Protected areas under pressure: decline, redistribution ...Giacomo Tavecchia2, Fernando Hiraldo1 1Department of Conservation Biology, Estación Biológica de Doñana − CSIC, C/Americo

ENDANGERED SPECIES RESEARCHEndang Species Res

Vol 38 189ndash204 2019httpsdoiorg103354esr00946

Published April 25

1 INTRODUCTION

Protected areas represent one of the pillars of theglobal conservation strategy (Watson et al 2014Beissinger et al 2017) Despite some recent debateover the breadth of their function (Naughton-Treveset al 2005) one of the main objectives of protected

areas is the protection of biodiversity and the con-servation of viable populations of imperiled species(eg IUCN 1994) However mounting pressureswithin and around protected areas often challengesuch goals (Hansen amp DeFries 2007) and after aperiod of necessary focus on reserve establishmentmore and more attention is being devoted to moni-

copy The authors 2019 Open Access under Creative Commons byAttribution Licence Use distribution and reproduction are un -restricted Authors and original publication must be credited

Publisher Inter-Research middot wwwint-rescom

Corresponding author fsergioebdcsices

Protected areas under pressure decline redistribution local eradication and projected extinction of a threatened predator the red kite in Dontildeana National Park Spain

Fabrizio Sergio1 Alessandro Tanferna1 Javier Chicano1 Julio Blas1 Giacomo Tavecchia2 Fernando Hiraldo1

1Department of Conservation Biology Estacioacuten Bioloacutegica de Dontildeana minus CSIC CAmerico Vespucio 26 41092 Seville Spain2Animal Demography and Ecology Unit Institute for Mediterranean Studies (IMEDEA) CSIC-UIB 07190 Esporles Spain

ABSTRACT After a period of overfocus on the establishment of reserves attention is increasinglybeing devoted to the capability of protected areas to maintain viable populations of endangeredspecies Here we examined the trends and reproduction of the red kite Milvus milvus a highlyendangered raptor near-endemic to Europe to illustrate the dual benefits and challenges faced bya national park to protect this iconic species Over the past 4 decades the kite population of south-ern Spain has declined steeply and has become progressively confined to Dontildeana National Parkand its buffering Natural Park areas Population deterioration was also evident within the pro-tected area through (1) spikes of rapid eradication of whole sub-populations from buffer areaslikely propelled by illegal poisoning and (2) more gradual but steady deterioration of numbersand reproduction especially in peripheral buffer areas probably caused by the interplay of sev-eral shocks related to food availability habitat degradation competition predation and chemicalcontamination The result was a 46minus55 decline with progressive confinement to the core Nat -ional Park and an alarming effective population size of lt10 pairs Demographic modelling sug-gested low adult survival and predicted further declines with possible extinction over the next 2decades We outline tentative goals for management but these will need urgent information onranging and mortality to provide more efficient targets These results illustrate how establishmentof a large park can prevent regional extinction but not necessarily guarantee species-safety lead-ing to protracted forms of extinction debt We suspect that similar dynamics will become morewidespread as anthropogenic pressures increase around protected areas and their performancemonitoring becomes more prevalent

KEY WORDS Matrix models middot European estimates middot Farmland intensification middot Protected area performance middot Reserves middot Review middot Population growth rate

OPENPEN ACCESSCCESS

Endang Species Res 38 189ndash204 2019

toring whether protected areas actually deliver bio-diversity benefits ie their performance (Gaston etal 2008) For example some studies have reporteddeclines and extinctions of key species in protectedareas and sometimes steeper declines within pro-tected areas than outside them (eg Brashares et al2001 Parks amp Harcourt 2002 Whitfield et al 2007Craigie et al 2010) This calls for the need to assessthe historical levels trends and viability of threat-ened species within protected areas in order toensure that they are properly preserved that theycan act as sources of colonizers for outside areas orthat potential declines are detected and haltedthrough management

Here we report a representative example of thebenefits and challenges provided and faced by pro-tected areas in attaining their biodiversity mainte-nance function In particular we focus on the long-term population dynamics of a highly endangeredavian predator the red kite Milvus milvus breedingin Dontildeana National Park in southern Spain

Dontildeana is one of the most biodiversity-rich parks inEurope currently listed as a Ramsar Site a WorldHeritage Site and a UNESCOrsquos Man and the Bio-sphere Reserve While it is usually lands of low eco-nomic value that are designated as protected areas(eg Scott et al 2001) Dontildeana is a relatively largepark (gt1000 km2) centered around a seasonal wet-land in prime fertile land along a river estuary Whileproviding clear biodiversity benefits its location onvery productive land has caused an exponentiallygrowing anthropogenic pressure at the park bordersespecially by rapid habitat conversion to intensivefarmland with consequent impacts on the watertable that sustains the reserve wetlands The park isalso famous worldwide for its dense populations ofcharismatic vertebrate predators which represent amajor attraction for over 300 000 annual tourists anda key focus for conservation management (eg Ferreramp Hiraldo 1991 Loacutepez et al 2009)

One component of such a predator assemblage isthe red kite a medium-sized raptor endemic to thewestern Palearctic with 95 of its world populationconfined to Europe (Knott et al 2009) This speciestypical of open landscapes with scattered woodlandhas suffered dramatic declines in recent decades inall its stronghold populations of Germany Franceand Spain which hold about 80 of the world popu-lation (Knott et al 2009) Due to such declines thered kite is currently classed as Near Threatened inthe IUCN Red List and included in Annex I of the EUBirds Directive and in Annex II of the Bern Conven-tion on the Conservation of European Wildlife and

Natural Habitats In Spain which holds the worldrsquosthird largest population the species has been clas -sified as lsquoat risk of extinctionrsquo since 2011 and as lsquoatcritical risk of extinctionrsquo within southern Spain(Andaluciacutea Autonomous Region) where the presentstudy was conducted In Dontildeana the breeding popu-lation is concentrated near the marshes (Sergio et al2005) where its generalist and opportunist diet isdominated by prey species typical of the marshes andof the ecotone between wetland and terrestrial habi-tats (eg rabbits Oryctolagus cuniculus waterbirdscarrion and reptiles Delibes amp Garciacutea 1984 Veiga ampHiraldo 1990) The preservation of this population isparticularly important for 3 main reasons (1) It is thelast reservoir of the species in southern Spain (seeSection 31) and thus a strategic key for avoiding fur-ther range contraction (a major goal of the EuropeanAction Plan for this species Knott et al 2009) (2) Itshigh dependence on a seasonal wetland makes this aunique population (Vintildeuela et al 1999 Seoane et al2003) with special adaptations to an unpredictablenatural disturbance regime of marsh inundationwhich would be lost in case of extinction with theconsequent erosion of functional biodiversity (3)During winter large numbers of red kite migrantsfrom central-northern Europe join the resident breed-ing population which can thus act as a sentinel formortality threats potentially impacting far away pop-ulations through carryover effects

