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A Demographic Description of the Recovery of the Vulnerable Spanish Imperial Eagle Aquila Adalberti

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A Demographic Description of the Recovery of the Vulnerable Spanish Imperial Eagle Aquila Adalberti A demographic description of the recovery of the Vulnerable Spanish imperial eagle Aquila adalberti E nric O rtega,Santi M a N˜ osa,Antoni M argalida,Roberto S A´ nchez J avier O ria and L uis M ariano G onzA´ lez Abstract The population of the Vulnerable Spanish im- strategies for conservation (Sarrazin & Legendre, 2000). perial eagle Aquila adalberti has experienced a gradual Such models can also be used to anticipate future population recovery from 38 pairs (1974) to 198 (2004). We analysed trends (Wootton & Bell, 1992). Demographic studies in birds the spatial and temporal variation of the demographic para- of prey have been carried out mainly in declining popula- meters for 1981–2004. Annual productivity was 1.19–1.32 tions (Whitfield et al., 2004), and less attention has been chicks per female and adult survival rate 0.918–0.986. paid to the demographic processes underlying recovering Survival during the post-fledging period was 0.894 and the populations (Wyllie & Newton, 1991; Ferrer & Dona´zar, annual survival rate of the dispersing individuals was 0.561. 1996). However, the indicators of demographic health or Three phases of population evolution were distinguished: sensitivity to demographic parameters in growing popula- growth (1981–1993), stability or slight decrease (1994– tions can differ from those observed in declining populations 1999) and growth (2000–2004). Variation in adult survival and, therefore, the diagnostic tools and the conservation seems to explain this pattern for the first two periods. strategies may be different (Katzner et al., 2007). However, a large disparity between the observed growth The Spanish imperial eagle Aquila adalberti is catego- rate and the modelled population growth in 2000–2004 is rized as Vulnerable on the IUCN Red List (Birdlife best explained if we assume that a decrease in the age of International, 2004;IUCN,2007) and recovered from 38 recruitment took place. This is supported by the recent pairs in the 1970stoc.200 pairs in 2004 (Gonza´lez & Oria, increase in the frequency of non-adult birds in breeding 2004). This offers the possibility of analysing the demo- pairs. The survival of unpaired eagles in dispersal areas is graphic processes associated with a population recovery. becoming more important for the maintenance of current The demography of the population of 8–15 pairs in Don˜ana population growth. Spatial variation of adult survival and has already been analysed (Ferrer & Caldero´n, 1990; breeding success is not congruent with the observed growth Ferrer & Dona´zar, 1996; Ferrer & Bisson, 2003; Ferrer rate of the population, which suggests the existence of an et al., 2003) but there is no published information on the important flow of individuals between populations. These demography of the largest part of the population. In this results highlight the importance of addressing the conser- context the aims of this research were to (1) describe and vation of the species from a global perspective and the need analyse the geographical variation of the species’ popula- to focus on adult survival in breeding territories and on tion trend during 1981–2004,(2) estimate the main de- non-adult survival in dispersal areas. mographic parameters and analyse their spatial and temporal variation, (3) build a demographic model to Keywords Aquila adalberti, demography, population dy- relate these variations to the species’ population trend, namics, Spain, Spanish imperial eagle. and (4) make recommendations for the conservation of the species. Introduction Methods emographic models are useful tools to understand Dpopulation trends and the influence of demographic All known breeding territories of the species up to 2004 parameters (Lebreton & Clobert, 1991) and to evaluate (Gonza´lez & Oria, 2004; Gonza´lez et al., 2006b) were grouped into five geographical areas: North, Centre, West, ENRIC ORTEGA and SANTI MAN˜ OSA Departament de Biologia Animal, South and Don˜ana (Fig. 1), based on the species’ breeding Universitat de Barcelona, Barcelona, Spain. area in 2004 (Gonza´lez et al., 2008). Breeding territories ANTONI MARGALIDA (Corresponding author) Bearded Vulture Study & were clumped in the same area if they were , 20 km apart Protection Group, Apdo. 43, E-25520 El Pont de Suert, Lleida, Spain. E-mail [email protected] and belonged to a similar geographical and habitat unit. 10 ROBERTO SA´ NCHEZ TRAGSA, c/Vela´zquez, Madrid, Spain. The data come from national censuses of the species, in 1974 (Garzo´n, 1974), 1981–1986 (Gonza´lez et al., 1987), JAVIER ORIA Boscaje S.L., Segovia, Spain. 