Bird population dynamics in the wintering season : the case of the mediterranean Gull Larus melanodephalus

Dinàmiques poblacional s en ocells durant la hivernada : el cas de la Gavina Capnegra Larus melanocephalus

Carles Carboneras Malet

A questa tesi doctoral està subjecta a la llicència Reconeixement - NoComercial 3.0. Espanya de Creative Commons .

Esta tesis doctoral e stá sujeta a la licencia Reconocimiento - NoComercial 3.0. España de Creative Commons .

Th is doctoral thesis is licensed under the Creative Commons Attribution - NonCommercia l 3.0. Spain License .

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   ACKNOWLEDGEMENTS

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INTRODUCTION ...... 9 Part I – Bird population dynamics in the wintering season ...... 9 Migration as a strategy to face environmental changes ...... 9 General rules, and exceptions, of birds’ migratory strategies ...... 11 Variations on a migratory theme: alloheimy and synheimy ...... 13 Winter site-fidelity and winter population structure ...... 15 Small-scale movements within the wintering season ...... 19 Part II – Conservation: a wintering area perspective ...... 19 Part III – Study species and study system ...... 21 OBJECTIVES ...... 32 SUPERVISORS’ REPORT ...... 37 Summary of results ...... ¡Error! Marcador no definido. Chapter I ...... 43 Inferring geographic origin, population structure and migration routes of a wintering population of Mediterranean gulls from resightings data...... 43 Chapter II ...... 45 Mediterranean Gulls Larus melanocephalus wintering in Spain and Portugal: one population or several? ...... 45 Chapter III ...... 47 Contrasting patterns of site-fidelity across spatial scales in wintering gulls...... 47 Chapter IV ...... 49 A new breeding population of Mediterranean Gulls Larus melanocephalus in the species’ main wintering area maintains independent spatial dynamics ...... 49 DISCUSSION ...... 51 CONCLUSIONS ...... 69 RESUM DE LA TESI DOCTORAL EN CATALÀ ...... 73 D. REFERENCES ...... 89 Appendix – Tables ...... 102

   INTRODUCTION

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Part I – Bird population dynamics in the wintering season

Migration as a strategy to face environmental changes

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  Fig. 1. Differential migration in several species of gulls as shown by the median latitude of the ringing recoveries of adult (crosses; n = 1160) and immature (black dots; n = 2125) birds from the British Isles. Notice that Larus argentatus and Larus marinus are non-migratory, and that immature birds are found further north in one species, Larus canus. Data from Siriwardena & Wernham (2002).

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 Variations on a migratory theme: alloheimy and synheimy

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Fig. 2. Distribution of the breeding and wintering areas of the Lesser White-fronted Goose (Anser erythropus) in Eurasia, with the migratory routes connecting them. This is a good example of alloheimy, or geographical segregation of populations of the same species. Map layout: Grid- Arendahl, Original data: Norwegian Ornithological Society, WWF Finland, Swedish Association of Hunting and Wildlife Management.

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 Fig. 3. Breeding areas and migratory routes of Mediterranean Gulls (Larus melanocephalus) wintering in the study system. The general wintering area (black box) is used by birds from different breeding populations, spanning over 30° of longitude and represented here by the centroid of their nesting colonies. The lack of geographical segregation may be indicative of synheimy. Map source: Carboneras et al. (2013).

Winter site-fidelity and winter population structure

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  Box 1. Demographic independence and metapopulation structure

Esler (2000) analysed the degree of demographic independence among subpopulations—and thus of the applicability of metapopulation theory—throughout the entire annual cycle of migratory birds. For metapopulation theory to apply, in its classical definition, (1) subpopulations must be sufficiently independent that extinction of a subpopulation can occur irrespective of the demographics of other subpopulations and (2) dispersal among subpopulations must be frequent enough that recolonisation of extinct subpopulations can occur. Subpopulations can be considered to be demographically independent when the above criteria apply; panmixia occurs when subpopulations, in the metapopulation context, do not exist because of a lack of independence of extinction probabilities. The term subpopulation applies to either breeding or wintering areas and may be defined as groupings of birds that are demographically independent. Instances of demographically distinct groups with no probability of exchange of individuals would be appropriately described as isolated and cannot be considered under the metapopulation construct.

Fig. 4. The four scenarios of spatial and temporal dynamics of migratory birds between breeding and wintering areas described by Esler (2000). Ovals represent geographically distinct groups, solid arrows represent migration, and dashed arrows represent dispersal.



 Small-scale movements within the wintering season

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Part II – Conservation: a wintering area perspective

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Fig. 5 Comparative size (using body mass as proxy) of the Mediterranean Gull in relation to the other five species in the “black-headed” or Ichthyaetus clade (above), and to other seven gull species with which it occurs sympatrically in the western Mediterranean region (below). Data from Handbook of the Birds of the World Alive (retrieved from http://www.hbw.com).

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Fig. 6. Location of the study system area showing, right, a schematic representation of the spatial distribution of Mediterranean Gull concentrations associated to the principal fishing harbours. Circle size is proportional to the average Mediterranean Gull numbers on a logarithm scale; there is one order of magnitude difference in the population size between Cambrils (black circle) and Vilanova i la Geltrú (grey circle), the two main localities sampled for marked birds; white circles indicate other sites. Site location is arranged according to the distance (in km) to Blanes (41º41’N, 2º48’E), at the N edge of the wintering area.





   Fig. 7. Current (2015) Fig. 7. Current of the distribution Gull in Mediterranean (black the breeding wintering dots) and (grey squares) in 50 plotted seasons, squares. x 50 km Updated from (1997). et al. Bekhuis is The species cells in 127 present bird as a breeding in the cells and in 128 wintering season.

   Fig. 8. Pre-1960s Fig. 8. Pre-1960s of the distribution Gull in the Mediterranean and dots) (black breeding squares) wintering (grey x 50 in 50 plotted seasons, At the time, km squares. in was present the species bird a breeding as 15 cells in the cells and in 102 wintering season.   7KH0HGLWHUUDQHDQ*XOOKDGEHHQXQNQRZQWRVFLHQFHXQWLODERXW\HDUVDJR

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 Fig. 9. Green 0E9, born on the French Mediterranean coast near Camargue, shown here in the study system in its first-winter plumage. Photo: Carles Carboneras.



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Chapter I

Inferring geographic origin, population structure and migration routes of a wintering population of Mediterranean gulls from resightings data

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   Popul Ecol DOI 10.1007/s10144-013-0362-9

ORIGINAL ARTICLE

Inferring geographic origin, population structure and migration routes of a wintering population of Mediterranean gulls from resightings data

Carles Carboneras • Giacomo Tavecchia • Meritxell Genovart • Susana Requena • Marc Olive´ • Daniel Oro

Received: 11 May 2012 / Accepted: 14 January 2013 The Society of Population Ecology and Springer Japan 2013

Abstract Winter congregations of migratory birds are were consistent with a migratory strategy that combines made by individuals of different origins and generally fluvial and coastal routes in an optimal way, seeking assumed to be variable across space and time, but the minimal distance along favourable terrain. We found that, demographic characteristics of these temporal populations after the first year, annual local survival at the wintering are poorly known. We used 2,216 observations of 472 site (0.81 on average) was comparable with the one esti- colour-ringed individuals to estimate the annual local sur- mated at the breeding colonies, indicating a high individual vival of Mediterranean gulls Larus melanocephalus win- fidelity to the areas used in winter. Our work shows that tering in NE Spain. In addition, by gathering the ringing winter groupings may behave as real populations, shaped information on the 19,856 individuals marked as fledglings by breeding output and survival, and that the geographic in 18 countries between 1990 and 2009, we were able to origin of wintering birds can be explained by a simple infer the composition of population in relation to the model. The study of winter congregations can help country of origin. We coupled these estimates with geo- understand a species’ population structure and movement graphic information to contrast hypotheses on the migra- strategies. tory pattern most likely used by the gulls in their first migration from their natal colonies to the wintering area. Keywords Capture–recapture Larus melanocephalus The probability of reaching the study area was negatively Movement strategy Site-fidelity Spatial variation associated with the distance from the natal colony. Data Survival

Electronic supplementary material The online version of this Introduction article (doi:10.1007/s10144-013-0362-9) contains supplementary material, which is available to authorized users. A large body of evidence indicates that the winter season is C. Carboneras (&) a critical period in the annual life cycle of most birds Departament de Biologia Animal, Universitat de Barcelona, wintering at high or temperate latitudes (Newton 1998; Avda. Diagonal 645, 08028 Barcelona, Spain Fort et al. 2009) and that overwinter mortality is the major e-mail: [email protected] determinant of population dynamics, even among those G. Tavecchia M. Genovart D. Oro species that migrate to warmer areas in autumn (Barbraud Institut Mediterrani d’Estudis Avanc¸ats IMEDEA (CSIC-UIB), et al. 1999; Tavecchia et al. 2001; Grande et al. 2009; Miquel Marque`s 21, 07190 Esporles (Mallorca), Spain Sergio et al. 2011). The winter period, however, is often S. Requena regarded as a ‘black box’ in the life cycle of many birds Institut de Cie`ncies del Mar (CSIC), Passeig Marı´tim de la because demographic processes during this season are Barceloneta 37-49, 08003 Barcelona, Spain largely unknown, particularly for migratory species that are assumed to be distributed over large wintering areas M. Olive´ Rambla de Sant Jordi 26, 4rt 1a, 08800 Vilanova i la Geltru´, (Frederiksen et al. 2012). This lack of knowledge is Spain reflected into conservation practices, which concentrate 123 Popul Ecol most of their efforts to protect populations during breeding. framework for studies on survival and population composition Recently, developments of tracking techniques have pro- in relation to origin and age. Here we tackle the problem of vided new insights into the migratory and wintering how wintering populations of gulls are formed and main- behaviour of medium- and even small-sized birds tained, and how they relate spatially to the breeding pop- (Baechler et al. 2010; Egevang et al. 2010; Catry et al. ulations. The mixing up of birds from different origins is 2011; Stenhouse et al. 2012). Results confirmed that spe- common among wintering seabirds (Gonza´lez-Solı´s et al. cific areas are often predictable as wintering and stop-over 2007; Reynolds et al. 2011; Frederiksen et al. 2012), but no sites (but see Gschweng et al. 2008) although considerable studies have so far analysed the relative composition of variation exists in individual site tenacity across years wintering populations linked to their distance from the (Marques et al. 2010; Dias et al. 2011; Guilford et al. birds’ natal areas. As colonies are expected to contribute 2011). Age- or sex-related differential migration is the differently in relation to migration costs, and these will norm among migratory birds (Cristol et al. 1999), and depend on the routes followed, we expect to be able to infer latitudinal segregation by age classes has been found in the most likely migratory strategy used by this long-lived most species of charadriiform seabirds (Siriwardena and gull. We use the information on 472 individually-marked Wernham 2002), including many gull species (Oro and Mediterranean gulls of known age and origin, recorded Martinez 1994; Marques et al. 2010). However, some repeatedly at a wintering area: (1) to estimate interannual seabirds distribute according to alternative migration pat- survival and winter site fidelity; (2) to estimate population terns, e.g., leap-frog system (Hallgrimsson et al. 2012) and composition on the wintering grounds in relation to the oriented chain migration (Fort et al. 2012). spatial distribution of natal colonies. The heritability of migratory routes (Pulido 2007; Mueller et al. 2011) and the repeatability of large-scale movement patterns (Gonza´lez-Solı´s et al. 2007; Dias et al. Materials and methods 2011) suggest that winter congregations may be structured according to bird origins. Hence, wintering populations of Study area and season birds can offer the opportunity to assess survival, popula- tion composition and origin of individuals. Population The 350 km of Mediterranean coast of NE Spain, between composition and winter site fidelity can be used to estimate the towns of Blanes (N41410,E2480) and Burriana annual survival probability and to infer migration routes, an (N39530,W0050), constitute one of the main wintering important information to understand population function- areas for the Mediterranean gull (Cama et al. 2011). The ing and guide management action (see for example area is delimited by two stretches of unoccupied, yet Hestbeck et al. 1991; Gauthier et al. 2001; Tavecchia et al. potentially adequate, habitat. In this defined area we con- 2005). ducted observations of individually marked birds for 5 The Mediterranean gull (Larus melanocephalus)isa consecutive winters, from 2005 to 2010. middle-sized, migratory gull whose current breeding pop- Records of ringed gulls were obtained by watching pre- ulation is confined entirely to Europe with the centre of roost flocks standing on beaches or in fishing harbour gravity in the Black sea, and significant populations in the premises, from a distance, using a telescope (Table 1). Low Countries, the Danube valley and the Mediterranean (Bekhuis et al. 1997). The recent expansion of the species Table 1 Summary of ringing and reading effort from Eastern Europe towards the Atlantic coast and the Country Ring Total chicks Recorded in Ring NW Mediterranean has been linked to the fluctuations of colour marked study area readings the Black sea stronghold (e.g., Sadoul 1997). The majority ATL White/ 6,420 15 81 of the population spends the winter around the Mediterra- green nean, mostly in Spain, France, Italy and Tunisia (Cama MED Green 708 132 809 et al. 2011). Its discontinuous winter distribution has ITA Blue 3,435 159 715 remained fairly constant since at least the 1950s (cf. HUN Red 2,729 80 316 Mayaud 1954; Shevareva 1955), despite the recent changes POL Red 246 2 4 in the coastal landscape and marine ecosystem (Coll et al. GRE Black 2,002 45 116 2010). Favoured by its aggregative nature, the species UKR Black 4,316 39 175 has received the attention of multiple colour-ringing Total 19,856 472 2,216 programmes along its expanding range, with [25,000 individuals (including 19,856 as chicks in 18 countries) The table shows, for each country ringing programme, the total number of chicks marked in the natal colonies from 1990 to 2009, and colour-ringed in 20 years, 1990–2009. This extensive the number of individual gulls observed and the total number of rings database, both spatially and temporally, provides an ideal read in the study area over 5 winter seasons, 2005/06–2009/10 123 Popul Ecol