Given all the above we reconstruct the historicalregional distribution of the red kite in southernSpain document its progressive extirpation and con-finement to Dontildeana demonstrate long-term declinesin abundance and performance of the park popula-tion across several decades provide projections offuture extinction risk and identify potential threats inurgent need of investigation and management

2 METHODS

21 Study area

The protected area of Dontildeana (hereafter Dontildeana)is located in southwestern Spain (36deg 56rsquo 51rsquorsquo N 6deg 21rsquo31rsquorsquo E) within the Autonomous Region of AndaluciacuteaIt is composed of 542 km2 of National Park institutedin 1969 and 538 km2 of buffering Natural Park witha less strict protection category instituted in 1989(Fig S1 and Table S1 in the Supplement at www int-res com articles suppl n038 p189 _ supp pdf) The Nat -ional Park is dominated by 5 main habitat types(1) seasonal marshland usually flooded during win-

190

Sergio et al Predator extinction debt in national park

ter and progressively drying during the springminus summer (2) scrubland a mixture of different degra-dation stages of autochthonous Mediterranean scrub -land (3) sparse grassland mainly located along theinterface between Mediterranean scrubland and themarshes (hereafter lsquoverarsquo) (4) mobile sand dunesalong the ocean shore and (5) stone pine (Pinuspinea) plantations Mature trees suitable for nestingkites are mainly present in the latter pinewoods andas isolated or small clumps of cork oaks Quercussuber and Eucalyptus trees The portions of the Nat-ural Park relevant for breeding kites are mainly thenorthern and eastern sectors of extensive pinewoodssurrounded by intensive farmland All farmland inand around Dontildeana has become progressively moreintensive especially with the spread of strawberrycultivation since the 1990s

22 Field procedures

Kite territories were censused by looking at territo-rial displays nest material transfers and presence ofbirds perched multiple times in or near their nest dur-ing pre-incubation in FebruaryminusMarch Wheneverpossible nest contents were checked at least 3 times(1) during incubation to assess clutch size (2) justafter hatching to estimate hatching success andbrood size and (3) when the nestlings were ~45 d oldto record the number of reared young (nestlingsfledge at ~55 d Bustamante 1993) Because kites areextremely vulnerable to poisoned baits (Villafuerte etal 1998 Vintildeuela et al 1999) we collected all avail-able data on kites found poisoned in Dontildeana Theseincluded (1) individuals reported to local authoritiesand confirmed as poisoned by subsequent eco-toxi-cological analysis and (2) individuals found dead inor under the nest Based on post-mortem chemical ana -lyses such cases are extremely likely to be caused bypoisoning (details in Hernaacutendez et al 2001)

23 Historical information

Kites were intensively surveyed until being rea-sonably sure to have detected the territorial pairs ofthe whole park since 2012 in collaboration with thepark authority and the Equipo de Seguimiento deProcesos Naturales of the Estacioacuten Bioloacutegica deDontildeana-CSIC (EBD-CSIC) which together surveyedthe peripheral sectors of Marismillas Sanlucarpinewood and Abalario (Fig S1) Before that inten-sive research on red kites was conducted in Dontildeana

in the 1980s and 1990s by a group led by F Hiraldoand subsequently by a group led by F Sergio and FHiraldo (eg Veiga amp Hiraldo 1990 Sergio et al2005) Work by these research groups focused espe-cially on certain portions of the park and thus inte-grated extensive information provided by (1) theEquipo de Seguimiento de Procesos Naturales of theEBD-CSIC (2) the Grupo de Conservacioacuten of Dontilde -ana National Park and (3) the field diaries of dozensof researchers field workers visitors volunteersgamekeepers park managers and wardens active inDontildeana since the 1960s Integrating all these sourcesespecially old field diaries from the 1960sminus 70srequired 2 yr of careful screening of locational infor-mation For example two different field ob serverssometimes visited the same territory in the same yearbut referred to the site using different local nameswhich often required meticulous diary inspection toreconstruct the route travelled by each observer onthat day In most years the census was not availablefor the whole park and the well surveyed sectors var-ied among years depending on research priorities orthe area of action of specific observers (eg wardensor field technicians deputed to certain sectors insome years and others in subsequent years) Becauseof such fragmented surveys we estimated the totalDontildeana population by decade as follows (1) wedivided the park into 12 sectors (Fig 1b) These weredefined on the basis of major differences in land-scape composition relevant for kite ecology (egextensive pinewoods and farmland in the north of thepark vs marsh ecotone in the park center) and delin-eations of private properties be cause these oftencaused abrupt management- driven differences inlandscape and because field workers were oftenassigned to survey whole properties in differentyears (2) For each well surveyed sector we calcu-lated the mean number of pairs per year for eachdecade (3) For each decade we summed up thesemean values to obtain a cumulative estimate for thatdecade (4) In the few cases in which the first com-plete survey of a sector was carried out late (eg inthe 1990s) we assumed that the population was stable in earlier decades (hereafter lsquostatic back- estimatesrsquo) This was done in order to enable a com-plete estimate of the whole population through alldecades and produced conservative estimates ofdeclines (5) In the few cases when complete surveysof a sector were not available for a given decade butthey were for earlier and later decades these wereinterpolated to estimate the missing value (hereafterlsquointerpolated estimatesrsquo) Finally (6) in a few periph-eral sectors complete surveys were available for cer-

191


tain portions of a sector in some years and other por-tions in different subsequent years In these cases wecalculated the mean number of pairs yrminus1 decademinus1 foreach subsector when well surveyed and summed upthese means to obtain a cumulative estimate de -cademinus1 for the whole sector (hereafter lsquocross-combinedestimatesrsquo) In all procedures above we ensured thatall alternative nests belonging to the same territorywere included within the same sector or subsector inorder to avoid biases caused by re-counting the sameterritory twice when combining different surveyunits Similarly in all cases of doubt we used the

estimate that minimized the historicalpopulation in order to provide conser-vative estimates of decline

To place declines and the potentialstrategic importance of the parkwithin a wider regional context wereconstructed the distribution andpopulation levels of the kite popula-tion for southern Spain (AndaluciacuteaAutonomous Region 87 268 km2) inthe 1970sminus1980s based on informa-tion found in Otero et al (1978) TorresEsquivias et al (1981) Ceballos ampGuimeraacute (1992) Vintildeuela et al (1999)Vintildeuela (2004) and conversationswith local experts Data for the currentred kite population outside Dontildeanawere provided by the Consejeriacutea deMedio Ambiente of the AndaluciacuteaAutonomous Region

