditions stabilized as high values and connected none or poor turbolences.

  • North and North East winds faciltate the arrivals and transits of flocks , and North West winds are prevalent at west area ( exit area ) close to the Marittime Alps at inland and to Gulf of the Lion (France) at the coasts .
  • The focused “area Liguria” is not an area where we can observe mass take-offs , but the not officially recorded evidence –in some peaks’ days – of big or very big flocks identified only by binoculars at very high altitude on the sea , is evidence of great populations flied from long-middle distances sites along the origins’ area ( K.Hobson -2009)










arriving and cotinuining  the migratory fly over the Gulf , specially during the days with stable high A.P. conditions.

  • In contrast of so many references concerning the relationships between migration behaviour and Air Pressure-meteo conditions , the item “ barometric/altimetric sense “ is  not or poorly developed and discussed.
  • Some Researchers have introducted the doubt that the study of flight’s height is not so important for the study of the migration phenology : it seems that so numerous specific  References collected in our Bibliography are a sufficient answer to this doubt.
  • The present research ( Liguria) can be considered preliminary, ready to be carried on the other MSM “ combs” in Central Italy .
  • Finally the key-consideration : if all these data are connected with a “barometric-altimetric” sense(AC,AE,AG) and related sensitive eco-system and supposed related anatomy ( Paratympanic Organ of Vitali ) , the challenge is in front of the Science to better investigate this sense. 


( marked References for specific interest

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2-Åkesson S, Weimerskirch H (2005) Albatross long distance navigation: comparing adults and juveniles. J.   Navigation 58:365–373. doi:10.1017/S0373463305003401

3-  Alerstam T (1979) Optimal use of wind by migrating birds: combined drift and  overcompensation. J Theor Biol 79:341–353. doi:10.1016/0022-5193(79)90351-5

4- Alerstam T (1990) Bird migration. Cambridge University Press, Cambridge

5- Alerstam T (2001) Detours in bird migration. J Theor Biol 209:319–331

6- Alerstam T (2011) Optimal bird migration revisited. J Ornithol 152:5–23. 7-

7- Alerstam T, Lindström A (1990) Optimal bird migration: the relative importance of time, energy, and safety. In:    Gwinner E (ed) Bird migration: physiology and ecophysiology. Springer, Berlin, pp 331–351

8- Allen PE, Goodrich LJ, Bildstein KL (1996) Within- and among-year effects of cold fronts on migrating raptors at Hawk Mountain, Pennsylvania, 1934-1991. Auk 113:329–338

9-Alves JA, Dias MP, Méndez V, Katrínardóttir B, Gunnarsson TG (2016) Very rapid long-distance sea crossing by  a migratory bird. Sci Rep. doi:10.1038/srep38154

10-American Meteorological Society (2016) Glossary of meteorology. Accessed 3 May 2017

11-Andueza M, Arizaga J, Belda EJ, Barba E (2013) The role of extrinsic and intrinsic factors on the departure decisions of a long-distance migratory passerine. Ardeola 60:59–72

12-Arizaga J, Belda EJ, Barba E (2011) Effect of fuel load, date, rain and wind on departure decisions of a migratory passerine. J Ornithol 152:991–999. doi:10.1007/s10336-011-0685-2

13-Bäckman J, Alerstam T (2003) Orientation scatter of free-flying nocturnal passerine migrants: components and causes. Anim Behav 65:987–996. doi:10.1006/anbe.2003.2119

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17-Berchtold A, Nightingale I, Vandermeer C, MacDougall-Shackleton Scott A (2017) Experimental temperature manipulations alter songbird autumnal nocturnal migratory restlessness. Anim Migr 4:1. doi:10.1515/ami-2017-0001

18-Beth P (1961) Weather and early spring migration in Iowa. Wilson Bull73:41–45

19-Bildstein KL, Zalles JI (2005) Old world versus new world long-distance migration in accipiters, buteos, and falcons. The interplay of migration ability and global biogeography. In: Greenberg R, Marra PP (eds) Birds of two worlds: the ecology and evolution of migration. Johns Hopkins University Press, Baltimore, pp 154–167

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21-Bohrer G, Brandes D, Mandel JT, Bildstein KL, Miller TA, Lanzone M, Katzner T, Maisonneuve C, Tremblay JA (2012) Estimating updraft velocity components over large spatial scales: contrasting migration strategies of golden eagles and turkey vultures. Ecol Lett 15:96–103. doi:10.1111/j.1461-0248.2011.01713.x

