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Resurgence of Egyptian Vultures in Western Pyrenees^ and Relationship

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Resurgence of Egyptian Vultures in Western Pyrenees^ and Relationship Resurgence of Egyptian Vultures in western Pyrenees^ and relationship with Griffon Vultures Jacques Carlon This paper is dedicated to the memory of Bernard Braillon, who died on 26th December 1986, in recognition of his pioneering work on this species on the northern slopes of the Pyrenees. ABSTRACT A long-term study of Egyptian Vultures Neophron percnopterus in southwest France showed a marked population recovery from 1985, the species having declined greatly during the previous decade and more. This resurgence was due in part to improved weather during critical periods of the breeding cycle, and also reflected improvements in the fortunes of the Spanish population. The species' relationship with the Griffon Vulture Gyps fulvus was also studied, and its response to nest-site usurpation by the larger vulture showed interesting adaptations. Our observations on Egyptian Vultures Neophron percnopterus were carried out in the province of Beam, in the eastern half of the departement of Pyrenees- Atlantiques, situated on the northern slope of the Pyrenees in southwest France (see Carlon 1996b). The study area comprised the valleys of the rivers Baretous, Aspe, Ossau and Ouzom, all in the southern half of the province and covering an area of approximately 860 km2. [Brit. Birds 91: 409-416, October 1998] © British Birds Ltd 1998 409 410 Carlon: Egyptian and Griffon Vultures in western Pyrenees During the 12 years 1984-95, a total of 2,366 hours of field observations was made by the author, with an additional 330 hours by colleagues. Altogether, 205 pairs of Egyptian Vultures were monitored at breeding sites over the 12 years, plus ten pairs in 1982 and 1983: making 215 pairs in all. Various findings of this study have already been published in French (Carlon 1989, 1992, 1993, 1996a). The present paper presents a summary in English of the remarkable resurgence of the species in Beam, together with interesting aspects of its relationship with the Griffon Vulture Gyps fulvus. Decline and regrowth of the population The population of the Egyptian Vulture in southern France and the Pyrenees underwent a strong decline in the years preceding the 1980s, but has since recovered to a healthier level. This was apparent not only in our study area, but also in the neighbouring regions of Provence, in southeast France (Bergier 1985), and Aragon, on the southern slope of the Pyrenees in Spain (Kostrzewa et al. 1986). During the main period of our study, regular checking of sites that had been abandoned for more than ten years, together with a systematic search of all potential breeding sites, enabled us to record the reoccupation of six sites, the discovery of ten new ones, and successful breeding at a site where a regular pair had taken a seven-year 'sabbatical' during 1977-83. Table 1 summarises the annual development of the Beam population from 1984 to 1995. The years 1982 and 1983, when our fieldwork was less intensive, are also included, to highlight the low point of this population (six pairs in 1984) and the subsequent spectacular recovery and growth (to 21 pairs in 1990). The population stabilised at 20-21 pairs during 1990-95. This apparent stagnation in the 1990s could, we believe, be linked with two factors. First, usurpation of eyries and competition for habitat by the Griffon Vulture represent an appreciable limiting factor (see below); and, secondly, observer effort and efficiency tend, through the process of habituation, to decrease over the course of the years, particularly with regard to searching for potential new breeding sites. It is possible, if not probable, that two or even three sites may have escaped detection in 1994 and 1995, bearing in mind the large extent of suitable breeding habitat available in the study area. Site fidelity A striking feature of this study was the great fidelity of pairs to breeding sites, so long as they were not subject to heavy disturbance or interspecific parasitism. Apart from instances of nest-site usurpation by Griffon Vultures (see below), we recorded only two definite cases of site desertion out of 205 pairs monitored. The first occurred at the highest site in the study area, at 1,450 m, within the montane and subalpine zone; this was a result of unusual weather conditions, with heavy snowfalls in two years out of three in April, the month when the pair was establishing territory. The second was in Vallee d'Aspe, very probably caused by disturbance from increasingly heavy traffic on the roadway; the pair involved re-established itself at a site 300 m higher up. British Birds, vol. 91, no. 10, October 1998 411 Factors responsible for the resurgence of the population Climatic influences The influence of climatic conditions on population fluctuations