24 Statistical analyses

To investigate how generalized thehistorical population declines acrossdifferent sectors of the park were weestimated whether the subpopulationof each sector increased or declinedbetween the earliest available surveyand the 2010s and then tested whetherdeclines were prevalent over stable orincreasing trends by means of a bino-mial test To explore temporal trendsin breeding rates we fitted general-ized linear models (GLMs) with de -cade sector and their interaction asexplanatory variables and with thelikelihood of laying eggs clutch sizehatching success brood reductionthe likelihood of successfully raising

chicks to fledgling and the number of young fledgedper territorial breeding and successful pair as re -sponse variables (Zuur et al 2009) In the aboveanalyses hatching success was defined as the pro-portion of eggs that hatched brood reduction was theproportion of hatched chicks lost before fledging aterritorial pair was one that held a territory a breed-ing pair one that laid at least 1 egg and a successfulpair one that raised at least 1 nestling to fledging age(gt45 d) In particular the likelihood of laying eggsand raising chicks to fledging were examinedthrough GLMs with binomial errors and a logit link

192

Fig 1 Approximate distribution range of the red kite in southern Spain (An-daluciacutea Autonomous Region) (a) in the 1980s (reconstructed on the basis ofsources cited in Section 23) compared to (b) the distribution as recorded in2017 The red sectors in (a) represent (1) the marshes of Dontildeana and Odiel inthe provinces of Huelva and Sevilla (2) the hills and low mountains of SierraMorena in the provinces of Huelva Sevilla and Cordoba and (3) the mountain

complex of Cazorla in the provinces of Granada and Jaeacuten


function while clutch size and the number of fledgedyoung were modelled through GLMs with Poissonerrors and a log link function Finally hatching suc-cess and brood reduction were modelled by GLMswith Poisson errors and a log link function using thenumber of hatched eggs and the number of disap-peared chicks as response variables and clutch sizeand brood size at hatching as offset variables (Zuur etal 2009)

The current expected growth of the Dontildeana popu-lation was estimated by building an age-structureddemographic model (Caswell 2001) M based on theage-dependent values of survival (see below) and themean fertility recorded over the last 5 yr in Dontildeana(0294 young per territorial pair) The matrix Mdescribes the transition of the population between 2consecutive years t and t+1 The asymptotic popula-tion growth λ is calculated as the dominant eigen-value of M (Caswell 2001) In a stable population λ isequal to 1 while lower values indicate a decliningtrend Because survival estimates were not availablefor the Dontildeana population we applied values pub-lished for other European populations (reviewed inTable 1) In all studies survival has been reported for3 age classes 1 yr 2 yr and adult (ge3 yr) so we main-tained the same age classes in the current assess-ment In particular to explore the whole range of sur-vival scenarios potentially experienced by ourpopulation we built 3 separate deterministic matrixmodels based on the minimum average and maxi-mum survival rates reported for other European pop-ulations respectively (Table 1) These exploratoryanalyses were conducted with deterministic modelsbecause uncertainty on parameter estimates de -pended largely on the population considered Wefurther assumed a 11 sex ratio and that all kites hadentered the breeding population by 3 yr of age

Based on the obtained λ we then projected the pop-ulation over the next 10minus20 yr assuming no densitydependence to explore its potential short-term via-bility or extinction over the coming decades Becauseof the assumptions outlined above and the parame-ters taken from other populations these projectionsshould be taken with caution and recalibrated asmore local data become available

Finally we used perturbation analyses to calcu-late the sensitivity S(θ) and elasticity E(θ) of thepopulation growth rate to a given parameter θ(Caswell 2001) Sensitivity is the change in pop -ulation growth rate in response to an absolutechange of a given parameter θ while elasticity isthe change in the population growth rate in re -sponse to a proportional change of a given param-eter θ Throughout all analyses were implementedwith R version 351 (R Development Core Team2018) using the libraries lsquopopbiorsquo and lsquolme4rsquoResults are given as means plusmn 1SE tests are 2-tailed and statistical significance was set at α le005

3 RESULTS

31 Regional distribution and the strategicimportance of the protected area

Red kites must originally have been common be -low 400minus600 m of elevation throughout Andaluciacuteawhich traditionally hosted a dense and well distrib-uted rabbit population as an important prey base(Loacutepez 1861) By the 1970sminus1980s legal and thenillegal shooting and poisoning had reduced the pop-ulation to 3 main nuclei (Fig 1a) (1) the marshes ofDontildeana and Odiel in the provinces of Huelva and

Sevilla (2) the hills and low moun-tains of Sierra Morena in theprovinces of Huelva Sevilla and Cor-doba and (3) the mountain complexof Cazorla in the provinces ofGranada and Jaeacuten The latter 2 nucleiwere reported as lsquoin steep declinersquo ordocumented as already extinct in the1990s (Vintildeuela et al 1999 Vintildeuela2004) so that the current regionalpopulation is virtually confined toDontildeana with 3 additional relict pairslocated in the Sierra de Huelva(Fig 1b) The nuclei outside the pro-tected area of Dontildeana were estimatedto ıhave suffered a 95 decline since

193

Area Age class Reference1 yr 2 yr ge 3 yr

England Midlands 080 094 093 Evans et al (1999)Scotland 037 072 087 Smart et al (2010)Wales uplands 060 079 094 Newton et al (1989)Switzerland 045 074 084 Schaub (2012)Spain Mallorca 072 082 079 Tavecchia et al (2012)Spain Menorca 054 083 084 Sanz Aguilar et al (2015)

Min European estimates 037 072 079Mean European estimates 058 081 087Max European estimates 080 094 094

Table 1 Published estimates of red kite survival by age class Minimum meanand maximum levels were used to set up different demographic scenarios for

matrix population models


the 1970sminus1980s (from a conservative estimate of40minus77 pairs to 2minus4 current pairs) Thus Dontildeanahosted 52minus63 of the regional population in the1970sminus80s and 95minus96 after 2010 increasing its rel-ative strategic importance

32 Abundance and trends in Dontildeana

Different sectors of the park showed differenttrends (Fig 2 Fig S2) Numbers appeared (1) stableor in moderate decline in the Reserva Biologica deDontildeana RBD (sectors 1 2 and 3) (2) stable or increas-ing in the sector of Puntal and in the vera of Algaida-Manecorro as well as its interior These areas werecharacterized by the removal of Eucalyptus planta-tions that converted large areas of woodland to opengrassland in the 1990s (sectors 4 and 5) or alwaysmaintained a very open landscape through extensivegrazing by livestock (sector 6) Finally (3) declines ofvarying magnitude were observed in all peripheralsectors to the north east and south While declineswere moderate in the eastern and southern sectorsthat were monitored less continuously dramatic de -clines were evident for all the northern sectors in -cluding portions of the National Park and its buffer-ing Natural Park In these sectors (7 8 9) a populationof 20minus25 pairs was basically eradicated in a few yearsbetween the late 1980s and early 1990s re mainingvirtually extinct to date (Fig 3) This rapid eradica-tion was accompanied by the detection of adultsfound dead by poisoning in or under their nest Over-all there was a significant preponderance of declinesbetween the numbers re corded in the earliest survey

available for each sector and the 2010s (binomial testp lt 0001 Fig 2) Correspondingly the surface of theoverall and core distribution range of the Dontildeanapopulation declined by 53 and 58 respectively(Fig 4)