We obtained resightings over nine sites within the study area. We began the analysis from a general model that Most records (96 %) came from three sites only: Vilanova i assumed all parameters to vary over the years, noted ‘t’in u0 u la Geltru´, Cambrils and Tarragona. Gulls were present at the model notation (model 11 in Table 2). This model, t t pt, study area between late July and late March each year; is an extension of the Cormack–Jolly–Seber model with two however, we restricted the analysis to the observations age-classes for survival. The fit of this model was assessed made from 15 September to 15 February. This was done to using the software U_CARE 2.2 (Choquet et al. 2009; see exclude transient birds that are mainly present from June to Appendix in Tavecchia et al. 2008 for more details on the mid-September and after mid-February. The goodness of fit goodness of fit test). In addition to a year-effect in detection test confirmed the absence of transient animals in our probability we considered the effect of the ring colour dataset (see below). We thus retained 2,216 observations of (noted ‘g’ in model notation; Table 1) because plastic rings 472 birds of known age and origin (13 countries). To esti- might show differences in their visibility and/or rate of mate the proportion of birds from each breeding region that deterioration. We also used a measure of the reading effort spent the winter in our study area, we obtained information (the number of marked birds recorded each winter, noted on the annual ringing effort of the 19,856 individuals ‘no. readings’) as a covariate for the detection probability. colour-ringed at fledgling in eighteen different countries Juvenile survival, u0, and adult survival, u, were set to vary from 1990 to 2009. To define the composition of the win- depending on the year (noted ‘t’) and on the area of origin tering population, we sorted the data according to 7 main (noted ‘c’). To account for the progressive deterioration of areas of origin by pooling together colonies lying in nearby plastic rings over time [M. Olive´, unpublished data, Table countries or regions. Birds ringed along the Atlantic coast of S1 in Electronic Supplementary Material (ESM)] we added France, Belgium, The Netherlands and Germany were a linear trend in survival, noted ‘T’ (see ‘‘Results’’). Finally, considered in a single group (noted ‘‘ATL’’, hereafter). A we used colony distance to explain the variability in sur- second group was formed by birds born in Mediterranean vival parameters (see below). France (noted ‘‘MED’’). The other groups were made by The resulting 20 models were fitted using program birds born in Italy (noted ‘‘ITA’’), Hungary, Serbia and MARK (White and Burnham 1999). We selected the more Slovakia (noted ‘‘HUN’’), Poland (noted ‘‘POL’’), Greece and parsimonious models using Akaike’s Information Criterion Turkey (noted ‘‘GRE’’) and Ukraine (noted ‘‘UKR’’), respec- corrected for sample size (AICc; Burnham and Anderson tively. Finally, we gathered, as supplementary information, the 1998). Models within 6 points of AICc were considered as occasional resightings along the migratory route and in alter- equivalent (Burnham and Anderson 1998). native wintering areas of individuals in our data set to help discussions and the interpretation of results. Population composition and migratory routes

Modelling survival/site fidelity and recapture In our analysis, survival parameters represent a combina- probability tion of the true survival, i.e., the probability to survive from one winter to the next, and the probability of reaching the Observations of marked birds recorded at the wintering wintering grounds. In this respect, permanent emigration, area during the five-year period were coded into individual i.e., the probability of never visiting the area again, would encounter histories. An encounter history is a series of be confounded with mortality. Assuming that juvenile 0 and 1 s indicating the absence and presence of a given survival is the same regardless of the colony of origin, the animal, respectively. The frequency of capture histo- variation in the parameter of juvenile survival, u0, should ries follows a multinomial distribution whose elementary reflect the different proportion of birds reaching the study parameters are the survival and recapture probabilities. area. This can be used to test hypotheses on movement These probabilities are estimates from the data using patterns (Tavecchia et al. 2005). Indeed, colonies are maximum likelihood procedures (see Lebreton et al. 1992 expected to contribute to the wintering population differ- for details of maximum likelihood estimation of parameters ently according to their distance; however, the distance and model selection methods). We modelled three sets of between each breeding region and the wintering area varies parameters: the first-year survival probability, noted u0, the depending on the migration route. Hence, if migration survival probability during subsequent years, noted u, and followed an exclusively coastal route, colonies from the the recapture—or detection—probability, noted p. The ‘GRE’ and ‘ITA’ areas would be expected to contribute local survival probability during the first year u0 referred to similarly because the distances to the wintering area would the joint probability that an individual survived until be similar (2,500 vs. 2,336 km). In contrast, if birds midwinter and reached the wintering site in NE Spain favoured an inland route along the main rivers, ITA-colo- during its first year of life, an interval shorter (ca. half) than nies would be expected to contribute many more birds than any subsequent ones. GRE-colonies, because the distance in the first case is 123 Popul Ecol

Table 2 Modelling apparent annual survival and resighting probability of Mediterranean gulls recorded in the wintering area in NE Spain Model Juvenile survival (u0) Adult survival (u) Detection probability (p) AICc DAICc AICc weight np Deviance

1 cTt 5,069.332 0.000 0.471 14 866.828 2 cct 5,071.562 2.230 0.155 19 859.040 3 ctt 5,071.819 2.487 0.136 16 865.309 4 ctg? t 5,072.419 3.086 0.101 22 853.883 5 c • t 5,074.919 5.587 0.023 13 874.418 6 ctg? no. readings 5,083.852 14.520 0.000 19 871.331 7 c • g ? t ? g 9 t 5,143.650 74.318 0.000 41 886.987 8 ••g ? t 5,237.592 168.260 0.000 13 1,037.091 9 ••g ? no. readings 5,244.820 175.488 0.000 10 1,050.327 10 ••g ? t ? g 9 t 5,251.083 181.751 0.000 34 1,008.476 11 ttt 5,377.167 307.835 0.000 12 1,178.669 12 t • t 5,388.623 319.291 0.000 10 1,194.130 13 • tt 5,418.612 349.280 0.000 9 1,226.121

14 u1;…; u7 t 5,424.672 355.340 0.000 12 1,226.174

15 u1;…; u5 t 5,426.440 357.107 0.000 10 1,231.946 16 ••readings 9 occasion-1 5,440.039 370.707 0.000 4 1,257.555 17 ••t 5,666.397 597.065 0.000 6 1,479.911 18 ••g ? t ? g 9 t 6,103.199 1,033.867 0.000 33 1,862.598 19 ••g 6,212.783 1,143.451 0.000 9 2,020.292 20 ••• 6,417.110 1,347.778 0.000 2 2,238.628 0 Parameters: juvenile survival (u ) = survival of gulls in their first year of life; adult survival (u) = survival of gulls after hatch-year; u1;…; un = survival of gulls variable in n age-classes; p = detection probability. Effects: ‘c’ = ‘‘country’’ (origin-dependence); ‘t’ = time-depen- dence; ‘T’ = linear trend; ‘g’ = group-dependence (ring colour combination); ‘no. readings’ = total readings of marked gulls; ‘read- ings 9 occasion-1’ = ratio of total ring readings to the number of occasions; ‘9’ = interaction between effects; ‘?’ = additive relation between effects; ‘•’ = constant. AICc = Akaike Information Criterion corrected for sample size; DAICc = difference with the lowest AICc value; AICc weight = the relative importance of each model; np = number of identifiable parameters in the model; Deviance = model deviance. Models in bold were used to obtain averaged estimates

much shorter (1,071 vs. 2,070 km). To contrast hypothesis GRE and UKR would reach the wintering area after fol- on migratory routes, we calculated the weighted centroid, lowing the Danube and Rhone; and hypothesis D (com- i.e., the centre of the area rather than the geometric cen- bined routes) predicted that gulls would combine rivers and troid, for each of the 7 groups of ringing colonies, by coastal routes in an optimal way minimising the distance applying the weighted value of the number of individuals while not crossing inhospitable terrain, thus making UKR, in our data set that had been ringed at each site. The dis- GRE and ITA birds fly chiefly W over land and sea, and tance, di, between the weighted centroid of the ringing area ATL birds cross over France, following the main rivers i to the study area was measured on the ETRS89 geodetic towards the Mediterranean (Fig. 1). The four hypotheses reference frame (Annoni et al. 2000) using the software generated a different ranking of the colonies based on their ARCGIS 10.0. We considered four possible layouts: under probability to contribute to the population wintering in the hypothesis A (Euclidean distance) gulls would reach the study area according to the expected distance calculated wintering grounds by flying in a straight line over land and under hypotheses, A, B, C and D, i.e., four different vectors u0 water; under the hypothesis B (coastal routes) gulls would of di values. We modelled the parameter i, where i is the fly to the coast on the shortest possible route and then ringing area, by the equation of the form: ÀÁ ÀÁ follow the coastline travelling exclusively over marine 0 logit u ¼ a þ b ln dH waters, so that UKR birds would cross the Bosporus and i i H the Aegean sea, ATL would migrate through the Straits where di is the vector of distances under hypothesis of Gibraltar and ITA birds would fly round the Italian H (with H = A, B, C or D; see above). The significance of peninsula; inversely, hypothesis C (river routes) favoured b and the total variance explained by the distance under inland routes along the main rivers: ATL would follow the each hypothesis were assessed using the ANODEV pro- Rhine and Rhone to the Mediterranean while HUN, POL, cedure in MARK.

123 Popul Ecol

Fig. 1 Hypothetical sketch of the migratory routes of first- winter Mediterranean gulls Larus melanocephalus from their colonies of birth (the weighted centroids of the 7 groups of ringing sites: ‘‘ATL’’, Atlantic France, Belgium, The Netherlands and Germany; ‘‘MED’’, Mediterranean France; ‘‘ITA’’, Italy; ‘‘HUN’’, Hungary, Serbia and Slovakia; ‘‘POL’’, Poland; ‘‘GRE’’, Greece and Turkey; ‘‘UKR’’, Ukraine) to the wintering area in NE Spain (inside box). Based on demographic model 1 (Table 2), which assumes the spatial variation of juvenile survival u0, and on migratory hypothesis D (Table 3) of a combination of coastal and inland (rivers, lakes) routes in an optimal way, i.e., minimal distance along favourable topography and positive taxis towards water

Results

Modelling annual survival/site fidelity and recapture probability

The goodness of fit test indicated that the general model fitted the data adequately and that the extra-binomial variance was not large (v2 = 45.13, df = 38, P = 0.20). Models with origin-dependent juvenile survival u0 (models 1–7; Table 2) had the lowest AICc values, suggesting that the survival/movement parameter during the first half-year of life varied strongly according to the area of origin. This effect was not present in later years, as the probability of local survival after the first winter was independent of the colony of birth (model 2). Note that the origin of birds is not known after the first year as gulls might have attended a colony other than that of birth. We tried further to refine the Fig. 2 Inter-annual apparent local survival of Mediterranean gulls probability of detection by incorporating different combi- after hatch-year as obtained from the time-dependent model (model 3 nations of reading effort (number of readings, number of in Table 2; square symbols with 95 % confidence interval). The line occasions) or of ring colour (group) as explanatory vari- shows the linear trend in survival equivalent to 4 % annual tag loss, as estimated by the retained model (model 1) ables, but such models failed to improve the accuracy of a time-dependent p, and were not retained (Table 2). Models with an age-dependent survival probability (model 14 for 7 negative trend in survival was due to the age of the ring and age classes and model 15 for 5 age classes) had little not to the age of the bird, the annual local survival would support. In contrast, a model assuming a linear trend—on a be 0.90 (95 % CI 0.82–0.95, estimates from model 1), logistic scale—in survival to describe the progressive corresponding to the one of a bird carrying a 2-year old deterioration of the ring (model 1) was preferred (Table 2; ring. In contrast, if the decrease on survival is a real effect Fig. 2). According to this model, apparent survival chan- of age or a mixture of the two, the average annual survival ged from 0.90 (95 % CI 0.82–0.95) to 0.78 (95 % CI would be 0.81 (95 % CI 0.79–0.84; estimates from model 0.74–0.81) 4 years later (Fig. 2). Assuming that this 5). Although there is evidence of ring loss (supplementary