and reproduction was discussed by Carlon (1992). Two interesting facts have emerged from this study: the low percentage of breeding failures, and the development of earlier and more successful fledging. In years with 'normal' weather conditions, breeding failures appear to be relatively rare. In Provence, Bergier & Cheylan (1980) recorded only four failures (6.8%) out of 59 breeding attempts. In Beam, of 66 breeding attempts over the five years 1985-88 & 1990, only five failed (7.5%, a figure roughly equivalent to theirs), even though the Mediterranean climate is more favourable to the species. On the other hand, the Provencale region is subject to much greater disturbance, owing to the greater accessibility of nesting sites. Breeding success across the entire northern slope of the Pyrenees in normal years during 1959-85 was shown to be 77% (Braillon 1986). In Beam, during 1985-90, we recorded only ten breeding failures (plus two suspected failures) out of 91 pairs, giving a success rate of 88%. This improvement appears to be due to better weather conditions during the early stages of development of the young. This was borne out in 1991, when six pairs failed out of a total of 21, all but one (which was probably due to human disturbance) at or soon after hatching. In that year, precipitation was 15.1% less than in the three preceding years, but temperature and total sunshine hours were both 14-15% lower, and these last two parameters would appear to be of the greatest importance during the critical period of hatching and early development. Note that cold, wet springs, such as occurred in 1988, do not necessarily lead to failure if hatching takes place after such weather has ceased. During the study period, a distinct advancement in fledging date was noted. Up to 1987, the mean fledging date in Beam was about 20th August; during the two drought years which followed (1989, 1990), it was 12th August, where it remained. Owing to much earlier fledging at some sites in 1994—2nd and 5th August, 25th and 30th July, and an unprecedented 11th July in Vallee d'Aspe (M. & R. Cruse in litt.)—the mean date for fledging has been brought forward to 25th July. We also recorded a small increase in productivity, with a higher number of juveniles reared per pair during 1988-95 (mean 1.26/pair) than during 1959- 85 (1.17/pair) (table 2). Given that 1989 and 1990 were drought years, we demonstrated (Carlon 1992) that higher temperatures, increased total sunshine hours and a drop in total precipitation were determining factors in this increase in fledging rate. As recording fledging success requires an enormous amount of time in the field, it was not possible to determine this throughout the entire 12-year period. Nevertheless, we were able to calculate it for six years (1988-90, 1991- 92 and 1995; table 2). These involved a significant number of breeding attempts (88), and a representative sample of climatic conditions which included two years (1989-90) of dry, hot weather, followed by two years (1991-92) with particularly cold and wet weather throughout the most critical 412 Carlon: Egyptian and Griffon Vultures in western Pyrenees stages (May-June: end of incubation, hatching and first days of life of the chicks), and then by one year of drought (1995). On the basis of these criteria, and a rigorous follow-up during the three periods, we consider our figure of 1.26 young/pair a realistic one, corroborated by and comparable with those published for Provence (1.38-1.40: Bergier & Cheylan 1980; Bergier 1985; Bayle 1990) and for the Spanish slope of the Pyrenees in Catalonia and Navarra (1.29-1.75: Donazar & Ceballos-Ruiz 1988). Note that this increase in fledging rate is due almost solely to the higher percentage of broods of two young in years when weather conditions were very favourable for this sun-loving species. Especially when climatic conditions have been propitious, fieldwork needs to be concentrated during and subsequent to the period when the young leave the nest (about the third week in July to late August); this is particularly true when the brood contains two young. Other possible influences As weE as these climatic factors that have played a major role on the two sides of the Pyrenean massif, particularly in Beam, there are others, not insignificant, that need to be examined (see Carlon 1989). The resurgence of the population may best be appreciated by dividing the period into two parts. In 1982-85, the combined number of occupied sites was 33 (possibly 35), or an annual mean of eight or nine; in 1986-88, this total rose to 46, or a mean of 15 or 16 per year (table 1). During 1979-83, Bergier (1985) recorded, in Provence, the recolonisation of a territory abandoned since 1965, the occupation of a new site, and the observation of several immatures during the breeding season, while in Aragon, Spain, in 1984-85, a population increase was confirmed for the first time (Kostrzewa et al.
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