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Updating specific  REFERENCES

212-Meier C.M. & co-Auth. (2018) What makes Alpine swift ascend  at twilight ? Novel geolocators reveal yearround flight behaviour . Behav.Ecology Sociobiol.(2018) 72:45 – related bibliography

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214Alerstam T.,Ulfstrand S. (1974) A radar study of the autumn migration of woodpigeons Columba palumbus  in southern Scandinavia  – IBIS  116.522-542 

215- Altshuler D.L. & co-Auth- (2015) The biophysics of bird flight : functional relationships integrate aerodynamics,morphology,kinematics,muscles,and sensors- Can.J.Zool. (2015)vol.93:961-975

216- Bagg A.M & co-Auth. ( 1950) Barometric pressure-patterns and Spring migration . Wilson Bull.(1950)vol.62.1 : 5 19

217- Becciu P. & co-Auth. ( 2019) Enviromental effects on flying migrants revealed by radar . Ecography (2019)42:1-14

218- Blomkvist O. & co-Auth. ( 2012 ) Mathematical modeling of flocking behaviour  Royal Institute Technology – Bachelor Yhesis  SA104X –SA108X- May 25,2012 pgg 1-51

219- Breuner C.W. & co-Auth. (2013) Environment,Behavior anPhysiology : do birds use Barometric Pressure ti predict storms ? Biol.Science Faculty Pub. Univ. Of Montana 6-1-2013 /pgg. 1982-1990 J. Exp.Biol.

220– Brown R.E.,Fedde M.R. (1993) Airflow sensors in the avian wing. J.Ex.Biol. (1993)179,13-30

221- Dokter A.M. & co-Auth. (2010) Bird migration flight altitudes studied by a network of operational weather radars . Jour.Royal Soc.Interface (2010) DOI :10.1098

222-Johnson K.P. & co-Auth ( 2010) The flight the Passenger Pigeon : Phylogenetics and biogeographic history of an extinct species . Molec.Philog. and Evol. (2010)57:455-458

223- Sachs G. (2009). Speed stability in birds . Mathematical Bioscence (2009)19,1,1:6. 

224- Åkesson S and Helm B (2020) Endogenous Programs and Flexibility in Bird Migration.
Front. Ecol. Evol. 8:78. doi: 10.3389/fevo.2020.00078

Special personal bibliography  and update March 2020  at Publications –extra-special (A – AA>AH)

  1. All papers (including 2020 ) in   >>


AA)  Cavina E.,Bucchi R.,Busse P. – (2018) – The General Pattern of Seasonal Dynamics of the Autumn Migration of the Wood Pigeon (Columba Palumbus ) in Italy – THE RING 40 (2018) 10.1515/ring-2018-0001and *****


AB) Cavina E.,Bucchi R.,Bianchi D.,Giovanetti G.,Feligetti V.,Giannerini S.,Bececco L –La Migrazione autunnale del Colombaccio ( Columba palumbus) in Italia –Monography – Ed.Aracne Editrice-Roma – 2017


PTO –special  Bibliography 

AC) Cavina E. : THE PARA-TYMPANIC ORGAN (PTO) of VITALI : a documental *

continuing forgotten request to the SCIENCE  ****

and related bibliograpphy 


  • Vitali G. Di un interessante derivato della prima fessura branchiale nel passero. Anat Anz. 1911;39:219–224.
  • Vitali G. Di un interessante derivato dell’ectoderma della prima fessura branchiale nel passero. Un organo nervoso di senso nell’orecchio medio degli uccelli. Anat Anz. 1912;40:631–639
  • Vitali G. Il comportamento dell’organo della prima fessura branchiale (placode epibranchiale) nei Selaci. Monit Zool Ital. 1925;36:122–130.
  • Vitali G. Sui disturbi funzionali e sulle lesioni istologiche dipendenti della distruzione dell’organo nervoso di senso da me descritto nell’orecchio medio degli Uccelli. Atti dell’Acc Dei Fisiocritici di Siena. 1915a;5 (7):291–302
  • Vitali G. Il comportamento dell’organo della prima fessura branchiale negli anfibi, nei rettili e nei mammiferi. Ricerche Morfol. 1924;4:191–220.