123 Popul Ecol material, Table 1), we cannot fully differentiate between different from 0 (Fig. 3; Table 3). Despite this general these two hypotheses. Detection probability fluctuated trend, the more distant colonies would be expected to between years, varying from 0.36 to 0.75, with an average contribute differently in relation to the migratory strat- value of 0.49 (95 % CI 0.42–0.55) in the retained model. egy considered. The Euclidean distance (hypothesis A) explained about a third (35.5 %) of the total variance of u0 across colonies but we retained a model assuming a com- Population composition and migration routes bination of routes (hypothesis D), which had the lowest AICc (Table 3). According to this model, distance The probability of reaching the study area during the first explained 48.8 % of the spatial component of u0. Note that winter varied largely across the different ringing areas, none of these models was preferred when compared to the ranging from 0.13 (area MED) to 0.01 (area ATL) (Fig. 3). one assuming a different parameter for each colony of As expected, the two nearest regions (MED and ITA), had origin (model 1; Table 2). So, despite the general trend, the highest probability to reach the area and contributed to there was an important variation that remained unex- the population wintering in the study area with nearly two- plained. This might be due to the presence of multiple thirds of the first-winter birds. In the probabilistic frame- 0 strategies occurring simultaneously in a single population work, the accuracy of the estimates of juvenile survival u (see ‘‘Discussion’’). (length of the 95 % CI bars in Fig. 3) was related to the ringing effort—the total number of juvenile birds ringed in each region (Table 1). The lowest probabilities of reaching Discussion the area corresponded to birds from the ATL and the UKR areas. Birds from HUN, GRE and POL showed interme- Consecutive resightings of the same gulls at the wintering diate probabilities; the wide confidence interval of the POL area in northern Spain gave us the opportunity to estimate group was a result of the small number of birds ringed (246 local survival, the product between true survival and site in total) and the small number of individuals present in the fidelity, of wintering birds. Also, by contrasting these study area (2) (Table 1). parameters against the total number of animals ringed at The main predictor of population composition was different breeding colonies we were able to assess the colony distance; indeed, in all hypotheses the slope of the 0 composition of the population according to bird origin. We relationship between distance and u was negative and additionally inferred the most likely migratory pattern by investigating if the colony-specific probability of reaching the wintering site was consistent with a direct migration route or whether birds deviated to follow rivers or coasts, or optimised their route in a combination of both. te Marvelde et al. (2009) found that local survival of Mediterranean gulls estimated at medium-sized (approxi- mately 600 pairs; L. te Marvelde, personal communication) breeding colonies was 0.86. We found a similar average estimate (0.81), indicating that fidelity to the wintering area is only ca. 5 % smaller than the propensity to return to the breeding colonies. Our analysis does not take into account a temporal emigration of individuals, which is normally reflected into the probability of recapture. Birds that were not seen in one season might well have visited an alter- native wintering area, but the high survival value indicates that they came back to the study site at least once more. The study of individual life histories (obtained from the Fig. 3 Plot of juvenile immigration probability against distance. ringing programme coordinators, listed in the Acknowl- Models with spatial variation in the survival of juvenile birds (first- time immigrants) obtained the lowest AICc values (Table 2). The edgements) provided supplementary information on tem- results for the retained model 1 are plotted here against the distance poral and permanent emigration that was in agreement with from the ringing areas, calculated according to the ‘optimal’ a high site fidelity: only 11 birds of 472 in the data set were combination of inland and coastal routes of hypothesis D (Table 3). recorded alive in other wintering areas (permanent emi- Bars represent 95 % confidence intervals. Notice the low probability of ATL for the estimated distance, which suggests that birds from this gration) and only 2 gulls visited alternative areas in sub- group may use alternative routes to reach the wintering area in the sequent winters and later returned to the study area Mediterranean, including the 4,000-km route round Gibraltar (temporal emigration). This strong fidelity to the wintering 123 Popul Ecol

Table 3 Proportion of the total variance explained by the distance between the colony of birth and the study area under the different migratory strategies (see also text) Hypothesis Expected colony ranking AICc Deviance Percentage Intercept Slope deviance (a; 95 % CI) (b; 95 % CI) explained

A MED \ ITA \ ATL \ HUN \ POL 5,295.40 226.62 35.5 4.1; 2.9/5.3 -1.03; -1.20/ \ \ - Direct flight GRE UKR 0.86 B MED \ ITA \ GRE \ HUN \ UKR 5,261.32 192.54 44.8 3.3; 2.4/2.2 -0.84; -0.96/ \ \ - Along coastlines ATL POL 0.72 C MED \ ITA \ ATL \ HUN \ POL 5,263.00 194.22 44.3 4.2; 3.2/5.3 -1.01; -1.2/ \ \ - Along rivers GRE UKR 0.87 D MED < ITA < ATL < HUN < GRE 5,248.17 179.39 48.4 4.8; 3.7/5.9 21.1; 21.3/ 2 Along rivers and < POL < UKR 0.95 coastlines Each hypothesis on movement patterns generated a different ranking of the colonies (2nd column). Differences in the ranking applied particularly to the ATL, GRE, UKR and POL groups. The model for hypothesis D, of a combination of river and coastal routes, had the lowest AICc and was selected (in bold) areas suggests that wintering individuals form a population, our model was able to generate predictions on population ‘‘a set of organisms belonging to the same species and composition in other wintering areas. For example, obser- occupying a clearly delimited space at the same time’’ vations of ringed gulls in Portugal (R. Flamant and (Wilson 2000). Also, the strong fidelity to the wintering C. Duponcheel, personal communication), confirmed the grounds despite major alterations in landscape in recent model’s prediction of a higher probability of ATL birds decades indicates a complicated spatial structure of win- and a lower probability of MED and ITA birds making up tering populations, where other factors besides the avail- that wintering population, relative to our study area. ability of suitable habitat could determine the occurrence of However, all models suggested a higher than expected a species. contribution of ATL colonies. It is possible that part of the We assumed that the true (as opposed to local) survival population from some colonies migrated along a ‘different- probability during the first 6 months was independent of than-optimal’ route. Cramp and Simmons (1983) suggested the colony of origin. This assumption permits to consider that certain passage might occur through the Straits of the variation in local survival as a consequence of the Gibraltar, on the basis of some adult birds moving along probability of reaching the wintering area. Thus, the spatial the western seaboard that returned to Black Sea colonies variation of juvenile survival u0 can be used to contrast through the Mediterranean. We cannot exclude this hypotheses on the selection of migratory routes (Tavecchia behaviour, but comparative studies in other wintering areas et al. 2005). As expected, u0 covaried with colony distance, should be done to answer this question. with the closest colonies contributing the most birds, but The spatial variation of juvenile survival u0 seems to the contribution of colonies further away would depend on reflect the historical route of expansion of the species as the migration route followed by the gulls. For example, if reported by Cramp and Simmons (1983). Gulls appear to birds followed coastlines only (hypothesis B), gulls from migrate along a combination of coastal and overland routes north-eastern European colonies (POL group) would be the on a broad front following a W–SW direction, not unlike least represented in the population (Table 3). None of these the pathway that led to the colonisation of wetlands in ultrastructural models was selected indicating that there central and Western Europe starting in the 1950s, rather was not a predominant strategy explaining the movement than travel directly to the wintering areas. Mayaud (1954) patterns of all groups. Instead, data suggested that birds and Shevareva (1955) analysed recoveries of Mediterra- adopted a mixed strategy migrating via rivers and coastal nean gulls ringed as chicks in Ukraine, at a time when routes in an optimal combination of minimal distance along breeding was restricted to the Black and Aegean seas. They favourable topography with positive taxis towards water. reported two findings that are relevant to our study: (a) that With this simplification we assumed that all birds from a our study area was already an important wintering ground given area migrated in the same way. This assumption is for Ukrainian birds in the 1940s and 1950s, and (b) that at likely to be wrong and is probably the reason why none of the time there was already a small flux of birds that reached the models including the distance as a covariate was pre- the Baltic and North Sea probably by following the main ferred. Nevertheless, simple and imperfect as it might be, European rivers (Dnieper, Danube, Rhine, Seine, Loire) in

123  Popul Ecol a westward direction. Thus, an incipient migratory pathway breeding birds: their distribution and abundance. T. & A.D. developed over inland Europe independently from the large Poyser, London Burnham KP, Anderson DR (1998) Model selection and inference. A expansion of the breeding range, which took place 3 dec- practical information-theoretic approach. Springer, New York ades later. Roughly the same migration routes are still Cama A, Josa P, Ferrer-Obiol J, Arcos JM (2011) Mediterranean maintained today, showing that the species probably has Gulls Larus melanocephalus wintering along the Mediterranean positive taxis towards flying over significant water bodies Iberian coast: numbers and activity rhythms in the species’ main winter quarters. J Ornithol 152:897–907 (rivers, lakes, coast) when on migration. However, the Catry I, Dias MP, Catry T, Afanasyev V, Fox J, Franco AMA, population composition on the wintering grounds today Sutherland WJ (2011) Individual variation in migratory move- must be different from the 1940s–1950s because the rela- ments and winter behaviour of Iberian Lesser Kestrels Falco tive distances from and to the breeding areas have changed naumanni revealed by geolocators. Ibis 153:154–164 Choquet R, Lebreton J-D, Gimenez O, Reboulet A-M, Pradel R (2009) greatly in the last 70 years. This ‘historical’ hypothesis U-CARE: utilities for performing goodness of fit tests and cannot be tested because it fails to generate objective manipulating CApture–REcapture data. Ecography 32:1071–1074 predictions on colony ranking. Coll M, Piroddi C, Steenbeek J, Kaschner K, Lasram FBR, Aguzzi J, Our results indicate that wintering aggregations form Ballesteros E, Bianchi CN, Corbera J, Dailianis T, Danovaro R, Estrada M, Froglia C, Galil BS, Gasol JM, Gertwagen R, Gil J, ‘real’ populations, and that these are not determined by the Guilhaumon F, Kesner-Reyes K, Kitsos M-S, Koukouras A, turnover of individuals as previously thought, but rather the Lampadariou N, Laxamana E, Lopez-Fe de la Cuadra CM, Lotze demographic processes of breeding output and survival. HK, Martin D, Mouillot D, Oro D, Raicevich S, Rius-Barile J, Indeed, winter groups of Mediterranean gulls are formed Ignacio Saiz-Salinas J, San Vicente C, Somot S, Templado J, Turon X, Vafidis D, Villanueva R, Voultsiadou E (2010) The by the same individuals that visit the same areas across biodiversity of the Mediterranean Sea: estimates, patterns, and time. Moreover, our results point out a model that may be threats. PLoS ONE 5:e11842 applied to predict the winter population origin and relative Cramp S, Simmons KE (1983) The birds of Western Paleartic, vol III. composition along its wintering distribution, and may help Oxford University Press, Oxford Cristol DA, Baker MB, Carbone C (1999) Differential migration to guarantee the long-term preservation of populations revisited: latitudinal segregation by age and sex class. In: Nolan throughout the species’ range and in all seasons. V Jr, Ketterson ED, Thompson CF (eds) Current ornithology. Kluwer Academic/Plenum Publishers, New York, pp 33–88 Acknowledgments We wish to thank the large number of people Dias MP, Granadeiro JP, Phillips RA, Alonso H, Catry P (2011) whose work made the present study possible, in particular the ringers Breaking the routine: individual Cory’s shearwaters shift winter and coordinators of the national ringing programmes: the late Tatyana destinations between hemispheres and across ocean basins. Proc Ardamatskaya, Nicola Baccetti, Martin Boschert, Albert Cama, Josef R Soc B Biol Sci 278:1786–1793 Chytil, Monika Czyzak, Camille Duponcheel, Joan Ferrer, Pete Egevang C, Stenhouse IJ, Phillips RA, Petersen A, Fox JW, Silk JRD Findley, Renaud Flamant, Vassilis Goutner, Lars Hansen, Su¨hendan (2010) Tracking of Arctic terns Sterna paradisaea reveals Karauz, Zsolt Karcza, Peter Meininger, Antonina Rudenko, Vladimir longest animal migration. Proc Natl Acad Sci USA 107:2078– Slobodnik, Jan Svetlik, Adriano Talamelli, Monika Zielinska, 2081 Andreas Zours and Antun Zuljevic. We are also indebted to the other Fort J, Porter WP, Gremillet D (2009) Thermodynamic modelling collaborators who kindly provided additional readings of Mediterra- predicts energetic bottleneck for seabirds wintering in the nean gulls. The following observers contributed [10 readings in the northwest Atlantic. J Exp Biol 212:2483–2490 study area during the winter season: Rau¨l Aymı´, Albert Cama, Fort J, Pettex E, Tremblay Y, Lorentsen S-H, Garthe S, Votier S, Pons Manuel Enrique Carballal, Joan Ferrer, Jose´ Luis Gren˜o, Ferran JB, Siorat F, Furness RW, Grecian WJ, Bearhop S, Montevecchi Lo´pez and Miguel Tirado. The study has received funds from the WA, Gremillet D (2012) Meta-population evidence of oriented Spanish Ministry of Science (Grant Ref. CGL2009–08298), the chain migration in northern gannets (Morus bassanus). Front Regional Government of the Balearic Islands and the European Ecol Environ 10:237–242 Regional Development Fund (ERDF). Frederiksen M, Moe B, Daunt F, Phillips RA, Barrett RT, Bogdanova MI, Boulinier T, Chardine JW, Chastel O, Chivers LS, Chris- tensen-Dalsgaard S, Clement-Chastel C, Colhoun K, Freeman R, Gaston AJ, Gonzalez-Solis J, Goutte A, Gremillet D, Guilford T, Jensen GH, Krasnov Y, Lorentsen S-H, Mallory ML, Newell M, References Olsen B, Shaw D, Steen H, Strom H, Systad GH, Thorarinsson TL, Anker-Nilssen T (2012) Multicolony tracking reveals the Annoni A, Luzet C, Megrin, Institute SA (2000) Spatial reference winter distribution of a pelagic seabird on an ocean basin scale. systems for Europe. European Commission, Joint Research Divers Distrib 18:530–542 Centre, Brussels Gauthier G, Pradel R, Menu S, Lebreton J-D (2001) Seasonal survival Baechler E, Hahn S, Schaub M, Arlettaz R, Jenni L, Fox JW, of Greater snow geese and effect of hunting under dependence in Afanasyev V, Liechti F (2010) Year-round tracking of small sighting probability. Ecology 82:3105–3119 trans-Saharan migrants using light-level geolocators. PLoS ONE Gonza´lez-Solı´s J, Croxall JP, Oro D, Ruiz X (2007) Trans-equatorial 5:e9566 migration and mixing in the wintering areas of a pelagic seabird. Barbraud C, Barbraud J-C, Barbraud M (1999) Population dynamics Front Ecol Environ 5:297–301 of the White Stork Ciconia ciconia in western France. Ibis Grande JM, Serrano D, Tavecchia G, Carrete M, Ceballos O, Dı´az- 141:469–479 Delgado R, Tella JL, Dona´zar JA (2009) Survival in a long-lived Bekhuis J, Meininger PL, Rudenko AG (1997) Mediterranean gull territorial migrant: effects of life-history traits and ecological Larus melanocephalus. In: The EBCC atlas of European conditions in wintering and breeding areas. Oikos 118:580–590