When summing up all sectors the overall Dontildeanapopulation was conservatively (see Section 23) esti-mated at 69 pairs (max peak of 82) in the 1980s 61(max 86) in the 1990s 53 (max 64) in 2000minus2009and 47 (max 70) in 2010minus2017 (Table 2) In 2017 thepopulation was composed of 37 pairs representing a46minus55 decline compared to the meanminusmaximumlevels of the 1980s ie in less than 40 yr Thus the~50 declining rate within the park was high butwas only almost half of the ~95 decline estimatedfor outside pairs over the same 4 decades (χ2 = 184p lt 00001)

33 Distribution in Dontildeana

Local eradication and differential declines pro-duced a contraction of the overall range and a shiftin the relative importance of some sectors Thusin the 1980s the population was well distributedthrough all sectors but more concentrated in thecore sectors of RBD vera Puntal and in the northernpinewoods of the Natural Park buffer (Fig 5aTable 2) From the 1990s onwards the populationbecame progressively more concentrated within theNational Park in particular in the ecotonal zonebetween the marsh and terrestrial habitats withvery low numbers or local extinctions in all theother sectors (Fig 5b Table 2)

34 Offspring production in Dontildeana

Except for the number of young persuccessful pair all breeding parame-ters deteriorated through time (all χ2 ge1733 all p lt 00017) The probabilityof laying eggs clutch size hatchingsuccess the likelihood of successfullyraising chicks to fledging and thenumber of young per territorialbreeding and successful pair declinedthrough the last 5 decades while theextent of brood reduction progres-sively increased (Fig 6) Deteriorationof reproduction was particularly pro-nounced for the probability of layingfor brood reduction for the probability

194

80 ndash100 ndash80 ndash60 ndash40 ndash20 0 20 40 60 100

10 Marismillas11 Sanlucar pinewood

12 AbalarioCumulative total

1 RBD vera2 RBD interior

3 RBD sand dunes

Change in breeding numbers compared to the 1980s ()

Sector

4 Vera Algaida-Manecorro5 Interior6 Puntal

7 Coto del Rey8 NW pinewoods9 NE pinewoods

Fig 2 Percentage change in breeding numbers of the Dontildeana red kite popu-lation by sector from the 1980s or since the earliest available record (1990s for

Puntal and Abalario) to 2017 RBD Reserva Biologica de Dontildeana


of successful breeding and for the number of fledgedyoung (Fig 7)

On average the whole population was estimated toproduce 103 fledglings per year in the 1980s this al-most halved to 57 in the 1990s 48 in 2000minus2009 andagain declined to more than halved to 19 in 2010minus2017In 2015 2016 and 2017 the population produced 7 7and 15 fledglings respectively Population decline and

breeding deterioration were accompa-nied by a change in the relative impor-tance of some sectors for offspring pro-duction (Fig 5 Table 2) In the 1980smost young were produced in thesector RBD vera and the northernpinewoods and most other sectors stillcontributed gt5 of the annual youngproduction (Fig 5c) In subsequentdecades fledgling production becameconcentrated in the marsh ecotone(Puntal RBD vera vera of Algaida-Manecorro) with additional inputsfrom some interior or peripheral sectorsand a larger portion of sectors with vir-tually zero contribution to future gener-ations (Fig 5d) Fig 3 illustrates howdiffuse successful reproduction was inthe 1980s compared to 2017

35 Comparison with otherEuropean populations

Compared to other areas of Europethe Dontildeana population showed a highdensity in the 1980s (well above theEuropean mean Table S3) This pro-gressively declined and has changedto below-average density since 2010All breeding parameters were wellbelow the mean European estimatesfrom the 1980s onwards (Table S3) Inparticular the percentage of success-ful pairs and the number of youngfledged per breeding pair are cur-rently 2 to 3 times lower than themean European values

36 Population growth andprojections for the Dontildeana

population

Observed declines were well cap-tured by the growth rate predicted by the populationmodel based on the minimum European survivalrates (Fig 8) The other 2 scenarios of maximum andmean survival rates predicted a stable population ordeclines from the year 2000 onwards only (Fig 8)For the 2010s the model estimated λ = 0849 0960and 1066 for the scenario based on minimum meanand maximum survival estimates respectively

195

Fig 3 Distribution of breeding territories in Dontildeana (a) in the 1980s and (b) in2017 Pairs that bred successfully (black points) were much more frequent andmore widely distributed especially in the northern sectors and in the NaturalPark in the 1980s than in 2017 when most pairs failed to raise chicks to fledg-ing (white points) Territory locations have been moved slightly for conserva-tion reasons and locations from the 1980s could only be approximate for some

territories especially those in the northern pinewoods


Based on the model with minimum survival rateswhich gave the best fit to past declines the popula-tion of 37 pairs censused in 2017 would be reduced to7 pairs within 10 yr and virtually extinct (le2 pairs) in17 yr The less supported scenario of mean Europeansurvival estimates projected a population of 25 pairsin 10 yr while the most optimistic scenario of maxi-mum survival predicted a full recovery to the 1980spopulation level of 70 pairs in 10 yr

Despite the current extremely lowbreeding success under all 3 scenar-ios of survival the population growthrate was least sensitive to changes infledglings production [E(F) = 0048minus0079 S(F) = 0137minus 0277] and mostsensitive to variations in adult survivalprobability [E(θ) = 0843minus 0904 S(θ) =0929minus 0955] In Fig 9 we plot thecombination of fledgling productionand adult survival that produce differ-ent levels of population growth Atcurrent levels of reproduction adultsurvival would have to be around090minus095 to attain a stable or slightlyincreasing population while restoringbreeding success to the levels of the1980s of 13 young per pair wouldimply a stable population even with arelatively low adult survival of ~077ie below the minimum ever recordedfor the species (Fig 9)

4 DISCUSSION

Our results showed that the currentdeteriorated condition of the Dontildeanared kite population was not a recentphenomenon but the tip of the ice-berg of decades of regional and localdegradation Although the protectedarea slowed the background levels ofregional decline and likely saved thespecies from local extinction its via-bility maintenance performance wasfar from satisfactory In a way the ob -served long-term deterioration actedas a protracted diluted form of extinc-tion debt (Tilman et al 1994) paidover a time-frame of several decadesWe suspect that similar conditionsmay be common in other protectedareas making this a classical example

of gradual underlying pressures diluted over such along time-frame that their severity becomes evidentonly through a large-scale long-term asses sment oronce population degradation has reached its extremeconsequences Note that for many parameters dete-rioration was not especially alarming from onedecade to the next making it difficult for the succes-sion of local field observers to appreciate the overallseverity of the long-term trends This difficulty in