1989 (Gonza´lez, 1991), 1994 (Gonza´lez, 1996), and 1999, LUIS MARIANO GONZA´ LEZ Direccio´n General para la Biodiversidad, Minis- 2000 2001 2002 2003 2004 2004 terio de Medio Ambiente, Madrid, Spain. , , , and (Gonza´lez & Oria, ); Received 11 June 2007. Revision requested 31 August 2007. each census covered the species’ whole range. We focus on Accepted 11 February 2008. North, Centre, West and South, which account for 95%of ª 2009 Fauna & Flora International, Oryx, 43(1), 113–121 doi:10.1017/S0030605307991048 Printed in the United Kingdom Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.14, on 24 Sep 2021 at 09:46:09, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605307991048 114 E. Ortega et al. FIG. 3 Leslie matrix of the model for the population of the Spanish imperial eagle i, where n1, n2, n3 and n4 are the numbers of females of each age group, t is time in years, So survival during post-fledging, Sd survival of the dispersers, S3 survival of the third to fourth age group, Sa survival of the breeders, b is the proportion of fledgling females, P3 and P4 are the productivity of the females in the third and fourth age classes, respectively, and 3 and 4 are the proportion of breeding females in each of these groups. FIG. 1 The five areas of the distribution of the Spanish imperial individuals 0.7 years old; age 2, individuals 1.7 years old; age 3, eagle (see text for further details); 95% of the population occurs individuals 2.7 years old; age 4, individuals $ 3.7 years old. in North, South, West and East, and only these areas were The model included the following parameters: considered in this study. So, survival from fledging until independence (the same value was assumed for all models); S , annual survival during the dispersal period, 0.7–2.7 the population (Fig. 2). A total of 220 territories, which d years of age, using the same value in all simulations; were occupied with variable frequency, were monitored in S , annual survival of breeders $ 3.7 years old; these censuses. Except for the 1974 census, LMG, JO and RS a S3, annual survival rate over 2.7–3.7 years, could not be participated in and coordinated the censuses. Previous directly estimated because of the lack of radio tracking data work has shown the existence of three periods of popula- for this period, and was therefore calculated in all cases tion growth during the recovery period: 1981–1993, 1994– as the average of S and S , weighted according to the 1999 and 2000–2004 (Gonza´lez, 1996; Gonza´lez & Oria, a d proportion of age 3 birds that are assumed to breed (c3; 2004). Comparisons of observed with expected trends were Fig. 3; in this way we assumed that survival before settle- conducted with a Student t-test for related samples, ment on a breeding territory was low and only rose after comparing results of the real counts with the number of settlement); pairs predicted by the model. P3, average number of chicks fledged by age 3 females; A Leslie matrix model structured in four age classes was P4, average number of chicks fledged by age 4 females; constructed (Fig. 3). We modelled only the female fraction r, proportion of females among fledglings, set as 0.5 in of the population. The model took the average date of birth all cases. to be 15 April and the beginning of the post-fledging period We used the software ULM 4.0 (Legendre, 2003) to run to be 1 July (Margalida et al., 2007). The dependence period the model. One hundred random trajectories were drawn of fledglings was assumed to last until 1 October (Alonso by randomly changing the productivity and survival pa- et al., 1987). At the beginning of each reproductive cycle, rameters each year, considering the productivity values P3 1 January, four possible age classes were considered: age 1, and P4 as stochastic variables with mean and variance following a log-normal distribution, and survival rates So, Sd, and Sa as stochastic variables with mean and variance following a beta distribution. We used the total variance, which integrates the inter-annual and inter-territorial variability of productivity. In this way, we estimated the average population growth rate ke and its associated stan- dard errors under different scenarios. Firstly, we built a global model considering all the geographical aggrega- tions as a single population going through three phases of population growth, 1981–1993, 1994–1999 and 2000–2004, each with its own estimated demographic parameters. The initial number of individuals in each age group (ni) at the 1981 FIG. 2 Changes in the number of pairs of the Spanish imperial start of the simulation in was estimated by interpola- eagle in the five areas (Fig. 1) from the results of 10 national tion from the results of the 1974 and 1986 censuses, as- censuses. suming that the population was in equilibrium. The initial ª 2009 Fauna & Flora International, Oryx, 43(1), 113–121 Downloaded from https://www.cambridge.org/core.
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