123 Popul Ecol

Gschweng M, Kalko EKV, Querner U, Fiedler W, Berthold P (2008) Sergio F, Tavecchia G, Blas J, Lo´pez L, Tanferna A, Hiraldo F (2011) All across Africa: highly individual migration routes of Eleono- Variation in age-structured vital rates of a long-lived raptor: ra’s falcon. Proc R Soc B Biol Sci 275:2887–2896 implications for population growth. Basic Appl Ecol 12:107–115 Guilford T, Freeman R, Boyle D, Dean B, Kirk H, Phillips R, Perrins Shevareva TP (1955) On the study of several biological features C (2011) A dispersive migration in the Atlantic Puffin and its of the Mediterranean Gull (Larus melanocephalus) by the implications for migratory navigation. PLoS ONE 6:e21336 ringing method. Trudy Byuro Kol’cevanija Moskva 8:46–90 Hallgrimsson G, Gunnarsson H, Torfason O, Buijs R-J, Camphuysen (in Russian) K (2012) Migration pattern of Icelandic Lesser Black-backed Siriwardena GM, Wernham CV (2002) Synthesis of the migration Gulls Larus fuscus graellsii: indications of a leap-frog system. patterns of British and Irish birds. In: Wernham CV, Toms MP, J Ornithol 153:603–609 Marchant JH, Clark JA, Siriwardena GM, Baillie SR (eds) The Hestbeck JB, Nichols JD, Malecki RA (1991) Estimates of movement migration atlas: movements of the birds of Britain and Ireland. and site fidelity using mark-resight data of wintering Canada T. & A.D. Poyser, London, pp 70–102 geese. Ecology 72:523–533 Stenhouse IJ, Egevang C, Phillips RA (2012) Trans-equatorial Lebreton JD, Burnham KP, Clobert J, Anderson DR (1992) Modeling migration, staging sites and wintering area of Sabine’s Gulls survival and testing biological hypotheses using marked animals: Larus sabini in the Atlantic Ocean. Ibis 154:42–51 a unified approach with case studies. Ecol Monogr 62:67–118 Tavecchia G, Pradel R, Boy V, Johnson A, Ce´zilly F (2001) Sex- and Marques PAM, Sowter D, Jorge PE (2010) Gulls can change their age-related variation in survival probability and the cost of the migratory behavior during lifetime. Oikos 119:946–951 first reproduction in breeding Greater Flamingos. Ecology Mayaud N (1954) Sur les migrations et l’hivernage de Larus 82:165–174 melanocephalus Temminck. Alauda 22:225–245 (in French) Tavecchia G, Serra L, Baccetti N (2005) Colony specific variation in Mueller JC, Pulido F, Kempenaers B (2011) Identification of a gene the use of a moulting site in the migratory little tern Sterna associated with avian migratory behaviour. Proc R Soc B Biol albifrons. J Avian Biol 36:501–509 Sci 278:2848–2856 Tavecchia G, Minguez E, De Leon A, Louzao M, Oro D (2008) Newton I (1998) Population limitation in birds. Academic Press, Living close, doing differently: small-scale asynchrony in London demographic parameters in two species of seabirds. Ecology Oro D, Martinez A (1994) Migration and dispersal of Audouin’s gull 89:77–85 Larus audouinii from the Ebro delta colony. Ostrich 65:225–230 te Marvelde L, Meininger PL, Flamant R, Dingemanse NJ (2009) Pulido F (2007) The genetics and evolution of avian migration. Age-specific density-dependent survival in Mediterranean Gulls Bioscience 57:165 Larus melanocephalus. Ardea 97:305–312 Reynolds TJ, Harris MP, King R, Swann RL, Jardine DC, Frederiksen White GC, Burnham KP (1999) Program MARK: survival estimation M, Wanless S (2011) Among-colony synchrony in the survival from populations of marked animals. Bird Study 46(suppl): of common guillemots Uria aalge reflects shared wintering 120–129 areas. Ibis 153:818–831 Wilson EO (2000) Sociobiology: the new synthesis, twenty-fifth Sadoul N (1997) The importance of spatial scales in long-term anniversary edn. Belknap Press of Harvard University Press, monitoring of colonial Charadriiformes in southern France. Cambridge Colon Waterbirds 20:330–338

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Chapter II

Mediterranean Gulls Larus melanocephalus wintering in Spain and Portugal: one population or several?

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Mediterranean Gulls Larus melanocephalus wintering in Spain and Portugal: one population or several?

Gaivotas-de-cabeça-preta Larus melanocephalus invernantes em Espanha e Portugal: uma ou várias populações?

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Chapter III

Contrasting patterns of site-fidelity across spatial scales in wintering gulls.

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   Contrasting patterns of site-fidelity across spatial scales in wintering gulls

Carles Carboneras, Giacomo Tavecchia, Meritxell Genovart and Albert Cama

C. Carboneras ([email protected]) and A. Cama, University of Barcelona, Department of Animal Biology, Diagonal 643, 08028 Barcelona, Spain. – C. Carboneras, G. Tavecchia and M. Genovart, Institut Mediterrani d’Estudis Avançats IMEDEA (CSIC-UIB), Miquel Marquès 21, 07190 Esporles, Mallorca, Spain. – C. Carboneras, Royal Society for the Protection of Birds, The Lodge, Sandy SG19 2DL, United Kingdom.

Site-fidelity is dependent on the size of the area to which the individual returns, so it is expected to vary according to scale. We examined spatial dynamics in a wintering population of Mediterranean Gulls known to show strong site-fidelity on a regional scale, but resightings of individually-coded colour-ringed birds indicated that they probably engaged in inter-site movements. To study interchange characteristics between localities, we pulled together resightings during 5 seasons from two near localities and used capture- recapture multi-site models to estimate the degree and directionality of movements at a small spatial scale. Results indicated that birds had a low monthly probability of remaining in their current locality (0.56) and that those changing location were equally likely to move to the other observed site (0.22) or to other areas. To estimate the total volume of birds at a single locality, we quantified local population turnover of individuals of all ages for each season independently. On average, 15800 birds visited the locality every season, a large proportion of the regional (43 %) and global (18 %) populations and 16-18 times above the standard midwinter counts. This is the first study of local-scale site-fidelity and within- season movement dynamics in any gull species; it reveals that although populations may be spatially stable and the seasonal presence of individuals may be predictable, they probably visited multiple localities in the course of a single season, including outside the designated protected areas. The high mobility of individuals found here and the high turnover of gulls in a given site has important consequences for the design of protected area networks.

The individual tendency to return to a given area habits, which may increase individual fitness and or site-fidelity is a biological trait with important overwinter survival (Robertson and Cooke 1999, consequences for survival and conservation Péron et al. 2010). Wintering has often been (Martínez-Abraín et al. 2001). Even if broadly considered a ‘black box’ in the annual cycle of studied during the breeding season, few studies migratory birds, while movements to and within have analysed the individual attachment to an the wintering area can reveal important area during the wintering season (but see (Harris individual strategies (Carboneras et al. 2013, et al. 2009, Grist et al. 2014). In the non-breeding Sergio et al. 2014). Migratory species are generally season, fidelity to the wintering area is also assumed to remain static during the summer and beneficial as a potential source of valuable local winter stationary periods, yet some bird species knowledge about food availability and predators’ show evidence of movements within the non- breeding season (Newton 2008). The study of probability of local survival and site-fidelity to the population turnover has revealed that some general wintering area (Carboneras et al. 2013) but wintering birds, particularly in the family observations of individually-coded colour-ringed Anatidae (ducks, geese and swans), regularly birds indicated that they probably engaged in change site within the same season, normally after inter-site movements. We analysed the volume, the depletion of food sources, extreme weather or direction and likelihood of movements between disturbance (Newton 2008); this results in the total two protected sites designated for the species. number of birds using a site being higher than the This setting was expected to provide information average number of birds present at any given time on the spatial scale of individual movements in (Giroux 1991, Pradel et al. 1997a, Frederiksen et al. relation to the network of protected areas. 2001, Rees 2006, Gourlay-Larour et al. 2013). From a conservation point of view, this means that Material and methods more individuals are afforded protection at a Study system given site than initially estimated, but also they are protected only for a short amount of time. We focused our investigation on two localities, Since the assessment of the importance of bird hereafter sites, about 50 km apart, Vilanova i la areas is generally based on instantaneous counts Geltrú (‘VNG’ hereafter; 41º14’N, 1º42’E) and and not on the total volume estimated (Heath et Cambrils (‘CAM’; 41º4’N, 1º3’E; Fig 1), on the al. 2000, Gourlay-Larour et al. 2013), there is a north eastern Mediterranean coast of Spain (Fig. potential mismatch between between the 1). Both sites host large numbers of wintering collective home range of a population and the size Mediterranean gull Larus melanocephalus with and location of the protected areas designated for average population in VNG of about 800 it. individuals, (EEA 2014) and in CAM of about A lot of effort has been put in recent years into 20,000 individuals, (Cama et al. 2011). Both sites defining the features, biotic and abiotic, that lead are listed in the Natura 2000 network under the to the identification of key sites in the terrestrial EU Birds Directive 2009/147/EC due to their and marine environments and into assessing their important populations of Mediterranean Gull. At adequate size and limits in order to protect them the centre of both sites are large fishing harbours (Boyd et al. 2008, Wilson et al. 2009, Ronconi et al. (Fig. 1, VNG: fleet = 83 vessels, average fish daily 2012). Most systems ultimately aim at identifying landings = 3·9t; CAM: fleet = 46 vessels, average the location of discrete areas of activity that are fish daily landing = 2·2t), which gulls attend to relevant for a substantial fraction of the feed on fish offal and discards. Previous work has population of a single species or a community of shown that the main wintering season of species, with a preference for multi-species Mediterranean Gulls in this area extends from 15 aggregation hotspots (Lascelles et al. 2012). That September to 15 February, and that stopover birds approach has the potential to leave part of the occur outside this period (Carboneras et al. 2013). area occupied by only one species unprotected. It Sites were thus visited regularly during this may also be unsuitable for highly mobile species period, from 2005 to 2010, in search of in which individuals shift location frequently and individually-marked birds as they concentrated move among neighbouring sites as part of their on beaches and around harbour premises before wintering strategy. they moved to roost at sea. Sampling effort was We looked at local-scale site-fidelity and within- more intense in VNG than in CAM, with 116 and season movement dynamics in a wintering 221 resighting days respectively. Observations population of Mediterranean Gulls Larus were made from a distance with the aid of a 20- melanocephalus. The species is known to show high 60x telescope, and there was no handling of any

bird. All gulls had been ringed during the site r, ϕr, and transition probability, ψrs, from site r breeding period in other countries over a large to site s. To maximize the number of individuals, area extending from the Black Sea to Belgium and we considered resightings made from the 8th to the Netherlands prior to their arrival in the study the 15th of each month (October to February) from system. 2005 to 2010. Our observations were only made on weekdays, when there was fishing activity. Before conducting the analysis, software U-CARE (Choquet et al. 2009) was used to assess the fit of the general model in which all parameters were site and time dependent. We assumed the presence of a non-observable site, ‘NOS’ hereafter, with the probability of resighting fixed to 0, to accommodate those individuals that could be alive but not seen neither in VNG nor in CAM (Hénaux et al. 2007). Hence, our computation allowed individuals to move freely among three sites, but only those in VNG and CAM would be visible. In this analysis, we considered four different models (ranked 1-4 in Table A1). Model 1 assumed equal movement probability among