196

Fig 4 Distribution range of red kites in Dontildeana (a) in the 1980s and (b) in2017 The pink outer polygon represents a 95 kernel constructed aroundthe territory locations while the red polygon is a 50 kernel showing the corearea of distribution The overall and core distribution range contracted by 534and 581 respectively Territory locations have been moved slightly for

conservation reasons

Sergio et al Predator extinction debt in national park 197

12

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1999

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276

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114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

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046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a


detecting the problem was likely exacerbated by thefollowing factors that could be defined as lsquoperceptionobstaclesrsquo (1) The fact that a population becomesmore and more restricted to a park is often used toillustrate the positive role of protected areas byfocusing their performance assessments on range

coverage ie the capability of protected areas tocover a large portion of a species range (eg Czech2005 Godet et al 2007) This diverts attention fromthe wider problems of range contraction and from theactual performance of the park in terms of viabilitymaintenance (Gaston et al 2008) (2) Confinement to

198

of nests in each sector

of annual young generated by each sector

1980s 2010s

1980s 2010s

a) b)

c) d)

Fig 5 Percentage of the total red kite population in each of 12 sectors of Dontildeana (a) in the 1980s and (b) in the 2010s percent-age of the annual young production generated by each sector (c) in the 1980s and (d) in the 2010s (a) In the 1980s the bulk ofthe population was concentrated in northern sectors (7 and 8) and in the southern shores of the marshes (sectors 1 6 and 10)(b) In the 2010s the population virtually disappeared from the northern sectors and became essentially confined to the na-tional park where it concentrated along the marshes (c) In the 1980s the population generated more than 100 young eachyear most of them produced by the northern sectors (7 8 9) and by the northern shores of the marshes (sectors 1 and 4) (d) Inthe 2010s the population generated less than 10 young mostly generated close to wetlands (sectors 1 4 11) with homo -geneously low numbers elsewhere Colors (yellow rarr orange rarr red) classify sectors by tertiles of the range of percentage

contribution


a protected area is usually accompanied by the auto-matic presumption that this will act as a source areawith limited anthropogenic impacts an assumptionthat is increasingly disproved (Hansen amp DeFries2007 Gaston et al 2008 Watson et al 2014) (3) Long-term assessments such as ours required the lengthy

collation of fragmented informationfrom a large number of sequentialfield workers and administrationseach of which had a very limited per-ception of the overall situation (4)Much monitoring was conducted inthe core areas of the park like theRBD where the species has alwaysbeen most abundant and relativelystable (eg Sergio et al 2005) con-fusing the perception of broaderdeclines (5) Finally large numbersof wintering migrants boost the pop-ulation in the winter and earlyspring often producing an optimisticimpression of apparent abundance

even in relatively expert members of the public or oflocal administrations as we frequently noted in pub-lic talks and private conversations

Overall the decline occurred at multiple levels redkites were originally widespread throughout south-ern Spain (Loacutepez 1861) but became progressively

199

2030405060708090

100110

Bree

ding

su

cces

sful

()

a breeding pairs successful pairs

202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade

c Hatching successBrood reduction

03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade

d Young breeding pairYoung successful pair

Fig 6 Reproductive parameters of the red kite population of Dontildeana (a) percentage of pairs laying eggs (black dots n = 804)and successfully raising fledglings (white dots n = 1185) (b) mean clutch size (n = 469) (c) hatching success (percentage ofeggs that hatched black dots n = 353) and brood reduction (percentage of chicks that disappeared during the nestling periodwhite dots n = 240) (d) mean number of young fledged per territorial pair (black dots n = 1145) and per successful pair (white

dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75

Young fledgedsuccessful pairYoung fledgedbreeding pairYoung fledgedterritorial pair

Probability of successful breedingBrood reduction

Hatching successClutch size

Probability of laying

Deterioration ()

Breeding parameter

Fig 7 Percentage deterioration of breeding parameters of the red kite popu-lation of Dontildeana between the 1980s and the 2010s


confined to the protected area of Dontildeana in recentdecades However the park did not confer automaticsafety and declines and range contraction also con-tinued within the protected area suggesting perva-sive pressures and range erosion at multiple spatialscales In particular almost one third of the Dontildeana

population was virtually eradicatedfrom the northern pinewoods of thebuffering Natural Park in 5 to 7 yrbetween the late 1980s and early1990s Several lines of evidence sug-gest that illegal poisoning was themain cause of this eradication (1) Sev-eral adults were found poisoned undertheir nest in those sectors and years(2) Large-scale poisoning campaignsof raptors were reported throughoutSpain in those exact same years asa consequence of huntersrsquo illegalattempts to control predator popula-tions to increase the rapidly deterio-rating numbers of small game (rabbitsand partridges Villafuerte et al 1998Vintildeuela amp Villafuerte 2003 Cano et al2008) In this sense the National Parksuffered much wider national-level

pressures (3) The northern sectors of the parkwhere the eradication took place included or wereclose to private small-game reserves and predatorcontrol was well known to occur in some of them (4)Finally the rapidity of the eradication of such a long-lived species (maximum longevity of 29 yr in Dontilde -ana) implied a causative agent that directly affectedadult mortality (Saether amp Bakke 2000) such as poi-soning The marked impact of poisoning on adult sur-vival and population growth are well recognized forthis species whose facultative scavenging habits andforaging method of meticulous low and slow quar-tering of the ground make it particularly sensitive tothis threat (Smart et al 2010 Tavecchia et al 2012Tenan et al 2012 Sanz-Aguilar et al 2015)

Spikes of rapid local eradication were superim-posed on more gradual but steady deterioration ofoffspring production Gradual and continuous ratherthan sudden degradation of performance suggestedthat poisoning was not the only threat in effect Inthis context several factors may have contributed togradually deteriorate ecological conditions for kitesin recent decades (1) Rabbit epidemics have causedprogressive declines in the availability of this keyprey (Vintildeuela amp Villafuerte 2003) (2) Lower preyavailability may have exacerbated competition withthe dense populations of other raptors with similardiets such as the ecologically similar and in creasingblack kite (3) Predation rates on adults and nestlingsmay have increased with the expansion of the popu-lations of potential predators such as eagle owls Bubobubo and goshawks Accipiter gentilis (4) Recentassessments have detected alarming levels of toxic