*      Ruffini A. Sull’organo nervoso paratimpanico di G. Vitali od organo del volo degli uccelli. Boll Sci med Soc med-chir

       Bologna. 1920:141–154.


AD) Francesco Giannessi,a,*Riccardo Ruffoli,a and Christopher S. von Bartheld – (2013 ) –Giovanni Vitali: Discoverer of the Paratympanic Organ – Ann Anat. 2013   Jan;195(1): 10.


AE) von Bartheld C.S.,Giannessi F.- (2011)- The paratympanic organ: a barometer and altimeter in the middle ear of   birds?- Exp. Zool. (Mol. Dev. Evol.) 316:402–408, 2011

  • Giannessi F. On the presence of reciprocal synapses in the paratympanic organ of the chicken. Anat Embryol (Berl) 1989;180:175–178.
  • Giannessi F. Ultrastructure of the supporting cells of the paratympanic organ in the chicken: a preliminary study. Boll Soc Ital Biol Sper. 1990;66:99–104
  • Giannessi F, Ruffoli R. The ultrastructure of the sensory hair cells of the paratympanic organ receptor cells in chicken. Anat Embryol. 1996b;193:569–575
  • Federici F. Ueber die Innervation des von Vitali entdeckten Sinnesorgans im Mittelohr der Voegel (sogen. paratympanisches Organ) Anat Anz. 1927;62:241–254.
  • Benjamins CE. Y a-t-il une relation entre l’organe paratympanique de Vitali et le vol des oiseaux? Arch Neerl Physiol. 1926;11:215–222.
  • Benjamins CE. L’organe paratympanique de Vitali, soi-disant organe du vol des oiseaux, chez le pingouin. Acta otolar. 1939;27:266–270.

        Giannessi F, Pera L. On the presence of fibers of probable efferent function in the paratympanic organ in chickens. Boll 

       SocBioSper. 1987;63:337–340  

  • Giannessi F, Fattori B, Ruffoli R, Gagliardo A. Homing experiments on pigeons subjected to bilateral destruction of the paratympanic organ. J Exp Biol. 1996;199:2035–2039.
  • Barry MA, Boord RL. The spiracular organ of sharks and skates: anatomical evidence indicating a mechanoreceptive role. 1984;226:990–992.
  •   Baker CVH, O’Neill P, McCole RB. Lateral line, otic and epibranchial placodes:developmental and evolutionary links? ExpZoolB Mol Dev Evol. 2008;310:370–383.
  • Petrash EW, Andres KH, von Düring M, Delius JD. Morphologische Aspekte des Vitali-Organs bei der Taube. Anat Anz. 1983;153:288.
  • Ruffoli R, Giambelluca MA, Giannessi F. Ultrastructure of the supporting cells in the paratympanic organ of chicken, Gallus gallus domesticus. J Morphol. 1998;236:65–73.
  • Simonetta A. L’organo di senso dello spiracolo e l’organo paratimpanico nella sistematica dei vertebrati. Arch Ital Anat Embryol. 1953;58:266–294
  • von Bartheld CS. Development and innervation of the paratympanic organ (Vitali organ) in chick embryos. Brain Behav Evol. 1990;35:1–15
  • von Bartheld CS. Functional morphology of the paratympanic organ in the middle ear of birds. Brain Behav Evol. 1994;44:61–73.
  • von Bartheld CS, Giannessi F. The paratympanic organ: a barometer and altimeter in the middle ear of birds? J Exp Zool B Mol Dev Evol. 2011;316:402–408. [PMC free article]
  • von Bartheld CS, Rubel EW. Paratympanic and spiracular sense organs: phylogenetic distribution and theories of function, including hearing. In: Webster DB, Fay RR, Popper AN, editors. Evolutionary Biology of Hearing.Springer; New York: 1992. pp. 582–583.
  • O’Neill P. Molecular mechanisms underlying paratympanic organ development and evolution. RIKEN FY2009 Foreign Postdoctoral Researchers. 2010;22:193–194.

AG ) Martin G. – (2017)- The sensory ecology of birds – Oxford Univ.Press 2017

AF ) Alerstam T.-(1997)- Bird Migration – Cambridge Univ.Press 1997

AH ) Newton I.- (2007) – The migration ecology of birds – Elsevier Pub. 2007