Fig. 1. Location of the wintering area showing, right, a the three states, models 2 and 3 assumed a schematic representation of the spatial distribution of different movement probability associated to each Mediterranean Gull concentrations associated to the state and model 4 assumed equal probability of principal fishing harbours. Circle size is proportional to reciprocal movement between pairs of states, that the average Mediterranean Gull numbers on a is ψVNGCAM=ψCAMVNG and ψCAMNOS=ψNOSCAM but logarithm scale; there is one order of magnitude ψVNGCAM≠ψCAMNOS. Model selection followed an difference in the population size between Cambrils information-theoretic approach based on the (black circle) and Vilanova i la Geltrú (grey circle), the Akaike Information Criterion adjusted for the two localities sampled for marked birds; white circles small sample size AICc (Burnham and Anderson indicate other sites. Site location is arranged according to the distance (in km) to Blanes (41º41’N, 2º48’E), at the 2002), in which the model with the lowest AICc N edge of the wintering area. The shaded grey area values should be considered as the best indicates the location of the Marine Protection Areas compromise between model fit and the number of designated as Natura 2000 sites in the region. estimable parameters. Models with AICc values below 4 points were arbitrarily considered as

equivalent (Burnham and Anderson 2002). Between-site dispersal Winter population volume and super- To investigate the direction and frequency of population size interchange movements within the study area, we We investigated the population volume and the built a dataset with the weekly observations of turnover in the smaller of the two localities colour-ringed birds made over 5 seasons (30 considered, VNG, for which we had more data, occasions) at both sites, VNG and CAM. We using the encounter histories of birds seen in this fitted multi-site capture-recapture models in site for five consecutive winters (2005/06 to program MARK (White and Burnham 1999) to 2009/10). The total number of animals in VNG at estimate site-dependent survival probability at any given time was estimated using the POPAN

model (Arnason and Schwarz 2002) available in deviances (Cooch and White 2012). Following the software MARK 7.0 (White and Burnham Silva et al. (2010) we corrected the estimates of 1999). The POPAN formulation is a particular population size for the proportion of colour- parameterization of the time-dependent Cormack- marked individuals relative to the total number Jolly-Seber model (CJS; (Schwarz and Arnason available for observation, m. This value was used 1996). By assuming a super-population, N, made to estimate the number of birds present on the site of the total number of animals ever present in the (Atkinson et al. 2007) as: population, it is possible to estimate at each N Ngull occasion, i, a recruitment parameter, bi, which m represents the probability of entry in the observed where Ngull is the estimated number of population, i.e., the probability that an animal Mediterranean Gulls (marked and unmarked) from this hypothetical super-population would using a given site and N is either Nˆ or Nˆ *, enter the observed population between occasion i where Nˆ is the monthly and Nˆ * is the annual and i+1 (Schwarz and Arnason 1996). Within each gross super-population size, that includes all year we identified intervals of 4 weeks (~ one individuals that enter and leave the population month) and pooled resightings obtained over the between consecutive surveys and thus are never first week only, to meet the assumption of no available for sampling (Williams et al. 2011). mortality during the resighting period (Lebreton Monthly and seasonal estimates of the number of et al. 1992). We considered the 5 wintering Mediterranean Gulls visiting VNG were seasons independently and constructed one calculated based on the values of Nˆ in the dataset per season with the encounter histories of retained model and the proportion of colour- marked animals. We assumed that ring loss during a single winter was negligible. Not all marked birds calculated on the site ( x = 1·397 · 102, parameters of the general time-dependent SE = 0·055 · 102). POPAN model are estimable (Cooch and White We collected data on the apparent total number of 2012). We applied the constraint proposed by birds present at the site by means a single-count Williams et al. (Williams et al. 2011) to set p1 = p2 during each individual sampling occasion (n = 92). and pk = pk1, where k is the number of occasions, in This value has been used as a comparison with order to make all survival and entry parameters the estimated N. We also used for comparison the estimable in the model. The probability of entry, b, data of the local mid-winter International was kept time-dependent because models with Waterbird Census (IWC) counts carried out in constant b have no sensible biological January of each year by an independent observer interpretation (Cooch and White 2012). For each (n = 5) using the same methodology. winter, we tested a combination of four candidate models: a fully time-dependent model, ϕ(t) p(t), a Results model with constant survival and encounter probability, ϕ(·) p(·), and two models in which Between-site dispersal either encounter probability or survival The GOF test detected a highly significant probability was time-dependent while the other presence of transients or permanent emigrants in was held constant, ϕ(t) p(·) and ϕ(·) p(t). The 2 the general multi-site model (test 3G.SR = goodness-of-fit, GOF, of the full time dependent F 36 model was assessed with program RELEASE. The 68·00, P = 0·001). To reduce the effect of an total chi-square value of the GOF tests, divided by excessive presence of transients in the general the degrees of freedom, can be taken as a variance model, we suppressed the first capture occasion inflation factor, cˆ , and used to correct model

leaving that locality, 22% of moving to the other site and 22% of going somewhere else in the general area. Equally, an individual in state NOS had a 44% probability of moving to VNG or CAM. The monthly probability of remaining on the site, calculated by subtraction, was 0·56 (95% CI, 0·42– 0·66) (fig. 4).

Winter population volume and super- population size \ Fig. 2. Transition probabilities (95% CI) among In total, we recorded 667 resightings (97-217 per VNG (Vilanova i la Geltrú), CAM (Cambrils) and season; average 100) of 501 individuals (76-146 NOS (non-observable site) estimated by multi-site per season; average 133). The goodness-of-fit tests model 1, based on equal transition probabilities indicated that the POPAN models fitted the data among the three sites that are constant over time adequately (Table A2). This is in contrast with the (Table A1). The probability of residence is calculated \ results from the first analysis possibly due to the by subtraction (1 – ). lower observation effort in CAM, with 40 % of occasions with p = 0. Nevertheless we used the (Pradel et al. 1997b), i.e. the first resighting. The appropriate cˆ for each dataset to adjust model reduced data set consisted of 438 encounters of deviance (Table A2). For each seasonal analysis, 195 individuals in 30 week-long sampling the model with the highest AIC weight was occasions during the 2005-2010 winters. The retained (Table A3). second GOF test indicated that the multi-site type fitted the corrected data adequately and that the assumptions of the Jolly-Movement (JMV) model 2 were being met ( = 106·71, P = 0·99). Of the 15 F144 models available in this analysis (Table A1), four (ranked 1–4) had relatively low differences in AICc weight (less than 4 points, or > 0.1%). The main difference between models 1–4 lied in the structure of the transition probabilities, \ . Model 1 had the highest AIC weight (0.39), more than twice that of similar model 3 (0.17), so it was judged to be the most likely. This model assumes equal movement probability among the three states (but cf. models 2 & 6). Its encounter probabilities were pVNG = 0·40 (95% CI. 0·34–0·47) for the VNG site, similar to the one found in the single-site approach, and pCAM = 0·10 (95% CI, Fig. 3. Boxplot comparison of the results of counting 0·08–0·16) for the CAM site. The transition wintering Mediterranean Gulls present in VNG following probabilities of this model were \ = 0·22 (95% different methodologies: direct counts, by the same CI, 0·17–0·29) for all transitions among VNG, observer, of the number of gulls present at the time of each sampling occasion (n = 92); International Waterbird Census CAM and ‘NOS’ (fig. 2). Hence, an individual in (IWC) counts (n = 5); and average estimates of the super- VNG or CAM had a 44% monthly probability of population size Nˆ at the time of each sampling occasion, as derived from the POPAN models (n = 23). The width of each box is proportional to the sample size.

The geometric mean of the monthly encounter and 14 085 (95% CI, 9 133–19 037) in 2009/10, with probabilities p was 0·36 (range, 0·30–0·48) and the a geometric mean of 15 754 (95% CI, 10 630–20 geometric mean of the monthly survival 683). This represents 85% of the population probabilities, ϕ, was 0·86 (range, 0·80–0·92). The wintering in the Catalonia region (Josa et al. 2011), models provided monthly estimates of the 43% of the total population wintering in the NW

population size, Nˆ i as a derived parameter. Mediterranean and 18% of the global population Monthly estimates of the number of of the species (Cama et al. 2011, Wetlands Mediterranean Gulls visiting VNG each month International 2015), as based on direct counts and were consistently high across winters, with peaks the extrapolation of observed densities. between October and January (fig. 2a). Mean Discussion estimates of Nˆ (x = 5504, SE = 470, n = 23) were about six times higher than the direct counts of Our study looked into the fine-scale spatial birds present during sampling occasions in the dynamics of a wintering population, and it wintering period ( x = 886, SE = 63, n = 92) and addresses, for the first time for any gull species, from the International Waterbird Census (IWC) hypotheses on small-scale movements using counts, January 2006–2010 ( x = 990, SE = 298, n = individual based data. The results revealed that 5) (Fig. 3). Across winters, the average monthly site fidelity appears to be dependent on the spatial probability of entry between sampling occasions, scale considered. Individuals were faithful to the b, was 0·36 in Oct, 0·24 in Nov, 0·12 in Dec and wintering area on a regional scale (hundreds of 0·08 in Jan (fig. 2b). This is the probability that a km) but showed little site fidelity on a local scale Mediterranean Gull already present in the super- (tens of km). Despite showing strong site-fidelity population entered the VNG site for the first time at both to the breeding grounds (Marvelde et al. those points in time. 2009) and to the wintering area (Carboneras et al. The volume estimates of the total number of 2013), individual Mediterranean Gulls in this Mediterranean Gulls visiting VNG during a study were not likely to spend all winter in a wintering season, Nˆ *, were: 14 796 (95% CI, 9 single site; their monthly probability of moving 579–20 013) in 2005/06, 13 398 (95% CI, 10 135–16 away from a site was almost as high as that of 661) in 2006/07, 18 449 (95% CI, 15 411–21 486) in staying. When they moved, birds did not follow 2007/08, 18 842 (95% CI, 9 934–27 749) in 2008/09 any preferred direction, and many colour-ringed