200

00

02

04

06

08

10

12

1980sndash1990s 1990sndash2000s 2000sndash2010s

Popu

latio

n gr

owth

rate

Period

Observed populationgrowth

Scenario 1 minimumEuropean survival

Scenario 2 meanEuropean survival

Scenario 3 maximumEuropean survival

Fig 8 Population growth of the Dontildeana red kite population during the past 4decades as predicted by matrix demographic models based on 3 scenarioswhich assumed that survival was equal to minimum mean or maximum sur-vival rates reported for other European populations (reviewed in Table 1)Predicted values are compared with observed declines from one decade to

the next

Fig 9 Population growth rate of red kites in relation to adultsurvival and breeding performance (mean number of youngfledged per territorial pair) based on a matrix populationmodel Note that the graph could be applied to any Euro-pean population The 3 points at the base of the graph repre-sent the minimum (black) mean (grey) and maximum (blue)survival published for European populations (reviewed in

Table 1)


chemicals in red kite eggs from Dontildeana (Goacutemara etal 2008) (5) Climate change and agricultural man-agement have lowered the water table with likelyimpacts on marsh inundation levels and aquatic prey(Garciacutea Novo amp Mariacuten Cabrera 2005 Diacuteaz-Paniaguaamp Aragoneacutes 2015 Schmidt et al 2017) (6) Conver-sion to intensive farmland in surrounding lands haseroded habitat and prey availability outside the pro-tected area which may have buffered kites fromperiods of low prey availability within the park (egduring droughts in the marshes) Some of the pat-terns of observed reproductive deterioration lendsupport to some of these impacts For exampledeclining clutch sizes and increasing rates of broodreduction suggest that food was not sufficient to sus-tain earlier reproductive levels while lowered hatch-ing success is often the result of chemical contamina-tion in raptors (review in Newton 1979) While eachof these threats will be quantitatively investigated indetail in the immediate future in the present contextthey illustrate the frequent and complex challengesfaced by (1) protected areas in maintaining biodiver-sity and viable populations of imperiled species and(2) conservation biologists in identifying causes ofdecline for subsequent management interventioneven in lsquoprivilegedrsquo sites such as reserves

Redistribution declining numbers and perform-ance had important repercussions for future viability(1) The population passed from producing gt100annual young to lt10minus20 which will make mainte-nance and recovery difficult (2) These young wereincreasingly produced in the core sectors of themarsh-ecotone within the National Park which prob-ably increased the dependence of the population onmarsh inundation in turn affected by problems oper-ating outside the park (Garciacutea Novo amp Mariacuten Cabrera2005 Diacuteaz-Paniagua amp Aragoneacutes 2015 Schmidt et al2017) (3) Steep declines in the percentage of suc-cessful pairs implied that future generations wereproduced by a progressively more restricted numberof pairs which will make the population more sub-ject to inbreeding and stochastic events For exam-ple in the last 3 yr all the annual young were pro-duced by 6 8 and 9 pairs respectively suggesting aneffective population size 4 to 9 times lower than theoverall abundance

Given all the above it is no surprise that demo-graphic models suggested a negative growth and apopulation incapable of self-sustenance In particu-lar the model that gave the best fit to past observeddeclines was the one based on minimum Europeansurvival estimates followed by the scenario based onmean European survival estimates Given the above-

mentioned threats declines and performance deteri-oration we would expect survival to be closer to min-imum than mean European levels as supported bythe observed fit to past and recent declines Theseresults have 2 main implications (1) If survival is aslow as we suspect demographic models predict vir-tual extinction within a few years demanding anurgent need for identification and management ofmortality threats as well as quantification of actualsurvival to refine predictions Note that virtual ex -tinction could be even earlier when taking intoaccount effective rather than total population sizeand a stochastic rather than deterministic populationmodel (2) Should actual survival be confirmed lowbut further declines not observed the populationwould be a sink sustained by external sources whichwould make it even more at risk from anthropogenicthreats outside the park and thus more challengingto manage (eg Hansen amp DeFries 2007 Blanco et al2017)

5 Conclusions and implications for conservation

The decades-long degradation that we depictedwas likely generated by a combination of factorsboth internal to the park (eg rabbit availabilitymarsh inundation rates predation and competitionwith other species) and external (eg illegal poi-soning contamination farmland intensification)Some of them will be amenable to managementand others less so but the first priority will be tofind rapid ways to identify the key factors thatdrive population growth so as to optimize conserva-tion action before it is too late For example focus-ing conservation lsquoblindlyrsquo on supplementary feedingcould be useless if the problem is chemical contam-ination The quickest way to obtain such informa-tion would be a satelliteGSM (Global System forMobile communication) tagging program that wouldallow to (1) obtain rapid estimates of health andsurvival for young and adults (eg Sergio et al2019) so as to recalibrate future projections andgoals (2) assess the relative contribution of mainmortality causes which is an essential first step inany recovery program (eg Gonzaacutelez et al 2007Margalida et al 2008 Smart et al 2010) and (3)identify the areas used by breeding and non-breed-ing individuals inside and outside of the park toassess their risk-exposure and intervene before orafter death (eg to prosecute perpetrators of illegalpoisoning or re trofit dangerous electricity pylons inareas of high use by kites)

201


Until this information becomes available below welist some tentative figures and priorities for conserva-tion First of all restoring the current population toformer levels would undoubtedly be challenging andprobably unrealistic given the profound changesincurred by the wider Dontildeana ecosystem in recentdecades Thus we propose an initial target of 60pairs as a long-term restoration goal Elasticity analy-sis suggested that optimal management should focuson improving adult survival which had the largestimpact on population growth as in other long-livedspecies (Saether amp Bakke 2000 Whitfield et al 2004Kruumlger et al 2010) This could be attempted by in -creasing food availability especially of key prey suchas rabbits and by curtailing mortality threats such asillegal poisoning which still continues despite mucheffort to eradicate it by the local administration Thetelemetry program outlined above could help toidentify local perpetrators as occurred in other pop-ulations (Smart et al 2010 Tavecchia et al 2012) Onthe other hand despite their low elasticities repro-ductive rates are currently so low that it would befundamental to increase them eg by increasingfood availability and ensuring proper inundation ofthe marshes Based on Fig 5 we propose as initialtargets for restoration (1) survival rates equal orhigher than the European mean (adult survivalge087) and (2) a number of young fledged per territo-rial pair above 10 which would produce an increas-ing population provided that adult survival is equalto or above the European mean Note that Fig 5could be applied to any European population as atool to check its potential viability Given the highlythreatened status of red kites the speciesrsquo limitedrange confined to Europe and its ongoing declinesviability assessments such as ours may sadly becomemore important for the management and recovery ofmany populations