Fig. 4a (left). Estimates of temporal super-population size ( Nˆ ) at the time of each monthly sampling occasion for the 5 seasons, 2005/06 – 2009/10, as derived from the POPAN models. The wintering period extends between 15 Sep and 15 Feb. Fig. 4b (right). Estimates of monthly entry probabilities β, as obtained from the selected POPAN models for the 5 seasons. Each point shows the probability that, in a four-week period, an individual present in the super-population N enters the VNG site for the first time in that season. The grey columns show the mean values for each month. individuals returned to one of the observation while food was still predictable at the site they localities after having spent time in the other, or were leaving. vice versa. Being mobile in winter implies In the POPAN model, the distribution of the b spending energy in travelling and perhaps having values of probability to enter the observed less detailed local knowledge. Despite the population, reflected the relative probability that energetic cost, small scale movements increase the a bird, ultimately a visitor to VNG, had been in chances of optimizing foraging strategies and of the general area without visiting the locality. This adapting to local availability of resources. probability was still high in December (12 %) and Our multi-site model highlighted the equal into January (8 %), which reflects the dynamism probability that, once individuals have moved, and the size of the wintering population they travel to one of the well-known wintering associated to the site. Monthly estimates of the net localities (shorter distance) or outside the study ˆ population size N fluctuated seasonally and did system (longer distance). Assuming that birds not fluctuate greatly between years (Fig. 2a). stayed in the same general area all winter However, the yearly peaks occurred at different (Carboneras et al. 2013), ‘elsewhere’ would mean months each year, reflecting that spatiotemporal any site on a 350-km long coastline. (Carboneras oscillations in the distribution occurred on a local et al. 2010) demonstrated that Mediterranean Gull scale, in the changing proportions of the populations distribute around the Iberian wintering population that spend time in VNG. Peninsula following a metapopulation pattern, Possibly through a combination of self-acquired clumping in wintering units >500 km apart that and socially-held knowledge, individual gulls do not share individuals other than through might become aware of the existence of foraging dispersal. Significant parts of the occupied opportunities at other sites within the general coastline are protected as MPAs or Natura 2000 wintering area. Conspecific density at the sites, on land or at sea (or both), but there are also destination point might, if known, also play an long stretches of unprotected suitable habitat unknown role in individual decision-making. The which the birds may use either as alternative sites, total number of birds present at the larger locality or during their travels. It is clear from our results (CAM) was on average ten times bigger than in that wintering in multiple sites on a local scale is a VNG, yet that did not seem to affect the common strategy in this species; we found that probability of movement in either direction. birds moved freely within their wintering areas, The two MPAs designated as Natura 2000 sites in on a spatial scale much larger than the average the study area for the conservation of several coastal MPA or Natura 2000 site. seabird species, including Mediterranean Gulls, Mediterranean Gulls follow trawlers offshore are separated on the coastline by a calcareous during the day, and into the harbours in the massif and comprise two different coastal evenings, to feed mostly on discards that are landscapes. They are sufficiently far apart to available only on weekdays (Cama et al. 2011). justify – to the human eyes – being considered as Food is less predictable during weekends, so we separate areas for the purpose of conservation, anticipated that changes of location would be and not as part of the same continuum. However, more likely then. To avoid that potential bias, we our study of Mediterranean Gulls revealed that excluded any weekend observations from our individuals in this species spent the wintering data and, as a consequence, our results reflect season not in one but in multiple localities, and changes that occurred, or were detected, only would travel relatively long distances even in the during weekdays. Thus, individuals in our study peak of the wintering season. There was no moved about and travelled >50 km distances obvious change in the environmental conditions

in any of the five seasons of our study to explain implications for standard management of the individual movements observed. Conversely, protected areas, such as MPAs and Natura 2000 the spatial dynamics of waterbirds such as ducks, sites. Our present results suggest that the geese, swans and waders, showed a tendency to population of Mediterranean Gulls wintering change site in the same season following a change along the eastern Spanish coast is occupying sites of conditions (Tamisier 1978, Hestbeck et al. 1991, outside the designed protected areas. On the other Swann et al. 2005, Luís and Goss-Custard 2005). hand, it seems unlikely to be able to protect all From the high mobility detected, we deduce that potential sites visited by the birds. Further most if not all Mediterranean Gulls in the research should be done to assess threats on wintering area would eventually spend time in unprotected sites. the current network of MPAs designed for its conservation. However, given the discontinuities Acknowledgements – We thank Joan Ferrer, Pere of the network and the mobility of the species, it is Josa and Marc Olivé for providing resightings also very likely that most individuals would data and Vittorio Pedrocchi for sharing data on equally spend time in places without protection. mid-winter gull counts. Susana Requena kindly Our study provides direct evidence of within- produced the map in Fig. 1. winter movement and as such it has direct

References Carboneras, C., Aymí, R., Cama, A., Duponcheel, C., Ferrer, J., Flamant, R., García, S., Garzón, J., Arnason, A. and Schwarz, C. 2002. POPAN-6: Gutiérrez, A., Olivé, M. and Poot, M. 2010. exploring convergence and estimate properties Mediterranean Gulls Larus melanocephalus with SIMULATE. - J. Appl. Stat. 29: 649–668. wintering in Spain and Portugal: one population or several? - Airo 20: 3–11. Atkinson, P., Choquet, R., Frederiksen, M., Gillings, S., Pradel, R. and Rehfisch, M. M. Carboneras, C., Tavecchia, G., Genovart, M., 2007. Towards developing thresholds for Requena, S., Olivé, M. and Oro, D. 2013. waterbirds that take into account turnover. Inferring geographic origin, population structure and migration routes of a wintering Boyd, C., Brooks, T. M., Butchart, S. H. M., Edgar, population of Mediterranean gulls from G. J., Da Fonseca, G. A. B., Hawkins, F., resightings data. - Popul. Ecol. 55: 343–351. Hoffmann, M., Sechrest, W., Stuart, S. N. and Van Dijk, P. P. 2008. Spatial scale and the Choquet, R., Lebreton, J.-D., Gimenez, O., conservation of threatened species. - Conserv. Reboulet, A.-M. and Pradel, R. 2009. U-CARE: Lett. 1: 37–43. Utilities for performing goodness of fit tests and manipulating CApture-REcapture data. - Burnham, K. P. and Anderson, D. R. 2002. Model Ecography 32: 1071–1074. selection and inference. A practical information-theoretic approach. - Springer. Cooch, E. G. and White, G. C. 2012. Program MARK: a gentle introduction. Cama, A., Josa, P., Ferrer-Obiol, J. and Arcos, J. M. 2011. Mediterranean Gulls Larus melanocephalus EEA 2014. Natura 2000 Standard Data Form - wintering along the Mediterranean Iberian ES5110020 Costes del Garraf. in press. coast: numbers and activity rhythms in the species’ main winter quarters. - J. Ornithol. 152: Frederiksen, M., Fox, A. D., Madsen, J. and 897–907. Colhoun, K. 2001. Estimating the total number of birds using a staging site. - J. Wildl. Manag. 65: 282–289.

Giroux, J.-F. 1991. Roost fidelity of Pink-footed Luís, A. and Goss-Custard, J. 2005. Spatial Geese Anser brachyrhynchus in north-east organization of the Dunlin Calidris alpina L. Scotland. - Bird Study 38: 112–117. during winter – the existence of functional units. - Bird Study 52: 97–103. Gourlay-Larour, M.-L., Pradel, R., Guillemain, M., Santin-Janin, H., L’hostis, M. and Caizergues, Martínez-Abraín, A., Viedma, C., Ramón, N. and A. 2013. Individual turnover in common Oro, D. 2001. A note on the potential role of pochards wintering in western France. - J. philopatry and conspecific attraction as Wildl. Manag. 77: 477–485. conservation tools in Audouin’s Gull Larus audouinii. - Bird Conserv. Int. 11: 143–147. Grist, H., Daunt, F., Wanless, S., Nelson, E. J., Harris, M. P., Newell, M., Burthe, S. and Reid, Marvelde, L. te, Meininger, P. L., Flamant, R. and J. M. 2014. Site Fidelity and Individual Dingemanse, N. J. 2009. Age-Specific Density- Variation in Winter Location in Partially Dependent Survival in Mediterranean Gulls Migratory European Shags. - PLoS ONE 9: Larus melanocephalus. - Ardea 97: 305–312. e98562. Newton, I. 2008. The migration ecology of birds. - Harris, M. P., Daunt, F., Newell, M., Phillips, R. A. Academic Press. and Wanless, S. 2009. Wintering areas of adult Atlantic puffins Fratercula arctica from a North Péron, G., Lebreton, J.-D. and Crochet, P.-A. 2010. Sea colony as revealed by geolocation Breeding dispersal in black-headed gull: the technology. - Mar. Biol. 157: 827–836. value of familiarity in a contrasted environment. - J. Anim. Ecol. 79: 317–326. 2000. Important Bird Areas in Europe: Priority Sites for Conservation (MF Heath, MI Evans, Pradel, R., Rioux, N., Tamisier, A. and Lebreton, J.- DG Hoccom, AJ Payne, and NB Peet, Eds.). - D. 1997a. Individual turnover among wintering BirdLife International. teal in Camargue: a mark-recapture study. - J. Wildl. Manag. 61: 816–821. Hénaux, V., Bregnballe, T. and Lebreton, J.-D. 2007. Dispersal and recruitment during population Pradel, R., Hines, J. E., Lebreton, J.-D. and Nichols, growth in a colonial bird, the great cormorant J. D. 1997b. Capture-recapture survival models Phalacrocorax carbo sinensis. - J. Avian Biol. taking account of transients. - Biometrics 53: 38: 44–57. 60–72.

Hestbeck, J. B., Nichols, J. D. and Malecki, R. A. Rees, E. 2006. Bewick’s Swan. - Poyser. 1991. Estimates of movement and site fidelity using mark-resight data of wintering canada Robertson, G. J. and Cooke, F. 1999. Winter geese. - Ecology 72: 523–533. Philopatry in Migratory Waterfowl. - The Auk 116: 20–34. Josa, P., Ferrer, J., Carboneras, C. and Cama, A. 2011. Gavina capnegra Larus melanocephalus. Ronconi, R. A., Lascelles, B. G., Langham, G. M., - In: Atles dels ocells de Catalunya a l’hivern Reid, J. B. and Oro, D. 2012. The role of 2006-2009. Institut Català d’Ornitologia, in seabirds in Marine Protected Area press. identification, delineation, and monitoring: Introduction and synthesis. - Biol. Conserv. Lebreton, J. D., Burnham, K. P., Clobert, J. and 156: 1–4. Anderson, D. R. 1992. Modeling survival and testing biological hypotheses using marked Schwarz, C. J. and Arnason, A. N. 1996. A General animals: a unified approach with case studies. - Methodology for the Analysis of Capture- Ecol. Monogr. 62: 67–118. Recapture Experiments in Open Populations. - Biometrics 52: 860–873.

Sergio, F., Tanferna, A., De Stephanis, R., Jiménez, Wetlands International 2015. Waterbird Population L. L., Blas, J., Tavecchia, G., Preatoni, D. and Estimates. Hiraldo, F. 2014. Individual improvements and selective mortality shape lifelong migratory White, G. C. and Burnham, K. P. 1999. Program performance. - Nature 515: 410–413. MARK: survival estimation from populations of marked animals. - Bird Study 46: 120–139. Swann, B. (R. L. ., Brockway, I. K., Frederiksen, M., Hearn, R. D., Mitchell, C. and Sigfússon, A. Williams, K. A., Frederick, P. C. and Nichols, J. D. 2005. Within-winter movements and site 2011. Use of the superpopulation approach to fidelity of Icelandic Greylag Geese Anser anser: estimate breeding population size: an example Capsule Levels of mobility and turnover in asynchronously breeding birds. - Ecology 92: changed at different stages of the non-breeding 821–828. season. - Bird Study 52: 25–36. Wilson, L. J., McSorley, C. A., Gray, C. M., Dean, B. Tamisier, A. 1978. The functional units of J., Dunn, T. E., Webb, A. and Reid, J. B. 2009. wintering ducks: a spatial integration of their Radio-telemetry as a tool to define protected comfort and feeding requirements. - areas for seabirds in the marine environment. - Verhandlungen Ornithol. Gesellshaft Bayern Biol. Conserv. 142: 1808–1817. 23: 229–238.

Appendix - Supplementary Material

Table A1. Modelling local survival and transition probabilities among multiple sites. Np = number of parameters, AICc = Akaike Information Criterion corrected for sample size, Δi = difference in AICc relative to the best model, wi = model weight, ϕ = survival, p = resighting probability, ψ = transition probability of entry. The selected model is shown in bold. Model notation: t = time, · = constant.

Table A2. Results of the goodness of fit (GOF) tests on the datasets used for the POPAN analysis. cˆ = variance inflation factor. Each dataset corresponded to a winter season and was analysed separately.

Table A3. Estimating total volume or super-population size ( Nˆ *) and entry probabilities β, in VNG site for the 5 seasons, 2005/06 – 2009/10 with POPAN. Np = number of parameters, QAICc = quasi- likelihood in AIC, Δi = difference in AIC relative to the best model, wi = model weight, ϕ = survival, p

= resighting probability, β = probability of entry, Nˆ *= super-population size. Selected models are shown in bold. Model notation: t = time, · = constant, · = parameter not used.

Table A1.