In conclusion red kites would probably be extinctin southern Spain without the institution of a largenational park In this sense this example well illus-trates how protected areas will continue to be corner-stones of global conservation of threatened taxaHowever their safety is not ensured by park institu-tion per se and performance monitoring and focusedaction will be needed to avoid potential extinctiondebts which may be more protracted than previouslyappreciated In our exemplary case despite the pres-ence of a large national park on prime fertile landwithin a developed country with adequate institu-tional and financial support a key threatened speciessuffered latent but steady declines for decadesobscured by problems of perception lack and frag-

mentation of information and likely caused by acomplex array of within-park threats and growingexternal pressures Viability maintenance and recov-ery will probably require concerted actions withinand outside of the park but only after detailed infor-mation on crucial aspects becomes available whichis probably the most urgent current priority We sus-pect that dynamics like the ones illustrated here aremore common than currently appreciated and willbecome more widespread as anthropogenic pres-sures increase around protected areas and as theirperformance monitoring becomes more prevalent

Acknowledgements We thank all the (too many to mention)researchers field workers visitors volunteers gamekeep-ers park managers and wardens that have provided datasince the 1960s In particular we thank F G Vilches L Gar-ciacutea and the Equipo de Seguimiento de Procesos Naturales ofthe Estacioacuten Bioloacutegica de Dontildeana-CSIC for help in the fieldSpecial thanks to F G Vilches for the priceless help with thecollation and organization of historical information Wethank J P Castellano I Redondo and all the lsquoArea de Con-servacioacutenrsquo of Dontildeana National Park for constant interest andsupport We thank R Cadenas J Madrid R Arenas J RBenitez and all the pertinent personnel of the Consejeriacutea deMedio Ambiente of the Junta de Andaluciacutea for informationon current nest locations in the Sierra de Huelva and fortheir constant attentive support We are grateful to Dr NBunnefeld and 2 anonymous reviewers for constructivecomments that improved the manuscript This study wasfunded by the Foundation Jaime Gonzaacutelez-Gordon and bythe research project 16022015 of the Spanish Ministry ofAgriculture Food and the Environment (National ParksAutonomous Agency)

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Goacutemara B Gonzaacutelez MJ Baos R Hiraldo F Abad E RiveraJ Jimeacutenez B (2008) Unexpected high PCB and total DDTlevels in the breeding population of red kite (Milvus mil-vus) from Dontildeana National Park south-western SpainEnviron Int 34 73minus78

Gonzaacutelez LM Margalida A Mantildeosa S Saacutenchez R and oth-ers (2007) Causes and spatio-temporal variations of non-natural mortality in the vulnerable Spanish imperialeagle Aquila adalberti during a recovery period Oryx41 495minus502

Hansen AJ DeFries R (2007) Ecological mechanisms linkingprotected areas to surrounding lands Ecol Appl 17 974minus988

Hernaacutendez FJ Donaacutezar JA Hiraldo F (2001) Evaluacioacuten deluso de venenos para el control de predadores en laComunidad Autoacutenoma Andaluza y su impacto en la con-servacioacuten de especies amenazadas de vertebrados Con-sejeria de Medio Ambiente and Estacioacuten Bioloacutegica deDontildeana Sevilla

IUCN (The World Conservation Union) (1994) Guidelines forprotected area management categories IUCN Gland

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Loacutepez V (1861) Cataacutelogo de las aves observadas enAndaluciacutea Madrid

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Margalida A Heredia R Razin M Hernaacutendez M (2008)Sources of variation in mortality of the bearded vultureGypaetus barbatus in Europe Bird Conserv Int 18 1minus10

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Newton I (1979) Population ecology of raptors T amp AD PoyserBerkhamsted

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Saeligther BE Bakke Oslash (2000) Avian life history variation andcontribution of demographic traits to the populationgrowth rate Ecology 81 642minus653

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204

Editorial responsibility Nils BunnefeldStirling UK

Submitted October 8 2018 Accepted January 28 2019Proofs received from author(s) April 7 2019


toring whether protected areas actually deliver bio-diversity benefits ie their performance (Gaston etal 2008) For example some studies have reporteddeclines and extinctions of key species in protectedareas and sometimes steeper declines within pro-tected areas than outside them (eg Brashares et al2001 Parks amp Harcourt 2002 Whitfield et al 2007Craigie et al 2010) This calls for the need to assessthe historical levels trends and viability of threat-ened species within protected areas in order toensure that they are properly preserved that theycan act as sources of colonizers for outside areas orthat potential declines are detected and haltedthrough management

Here we report a representative example of thebenefits and challenges provided and faced by pro-tected areas in attaining their biodiversity mainte-nance function In particular we focus on the long-term population dynamics of a highly endangeredavian predator the red kite Milvus milvus breedingin Dontildeana National Park in southern Spain

Dontildeana is one of the most biodiversity-rich parks inEurope currently listed as a Ramsar Site a WorldHeritage Site and a UNESCOrsquos Man and the Bio-sphere Reserve While it is usually lands of low eco-nomic value that are designated as protected areas(eg Scott et al 2001) Dontildeana is a relatively largepark (gt1000 km2) centered around a seasonal wet-land in prime fertile land along a river estuary Whileproviding clear biodiversity benefits its location onvery productive land has caused an exponentiallygrowing anthropogenic pressure at the park bordersespecially by rapid habitat conversion to intensivefarmland with consequent impacts on the watertable that sustains the reserve wetlands The park isalso famous worldwide for its dense populations ofcharismatic vertebrate predators which represent amajor attraction for over 300 000 annual tourists anda key focus for conservation management (eg Ferreramp Hiraldo 1991 Loacutepez et al 2009)

One component of such a predator assemblage isthe red kite a medium-sized raptor endemic to thewestern Palearctic with 95 of its world populationconfined to Europe (Knott et al 2009) This speciestypical of open landscapes with scattered woodlandhas suffered dramatic declines in recent decades inall its stronghold populations of Germany Franceand Spain which hold about 80 of the world popu-lation (Knott et al 2009) Due to such declines thered kite is currently classed as Near Threatened inthe IUCN Red List and included in Annex I of the EUBirds Directive and in Annex II of the Bern Conven-tion on the Conservation of European Wildlife and

Natural Habitats In Spain which holds the worldrsquosthird largest population the species has been clas -sified as lsquoat risk of extinctionrsquo since 2011 and as lsquoatcritical risk of extinctionrsquo within southern Spain(Andaluciacutea Autonomous Region) where the presentstudy was conducted In Dontildeana the breeding popu-lation is concentrated near the marshes (Sergio et al2005) where its generalist and opportunist diet isdominated by prey species typical of the marshes andof the ecotone between wetland and terrestrial habi-tats (eg rabbits Oryctolagus cuniculus waterbirdscarrion and reptiles Delibes amp Garciacutea 1984 Veiga ampHiraldo 1990) The preservation of this population isparticularly important for 3 main reasons (1) It is thelast reservoir of the species in southern Spain (seeSection 31) and thus a strategic key for avoiding fur-ther range contraction (a major goal of the EuropeanAction Plan for this species Knott et al 2009) (2) Itshigh dependence on a seasonal wetland makes this aunique population (Vintildeuela et al 1999 Seoane et al2003) with special adaptations to an unpredictablenatural disturbance regime of marsh inundationwhich would be lost in case of extinction with theconsequent erosion of functional biodiversity (3)During winter large numbers of red kite migrantsfrom central-northern Europe join the resident breed-ing population which can thus act as a sentinel formortality threats potentially impacting far away pop-ulations through carryover effects