Rank Np AICc Δi wi Deviance ϕ p ψ

1 10 2137·79 0·00 0·39 1621·72 VNG=CAM=NOS (t) VNG,CAM,NOS (·) VNG=CAM=NOS (·)

2 12 2137·90 0·11 0·37 1617·64 VNG=CAM, NOS (t) VNG,CAM,NOS (·) VNG,CAM,NOS (·)

3 13 2139·49 1·70 0·17 1617·12 VNG=CAM=NOS (t) VNG,CAM,NOS (·) VNG,CAM,NOS (·)

4 13 2141·29 3·50 0·07 1618·93 VNG=CAM=NOS (t) VNG,CAM,NOS (·) VNGCAM=CAMVNG, CAMC=CCAM (·)

5 11 2160·22 22·43 0·00 1642·07 VNG,CAM,NOS (t) VNG,CAM,NOS (·) VNG,CAM,NOS (·)

6 9 2169·11 31·31 0·00 1655·12 VNG=CAM, C (t) VNG,CAM,NOS (·) VNG=CAM=NOS (·)

7 7 2175·61 37·81 0·00 1665·76 VNG=CAM (·), VNG,CAM,NOS (·) VNG,CAM,NOS (·) C (t)

8 12 2181·39 43·59 0·00 1661·13 VNG=CAM, C (t) VNG,CAM,NOS (·) VNGCAM=CAMVNG, CAMC=CCAM (·)

9 4 2191·63 53·84 0·00 1687·93 VNG=CAM=NOS (·) VNG,CAM,NOS (·) VNG=CAM=NOS (·)

10 6 2193·08 55·29 0·00 1685·30 VNG=CAM=NOS (·) VNG,CAM,NOS (·) VNG,CAM,NOS (·)

11 6 2193·52 55·73 0·00 1685·73 VNG,CAM,NOS (·) VNG,CAM,NOS (·) VNG,CAM,NOS (·)

12 9 2208·68 70·89 0·00 1694·69 VNG=CAM=NOS (t) VNG=CAM, C (·) VNG=CAM=NOS (·)

13 21 2335·54 197·75 0·00 1795·99 VNG=CAM=NOS (·) VNG,CAM,NOS (·) VNG,CAM,NOS (t)

14 39 79582·10 77444·31 0·00 79001·70 VNG,CAM,NOS (t) VNG,CAM (t), VNG,CAM,NOS (·) C (·)

15 80 79742·10 77604·31 0·00 79055·56 VNG,CAM,NOS (t) VNG,CAM,NOS (t) VNG,CAM,NOS (t)

Table A2.

no. GOF Season df P cˆ occasions Χ2 (test 2 + test 3)

2005/06 4 0·23 4 0·99 <1

2006/07 5 8·23 7 0·31 1.18

2007/08 5 11·66 6 0·07 1·94

2008/09 4 3·36 1 0.07 3.36

2009/10 5 5·92 4 0·21 1·48

Table A3.

Rank Np QAICc Δi wi ϕ p β Nˆ *(±SE) 2005/06 season 1 8 145·23 0·00 0·38 t · t 207 (±37) 2 6 145·61 0·38 0·32 · t t 3 5 146·86 1·62 0·17 · · t 4 9 147·40 2·17 0·13 t t t 2006/07 season 1 6 184·57 0·00 0·75 · · t 187 (±23) 2 10 187·84 3·27 0·15 t · t 3 8 189·05 4·48 0·08 · t t 4 12 191·95 7·38 0·02 t t t 2007/08 season 1 7 185·82 0·00 0·58 · · t 258 (±22) 2 8 186·74 0·93 0·36 · t t 3 11 191·55 5·74 0·03 t t t 4 10 191·94 6·12 0·03 t · t 2008/09 season 1 6 121·30 0·00 0·62 · · t 263 (±63) 2 7 123·02 1·72 0·26 · t t 3 8 125·31 4·01 0·08 t · t 4 9 127·64 6·34 0·03 t t t 2009/10 season 1 5 114·87 0·00 0·39 · · t 197(±35) 2 6 115·18 0·31 0·33 · t t 3 8 116·41 1·54 0·18 t · t 4 10 117·43 2·55 0·11 t t t

Chapter IV

A new breeding population of Mediterranean Gulls Larus melanocephalus in the species’ main wintering area maintains independent spatial dynamics

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Short communication ing good connectivity with the wintering areas may improve individual fitness during the non-breeding sea- son. In the Mediterranean Gull Larus melanocephalus, little is known about the demographics behind the spec- A new breeding population tacular expansion of its breeding range from SE to NW Europe during the 20th century. Initially confined to the of Mediterranean Gulls Black Sea coast, in the period 1965–90 it expanded 30° Larus melanocephalus in of longitude westwards across Europe to reach the Bal- tic, Atlantic and North Sea (Cramp & Simmons 1982, the species’ main Bekhuis et al. 1997). Breeding numbers in the Nether- lands, Belgium, NW France and UK (hereafter, ‘Atlantic’ wintering area maintains colonies) grew exponentially during the 1990s (to over independent spatial 2500 pairs in 2002, BirdLife International 2004), and continue to increase. This Atlantic population developed dynamics a westerly migration route, wintering mainly in NW CARLES CARBONERAS1,2,3* & JOSE IGNACIO DIES4 Spain, Portugal and in the Mediterranean as far east as 1Department of Animal Biology, University of Malaga (Fig. 1) (Poot & Flamant 2006, Carboneras et al. Barcelona, Barcelona, Spain 2010). 2Population Ecology Group, Institut Mediterrani In parallel, new breeding sites were established along d’Estudis Avancßats, IMEDEA (CSIC-UIB), Esporles, the western Mediterranean in France (first breeding Mallorca, Spain 1965, c. 3600 breeding pairs in 2010; Cadiou 2011), 3Royal Society for the Protection of Birds, The Lodge, and Italy (first breeding 1978, 2000–2100 pairs in 2002; Sandy, Bedfordshire SG19 2DL, UK Serra & Brichetti 2005). Recently established colonies in 4Servici Devesa-Albufera, Ajuntament de Valencia, the Valencia region of eastern Spain (Dies & Dies Valencia, Spain 2004), initially assumed to be part of the same process, represented a significant expansion of the breeding range towards the southwest. That population rose from a sin- gle pair that attempted breeding in 2001 to 447 breed- We studied a newly established breeding population of ing pairs in four colonies in 2014. the range-expanding Mediterranean Gull Larus melano- The Spanish Mediterranean coast has traditionally cephalus in eastern Spain, situated in close proximity to been the main winter quarters of the species, currently et al. the species’ main wintering area. By investigating the holding over 50% of the global population (Cama origin, population composition and wintering area of the 2011). Those birds return to breed mostly along the new breeders, we found that recruitment from locally Mediterranean and Black Sea coasts. In this context, the wintering birds was unlikely and that the emerging colo- emergence of a new breeding colony provided a rare nies were probably attracting birds from populations opportunity to study the relation between a long-stand- wintering 700–1200 km away in Portugal and southern ing, spatially stable wintering aggregation, formed exclu- Spain. Our findings reveal that expanding populations sively of migratory birds, and a locally breeding may follow their own dynamics, independently of other population of the same species. To elucidate whether fl populations of the same species, and may consist of dif- one population or strategy in uenced the other, we ferent individuals altogether. studied the source population of any new recruits, the wintering area of the new breeders and the degree of mixing between the two populations. Keywords: breeding range, expanding species, migration, population ecology, recruitment, METHODS wintering area. We collected and analysed data on the location of Mediterranean Gulls wearing individually coded colour Expanding populations of migratory birds face novel rings on the Mediterranean coast of Spain. Our breeding challenges as they establish themselves in new areas. A population dataset consisted of all the birds observed at tendency to return to the traditional wintering grounds and around the breeding colony established at the Raco should be expected, at least initially, because maintain- de l’Olla reserve, Parc Natural L’Albufera near Valencia, Spain (39°200N, 0°190W). This area, monitored daily since the 1990s, held 275 nesting pairs of Mediterranean *Corresponding author. Gull in 2012. Forty-two birds of known age and birth- Email: [email protected] place were recorded visiting, displaying or nesting at the

© 2015 British Ornithologists’ Union 2 C. Carboneras & J. I. Dies

20°W 0° 20°E 40°E 45°N 45°N

Wintering 30°N Breeding - consolidated

30°N Breeding - other 0200400600km Valencia breeding site CRS: ETRS89-LAEA Europe

0° 20°E

Figure 1. Distribution of breeding and wintering Mediterranean Gulls plotted according to the European Ornithological Atlas 50 9 50 km grid, updated from Bekhuis et al. (1997). Consolidated breeding sites held > 25 breeding pairs for > 3 years; shown win- tering cells regularly held > 50 individuals. The dotted line indicates the likely migratory route of the western European (‘Atlantic’) population southwards to Iberia. colony during the breeding season (April to June) in before moving to roost at sea; no birds were handled 2003–2012. during our study. Our wintering season dataset contained observations To assess whether the newly established population of colour-ringed Mediterranean Gulls recorded during was growing as a result of its own reproductive output five winter seasons (15 September to 15 February) in or of immigration, we followed the count-based popula- 2005–2010 (625 individuals; 2925 ring-readings), in the tion viability analysis (PVA) methodology described in 350-km-long stretch of coast between the towns of Morris and Doak (2002). We fitted a linear regression to Blanes (41°410N) and Burriana (39°540N) (Fig. 2), the the number of pairs counted during the annual census winter stronghold for the species (c. 40 000 birds; Cama of breeding waterbirds in the Valencia region in 2001– et al. 2011) and known since at least the 1940s (Mayaud 2014 and assumed that the number of nests was equal 1954). The distance between this area and the Valencia to the number of females. We used the slope of the colony site is approximately 65 km. regression line as an estimate of the annual growth rate The species is subject to an international colour-ring- (l) and the mean squared residual as an estimate of its 2 ing programme comprising all of its known breeding variance (r ) (Morris & Doak 2002). populations. All birds in our dataset had been individu- We inferred the origin of the recruiters to the breed- ally marked in breeding colonies outside Spain prior to ing colony by comparing its population composition their arrival in the study area. Observations were made (ratio of birds born in colonies west and east of from a distance with a 20–609 telescope when Gulls Gibraltar) with the local wintering population, described concentrated on beaches and around harbour premises in detail in Carboneras et al. (2013). Furthermore, we

© 2015 British Ornithologists’ Union Dynamics of expanding Mediterranean Gull populations 3

10°W 0° 10°E 50°N

! Natal colony, 2004 50°N 40°N Blanes

Burriana Dec 2005 Breeding Valencia 40°N 2009, 2011, 2012

Jan 2010 Oct 2011 Jul 2009 Jan 2013 Jan 2009

km 0 100 200 300 CRS: ETRS89-LAEA Europe

10°W 0° 10°E

Figure 2. Life history locations of colour-ringed Mediterranean Gull ‘Green 30VL’ born in Belgium (2004), recorded wintering in Por- tugal and Spain (2005–2013) and present as a breeding bird in Valencia (2009–2012). Grey cells indicate the species’ general winter- ing areas as in Figure 1. were able to compare the same ratio with that of other The natal colonies of the two sets of individuals dif- discrete Mediterranean Gull populations wintering in fered substantially. Although only about 1.5% of the the Iberian Peninsula, as it is known to follow a west to Mediterranean Gulls in the wintering area originated east gradient (Carboneras et al. 2010). from Atlantic colonies (Carboneras et al. 2013), they made up 57% of the breeders and visitors to the Valen- cia nesting site (Table 1). In contrast, birds born in RESULTS Mediterranean French and Italian colonies, which form The average annual growth rate, l, estimated by the two-thirds of the wintering population, only represented PVA was 0.47 and its variance, r2, 0.22 (P < 0.01). This 26% of the summering birds. represents an annual increase of 47%. The location of the wintering grounds of the Valencia The two sets of 42 summering and 625 wintering breeding birds remains largely unknown. Only one indi- individually marked Mediterranean Gulls contained no vidual, Belgium-born ‘Green 30VL’, provided numerous individual common to both sets, despite occurring in summering and wintering locations in 2004–2013. It close proximity. None of the breeding birds was ever moved between the Valencia breeding colony and win- recorded in the area during the winter months, and none tering grounds in SW Portugal and Malaga, southern of the marked wintering birds was ever recorded at the Spain (Fig. 2). Valencia nesting colonies during the breeding season. Individual encounter histories and field observations DISCUSSION showed that the wintering area was vacated completely by late March every year, as birds left for their breeding The emergence of a breeding population near a tradi- grounds. A few weeks later, the Valencia breeders were tional wintering site offered a unique opportunity to back at the nesting site in mid-April and May. study potential changes in the spatial dynamics of those