Given all the above we reconstruct the historicalregional distribution of the red kite in southernSpain document its progressive extirpation and con-finement to Dontildeana demonstrate long-term declinesin abundance and performance of the park popula-tion across several decades provide projections offuture extinction risk and identify potential threats inurgent need of investigation and management

2 METHODS

21 Study area

The protected area of Dontildeana (hereafter Dontildeana)is located in southwestern Spain (36deg 56rsquo 51rsquorsquo N 6deg 21rsquo31rsquorsquo E) within the Autonomous Region of AndaluciacuteaIt is composed of 542 km2 of National Park institutedin 1969 and 538 km2 of buffering Natural Park witha less strict protection category instituted in 1989(Fig S1 and Table S1 in the Supplement at www int-res com articles suppl n038 p189 _ supp pdf) The Nat -ional Park is dominated by 5 main habitat types(1) seasonal marshland usually flooded during win-

190



22 Field procedures





191








192








3 RESULTS




193
















194





3 RBD sand dunes


Sector












population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204





22 Field procedures





191








192








3 RESULTS




193
















194





3 RBD sand dunes


Sector












population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204










192








3 RESULTS




193
















194





3 RBD sand dunes


Sector












population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204








3 RESULTS




193
















194





3 RBD sand dunes


Sector












population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204













194





3 RBD sand dunes


Sector












population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204










population


195






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204






4 DISCUSSION



196




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204




12

3 R

BD

4 V

era

56

7 C

oto

8 N

W9

NE

10

11

12

Tot

al

RB

DR

BD

san

dA

lgai

da

Inte

rior

Pu

nta

ld

elp

ine-

pin

e-M

aris

-S

anlu

car

Ab

alar

iod

ecli

ne

vera

inte

rior

du

nes

Man

ecor

roR

eyw

ood

sw

ood

sm

illa

sp

inew

ood

a) M

ean

no

of

pai

rs19

80minus

1989

141

52

33

40a

20a

80b

50

107

44

50

30

40b

687

1990

minus19

9915

35

11

37

74

08

03

34

40

74

5e2

5c4

0a60

8minus

115

2000

minus20

0916

05

21

47

05

06

31

01

00

14

5e2

03

1a52

5minus

236

2010

minus20

1713

03

42

56

63

98

41

90

50

13

01

72

147

1minus

315

2017

100

35

211

00

03

12

37minus

462

b)

No

of

you

ng

pro

du

ced

an

nu

ally

1980

minus19

8916

14

25

66

43

09

610

020

311

06

04

56

010

26

1990

minus19

999

82

60

68

44

06

92

46

60

95

13

86

056

9minus

445

2000

minus20

0913

21

61

27

25

83

71

21

60

24

53

04

647

7minus

535

2010

minus20

172

60

61

11

82

33

50

90

30

01

62

91

919

4minus

811

2017

10

03

07

00

00

22

150

minus85

4

c)

of

the

tota

l p

op

ula

tio

n i

n e

ach

sec

tor

1980

minus19

8920

57

64

85

82

911

67

315

56

47

34

45

819

90minus

1999

251

84

21

126

66

131

55

73

12

74

41

66

2000

minus20

0929

49

62

612

99

211

51

81

80

28

33

79

120

10minus

2017

276

72

53

141

82

178

40

11

03

64

36

45

2017

270

00

81

135

54

297

00

00

00

81

27

54

d)

c

on

trib

uti

on

of

each

sec

tor

to t

ota

l an

nu

al y

ou

ng

pro

du

ctio

n19

80minus

1989

157

41

54

94

62

29

97

198

107

58

44

58

1990

minus19

9917

24

61

112

114

77

04

211

61

68

96

610

520

00minus

2009

277

33

24

78

151

121

24

34

04

94

63

96

2010

minus20

1713

43

35

418

19

411

74

81

30

08

215

09

620

176

70

00

046

720

00

00

00

00

00

013

313

3

a Cro

ss-c

omb

ined

est

imat

e b

Sta

tic

bac

k-e

stim

ate

c In

terp

olat

ed e

stim

ate

See

Sec

tion

23

for

det

ails

Tab

le 2

(a

) M

ean

nu

mb

er o

f p

airs

an

d (

b)

cum

ula

tive

nu

mb

er o

f of

fsp

rin

g p

rod

uce

d a

nn

ual

ly b

y th

e D

ontildean

a re

d k

ite

pop

ula

tion

wit

hin

eac

h s

ecto

r an

d i

n t

he

wh

ole

par

k o

ver

the

pas

t d

ecad

es a

nd

in 2

017

Per

cen

tag

e of

(c)

pai

rs a

nd

(d

) yo

un

g c

ontr

ibu

ted

by

each

sec

tor

to t

he

over

all p

opu

lati

on S

ecto

rs c

ontr

ibu

tin

g m

ore

than

10

ar

e h

igh

lig

hte

d i

n b

old

T

otal

ref

ers

to t

he

sum

of

all

sect

ors

d

ecli

ne

is t

he

dec

lin

e co

mp

ared

to

the

1980

s P

rop

orti

onal

con

trib

uti

ons

are

also

ill

ust

rate

d i

n F

ig

5

RB

D R

eser

va B

iolo

gic

a d

e D

ontildean

a




198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204






198



1980s 2010s

1980s 2010s

a) b)

c) d)


contribution






199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204








199

2030405060708090

100110

Bree

ding

su

cces

sful

()


202122232425262728

Clut

ch si

ze

b

25

35

45

55

65

75

85

1970-79 1980-89 1990-99 2000-09 2010-17

Hatc

hing

succ

ess

broo

d re

duct

ion

()

Decade


03

06

09

12

15

18

21

1970-79 1980-89 1990-99 2000-09 2010-17

Youn

g fle

dged

Decade



dots n = 576)

0 5 10 15 20 25 30 35 40 45 50 55 60 65 70 75





Deterioration ()

Breeding parameter







200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204








200

00

02

04

06

08

10

12


Popu

latio

n gr

owth

rate

Period






the next


Table 1)








201






LITERATURE CITED









202










































203











204










201






LITERATURE CITED









202










































203











204








LITERATURE CITED









202










































203











204












































203











204













204



Documents

Protected areas under pressure: decline, redistribution ...Giacomo Tavecchia2, Fernando Hiraldo1 1Department of Conservation Biology, Estación Biológica de Doñana − CSIC, C/Americo