© 2015 British Ornithologists’ Union 4 C. Carboneras & J. I. Dies

Table 1. Population assignment of the 42 colour-ringed birds A yearly increase of 47% can only occur through observed (‘obs’, in bold) at the breeding colony in Valencia, large-scale immigration. Using a Leslie matrix model and eastern Spain, in 2003–2012 compared with the expected demographic data, Sadoul (1997) demonstrated that the number of individuals if the population followed the same pro- 20% annual increase he observed in the Mediterranean portions observed in the locally wintering population by Car- Gull colony in Camargue, SE France, was due to immi- boneras et al. (2013). There is a highly significant gration. Otherwise, it would have required an unrealistic overrepresentation of birds from ‘Atlantic’ colonies (ATL), indi- cating the probable origin of the summering birds, and an mean breeding success of at least 1.6, sustained over an underrepresentation of birds from Italy (ITA), the French extended period. The observed average productivity in Mediterranean (MED), and eastern European colonies in Hun- France was consistently lower (1.0–1.6 young per breed- gary, Greece, Ukraine and Poland (E Eur). ing pair in predator-free colonies but 0.1–0.5 in dis- turbed colonies; Isenmann et al. 2004). Assuming that Fisher the productivity values in Valencia were not different E exact from the French colonies, we infer that only external MED ITA Eur ATL Total test immigration could explain their increase. The distribution of Mediterranean Gulls in winter Breeders + exp 3 3 3 0 9 0.02 was shown to follow a probability function of distance prospectors obs 1116 – Visitors exp 12 10 11 0 33 < 0.001 along the main northeast southwest route (Carboneras et al. obs 36618 2013); thus, in the main wintering area in E-NE Total exp 15 13 14 0 42 Spain, birds were most likely to come from colonies in obs 47724 nearby France and Italy, and secondly from the eastern Mediterranean. This study reveals, in strong contrast, that the recent colonization has been extending in the populations as a result of their physical proximity. The opposite direction, from west to east, and that it has new breeding colony of Mediterranean Gulls was able to consisted mostly of birds born in Atlantic colonies that grow very quickly by attracting immigrants. Unexpect- penetrated and established themselves in the western edly, those birds did not come from the spatially stable Mediterranean. population wintering in the vicinity. Instead, our data Our findings reveal that the large-scale expansion of suggest a more distant origin. a species’ range may involve only a fraction of all the Differences in the geographical composition of both individuals available, and that different populations may populations were considerable. Of all Mediterranean follow independent spatial dynamics, even where they Gulls ringed up to 2009, the proportion of Atlantic vs. co-occur. The birds forming part of the spatially stable Mediterranean birds on a global scale was 40 : 60; wintering population did not take advantage of the among the populations wintering in the Iberian Penin- emergence of new breeding opportunities nearby and sula, that proportion followed a west to east gradient: consistently migrated to their distant breeding grounds. 73% of the Mediterranean Gulls in Galicia–NW Spain; In parallel, the new breeders continued to behave as if 66–67% in W Portugal; 40% in Malaga but only 3–6% the important wintering area in the vicinity did not in E-NE Spain were of Atlantic origin (Carboneras et al. exist, and probably returned to their traditional areas. 2010). The proportion of Valencia breeders and visitors This reveals a strong tendency for populations to main- born in Atlantic colonies (57%) did not match that of tain their momentum and spatial strategies, and suggests the population wintering in E-NE Spain, pointing to a that shifts in distribution (e.g. those expected as a result more likely origin of those birds from either the popula- of a changing environment) may not necessarily be sim- tion wintering in Portugal (over 1200 km away) or from ple, linear processes. Malaga (over 700 km). Interestingly, all of the wintering records of Green 30VL came from those two regions. We thank Giacomo Tavecchia, the editors Ruedi Nager and Ross We put considerable effort into ring-reading in win- Wanless, as well as two anonymous reviewers, for providing valu- ter. A previous study on a subset of our data (Carbon- able comments that improved an earlier version of the manu- eras et al. 2013) estimated an average annual resighting script. Susana Requena kindly produced the maps in Figures 1 and 2. Miguel Chardı helped with survey work at the breeding probability of 0.49 (95% confidence interval (CI) 0.42– site, as a part of the management tasks carried out on behalf of 0.55). This is the probability that an individual, being Servici Devesa-Albufera of the city council of Valencia, under alive and in the area, was recorded at least once during the guidance of Amelia Quintana and Joan Miquel Benavent. the winter season. The inverse probability (that the bird wintered in the area but was not detected) would be 0.51. Therefore, of our set of 42 summering birds, we REFERENCES would expect to see around 21 at least once every win- Bekhuis, J, Meininger, P.L. & Rudenko, A.G. 1997. ter. However, none was seen, so we deduce that they Mediterranean Gull Larus melanocephalus. In Hagemeijer, were more likely to have wintered somewhere else.

© 2015 British Ornithologists’ Union Dynamics of expanding Mediterranean Gull populations 5

E.J.M. & Blair, M.J. (eds) The EBCC Atlas of European Dies, J.I. & Dies, B. 2004. The breeding biology of Breeding Birds: Their Distribution and Abundance: 903. Mediterranean Gull at l’Albufera de Valencia (Western London: T. & A.D. Poyser. Mediterranean). Ardeola 51: 221–224. BirdLife International 2004. Birds in Europe: Population Isenmann, P., Sadoul, N., Walmsley, J. & Yesou, P. 2004. Estimates, Trends and Conservation Status. Cambridge: Mouette melanocephale Larus melanocephalus. In Cadiou, BirdLife International. B., Pons, J.-M. & Yesou, P. (eds) Oiseaux Marins Nicheurs Cadiou, B. 2011. Cinquieme recensement national des de France Metropolitaine: 1960–2000:92–96. Meze oiseaux marins nicheurs de France metropolitaine 2009– (Herault): Biotope Editions. 2011, 1ere synthese: bilan intermediaire 2009–2010. Brest: Mayaud, N. 1954. Sur les migrations et l’hivernage de Larus Gisom et AAMP. melanocephalus Temminck. Alauda 22: 225–245. Cama, A., Josa, P., Ferrer-Obiol, J. & Arcos, J.M. 2011. Morris, W.F. & Doak, D.F. 2002. Quantitative Conservation Mediterranean Gulls Larus melanocephalus wintering along Biology: Theory and Practice of Population Viability the Mediterranean Iberian coast: numbers and activity Analysis, 1st edn. Sunderland, MA: Sinauer Associates. rhythms in the species’ main winter quarters. J. Ornithol. Poot, M. & Flamant, R. 2006. Numbers, behaviour and origin 152: 897–907. of Mediterranean Gulls Larus melanocephalus wintering Carboneras, C., Aymı, R., Cama, A., Duponcheel, C., along the west coast of Southern Portugal. Airo 16:13–22. Ferrer, J., Flamant, R., García, S., Garzón, J., Gutierrez, Sadoul, N. 1997. The importance of spatial scales in long- A., Olivé, M., & Poot, M. 2010. Mediterranean Gulls Larus term monitoring of colonial Charadriiformes in Southern melanocephalus wintering in Spain and Portugal: one France. Colon. Waterbirds 20: 330–338. population or several? Airo 20:3–11. Serra, L. & Brichetti, P. 2005. Popolazioni di Uccelli Acquatici Carboneras, C., Tavecchia, G., Genovart, M., Requena, S., Nidificanti in Italia – Resoconto 2002. Avocetta 29:41–44. Olive, M. & Oro, D. 2013. Inferring geographic origin, population structure and migration routes of a wintering Received 15 May 2015; population of Mediterranean Gulls from resightings data. revision accepted 18 October 2015. Popul. Ecol. 55: 343–351. Associate Editor: Ross Wanless. Cramp, S. & Simmons, K.E. 1982. The Birds of the Western Palearctic, 3rd edn. Oxford: Oxford University Press.

© 2015 British Ornithologists’ Union

DISCUSSION

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56 Fig. 10. Schematic representation of the major migratory routes of Mediterranean Gulls from the breeding colonies to the main wintering areas in NE Spain and Málaga, S Spain (green lines). The spring migration is probably along the same routes, except in a minority of cases (dashed blue lines) of birds that likely return from Málaga following a more direct northward route.The current model predicts the future occurrence of resighitngs inland, along the main rivers and sometimes far from seashores (especialy in shouthern Germany, Austria and in other Eastern European countries).

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58 

Fig. 11. Blue [A24] (right), born in Italy in 1990, was the oldest known individual of the species when it was last recorded in the study system in 2011. It is seen here next to Black [P8K] (left), ringed as a fully-grown adult at a nesting colony in the Black Sea. Notice the partly broken colour-ring of the latter, which it eventually lost completely; since then, it was only identifiable through its metal ring (not visible in this picture). Photo: Carles Carboneras.

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Fig. 12. Evidence of actual threats faced by Mediterranean Gulls in the study system: above, adult bird with amputated foot, probably as a consequence of interaction with a fishing vessel; middle, severe instance of oiling in a second-winter bird; below, adult bird with swallowed hook; the line can be seen protruding from its mouth. Photos: Carles Carboneras.





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6 82 0 0 60 0 88 118 124 300 2007 155 237 230 891 230 237 0 0 6 0 0 0 1 0 0 0 1 0 0 0

4 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 11 236 249322 0 30 0 0 30 0 0 28 11 21 0 3 0 0 3 21 26 74 56 47 56 74 26 163 125 104 122 2006 140 IO\LQJELUGVRIDQ\DJH 253 0 0 0 253 448 325 71 57 71 325 448 0 7 0 6 8 0 4 56 0 0 5 5

6 0 40 189 292147 6 3 0 97 0 0 0 0 0 0 0 14 35 78 19 31 19 78 35 14 42 38 162 141 2005 0 0 0 0 0

0 0 0 0 50 0 11 0 10 0 29 20 57 9 8 10 26 10 9 8 57 20 54 240 2004 0 0 2 0 2 2

0 0 0 6 32 0 15 20 53 178 300 354 2003 1 0 0 0 2 0

0 0 0 3 59 0 0 19 12 684 123 2002 FKLFNVಫIOHGJHGಬ IXOO 1544 0 0 4 0

0 0 0 0 0 83 0 0 19 14 20 30 102 123 2001 0 0 3

0 0 1 0 0 6 1 0 13 35 66 12 49 566 148 2000 0 0

0 0 0 0 0  2 0 0 0 26 84 67 44 19 975 118 1999 

0 0 0 0 0 222 0 0 0 233 0 0 0 222 0 0 0 0 0 0 0 0 0 0 0 15 2 0 15 0 0 0 0 0 0 0 6 0 0 4 0 0 0 0 0 0 0 0 0 0 0 0 40 12 2 9 8 22 14 9 21 9 21 9 14 22 2 9 8 12 35 208 205 300 1998 ಫ3XOOLಬ XQIOHGJHG

0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 90 0 0 0 0 0 0 0 90 0 0 0 0 47 26 8 22 144 72 105 52 181 23 172 60 22 21 43 95 103 646 324 31 131 249 131 31 324 646 103 95 43 21 14 0 15 0 4 0 31 0 0 31 0 4 0 15 0 14 79 225 107 77 0 137 0 100 80 100 0 137 0 77 107 225 79 160 208 263 43 22 80 197100 139 74 165 458 281264 87 226281 266 145 1997 \

0 0 0 0 0 34 9 20 0 5 21 0 0 0 0 0 0 0 0 3 9 3 12 13 12 0 0 0 0 3 9 3 0 0 0 0 0 0 0 8 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 19 0 0 0 0 0 0 0 0 30 71 139 1996

0 0 0 0 0 9 0 0 0 0 0 0 0 0 0 0 56 78 1995

0 0 0 0 0 1 0 0 0 1 0 0 0 0 0 18 98 292 1994

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 66 109 1993

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 15 27 30 1992

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 23 42 1991 0HGLWHUUDQHDQ*XOOVSHUFRXQWU

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 35 1990

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1989

chick 0 0 0 0 0 0 0 0 0 chick chick 0 0 0 0 0 0 3 5 chick 0 0 0 0 0 0 0 0 0 chick chick chick chick 0 0 0 0 0 0 0 0 0 chick chick 0 0 0 0 0 107180 204 chick chick chick chick chick 0 0 0 0 0 0 0 chick flying flying chick 0 0 0 0 0 0 3 chick flying flying 1 0 1 0 0 1 0 0 1 0 0 1 0 1 flying flying flying 0 24 11 11 0 14 3 11 0 24 flying flying chick 0 104167 193146 66 28 35 flying flying 0 0 0 0 1 0 1 45 0 0 0 0 1 0 1 flying chick chick chick chick chick chick flying chick chick chick chick flying chick chick chick 0 0 0 0 0 0 0 0 chick flying chick 0 0 0 0 0 0 0 chick flying 0 0 0 0 0 0 0 0 0 0 0 0 0 0 flying flying 0 0 0 0 0 0 0 0 0 flying flying 0 0 0 0 0 0 0 0 0 0 0 0 0 0 flying flying 0 0 0 0 0 0 0 0 0 0 0 0 0 0 flying flying 0 0 0 0 0 4 0 0 4 0 0 0 0 0 flying YEAR

 UK Italy Spain Poland Ireland Russia Turkey Turkey TOTAL TOTAL Greece Norway Ukraine Belgium Hungary Hungary Slovakia Denmark Romania Germany Czech Rep.Czech Netherlands France-MED Serbia-Mont. Serbia-Mont. France ATL - 7DEOH1XPEHURIFRORXUULQJHG