Nest sites
Historical references provided information on the geographical
location of 10 nests (see Fig. 1 and Appendix 1, Nos 16, 30, 33;
Appendix 2, Nos 1, 3, 4, 6, 7, 12, 13). In most cases, this
information was quite general, i.e. the name of a mountain or a pass,
while more precise details were lacking. As an exception, we quote No.
16 (Appendix 1), referring to the nest destroyed in the Vallon de
Ceillac (F) in April 1872, for which a quite precise site description
was found in an unpublished document (P. G. Berlie, ms). Even so, this
site is not precisely identifiable today (J. F. Terrasse, pers. comm.).
We carried out special research into three other sites, Nos 7, 12, 13
in Appendix 2; only the first one, on the eastern cliffs of the Signal
de St Chaffrey (Briançon, F), was in some way recognizable.
Nevertheless, the historical data appeared sufficiently detailed to
attempt an evaluation of the distance between neighbouring nest sites,
such as the three well-documented ones for the Briançon region, i.e.
No. 30, Appendix 1 (Grand Aréa, Vallée de la Guisanne), No. 7,
Appendix 2 (Signal de St Chaffrey) and No. 13, Appendix 2 (Montgenèvre
Pass), 9 km (No. 30 vs No. 7) and 7 km (No. 7 vs No. 13) apart.
However, we do not know whether the three nest sites were ever
simultaneously occupied since the data referred to a time span longer
than 20 years.
Specimens still preserved in museums and collections
Thirty-two specimens shot in the western Alps are still preserved in
natural history museums and collections of France (18), Italy (12) and
the United Kingdom (two) (23 were killed in the French Alps, seven in
the Italian and two on the border of the two countries; see Appendix
1).
Despite our checking, some uncertainty regarding this list still
remains. In fact, bibliographical and museological data did not always
correspond, as shown by the four specimens quoted for the Grenoble
Museum (F): only three of them, with incomplete and only partially
matching data labels, were present in this collection (M. Dunand,
pers. comm.).
Six
more specimens, lacking data labels, are preserved in French museums
and collections: one juvenile in Nîmes Museum (G. Gory, pers. comm.),
one individual in Guimet Museum, Lyon (J. Clary, pers. comm.), one
individual in Hyeres Museum (C. Nicolai, pers. comm.), two individuals
in Marseille Museum (A. Delcourt, pers. comm.) and one adult in a
private collection in Barcellonnette (C. Joulot, pers. comm.). The
origin of these specimens remains uncertain, even though they were
probably shot in the western Alps.
DISCUSSION
Distribution of records
Figure 1 shows a tendency of records to cluster in different alpine
sectors, namely the area including the Valdieri valleys and the Tende
Pass in the Italian Maritime Alps (12 records at least), the Briançon
region in the French Cottian Alps (12 records) and the Gran Paradiso
Massif in the Italian Graian Alps (nine records).
The
two Italian alpine sectors (both protected areas today) were formerly
royal game preserves of the House of Savoy; as a consequence, two
factors should explain their richness of records. First, favourable
habitat conditions and a consequently high population density of the
species: these zones were certainly richer in wild ungulate
populations (i.e. chamois Rupicapra rupicapra and ibex Capra
ibex) - potential sources of food - than the surrounding areas
(Richard, 1914). Thus, their ecological suitability and attractiveness
had to be greater for the vulture, just as it is nowadays (almost all
of its spontaneous reappearances in the western Italian Alps between
1977 and 1980 were recorded here, Mingozzi, 1981). Second, a better
availability of historical information: the shooting of every large
carnivore was encouraged (see below) and normally reported to the
Royal Zoological Museum of Turin.
The
richness of records of the Briançon region cannot be attributed to a
special source of information but was likely due to a previous high
population density. On the assumption that the three reported nest
sites were simultaneously occupied by neighbouring pairs, the
distances between them correspond to a breeding density as high as
presently estimated in the Central Spanish Pyrenees (average distance
= 8.813 km; Donazar et al., 1993).
Extirpation time pattern
Bailly (1853) and Salvadori (1872) stated that the bearded vulture
exhibited an initial population decrease in the first half of the 19th
century. On the whole, historical evidence (see also Roux, 1825;
Temminck, 1835; Durazzo, 1840; Jaubert & Lapommeraye, 1859) indicates
that the species was widespread and common at the end of the 18th to
beginning of the 19th centuries and then, 50-60 years later, quite
rare in different parts of the western Alps.
Nevertheless, around 1850-1860 the total population may still have
been quite substantial, even if it was probably reduced in range. A
lot of birds were in fact still shot in the following 60-70 years and
the vulture was still not considered rare around 1860-1870 in some
alpine sectors, in particular Ubaye and Briançonnais (Berlie, 1898;
Rérolle, 1900).
However, a clear pattern of rarefaction has been depicted by
historical data only since the end of the 19th century. Indeed, Fig. 3
reveals a great increase in the number of birds shot during the decade
1890-1899 and then a rapid decline until final extirpation, 30 years
later.
It
is difficult to confirm the reality of this trend, although a very
similar extirpation pattern was depicted for another alpine species,
the lynx (see Mingozzi, 1995). Clearly, information was lacking before
the decade 1890-1899 since shooting became an event worth reporting at
the same rate as bird rarity increased.
In
any case, the last juvenile vulture was shot in 1904 in the Valdieri
valleys (No. 49, Appendix 1) while the last evidence of breeding was
in 1907 at Montgenèvre Pass in the Cottian Alps (No. 13, Appendix 2).
Since then there has been only one breeding record, more indefinite
than the former and relating to the Italian Maritime Alps for 1914
(No. 9, Appendix 3), but no further killing or live observations of
juvenile birds were recorded. Therefore, the data seem to indicate
that reproduction stopped around 1910, while the species probably
survived for a further 15-20 years (the normal life life expectancy of
a large raptor). Two birds were shot in 1920 in the French Hautes
Alpes (see above and Appendices 1 and 2) and it seems that one or two
individuals were present in the Gran Paradiso National Park between
1924 and 1930 (see above and Appendix 2). No further reliable records
appeared after 1930 until the well-known spontaneous reappearance of
the species between 1970 and 1980 (Tosi, 1978; Huboux, 1980; Tosi &
Piantanida, 1980; Géroudet, 1981; Mingozzi, 1981).
Causes of extirpation
Past authors (Rérolle, 1900; Lavauden, 1911) attributed the decrease
of the bearded vulture populations to the disappearance of other large
carnivorous vertebrates, mainly the wolf, but also the brown
bear and the lynx. However, one may note that the bird also lives in
areas where none of these three mammals has been present since
historical times (e.g. Corsica, Crete) or where they are very scarce
(Pyrenees).
Recent authors (see, for example, Géroudet, 1974; Hiraldo et al.,
1979; Walter, in Mundy, 1985) have stated that deliberate human
persecution was the main reason for extirpation in the Alps; the same
opinion was also held by Hiraldo et al. (1979) for Spain.
Haller (1983) postulated that climatic cooling at the beginning of the
17th and early 19th centuries in Europe probably had a negative
influence on living conditions of the species.
Moreover, Coton and Estève (1990) listed the decrease of wild ungulate
populations among the possible extirpation causes. During the 19th
century the ibex became almost extinct in the whole alpine chain and
the chamois was certainly not as common as nowadays (see de
Tschudi, 1859; Perlini, 1923). However, as pointed out by past authors
(e.g. Bailly, 1853; de Tschudi, 1859; Rérolle, 1900), sheep, goats,
cattle and other livestock, rather than wild ungulates, were a much
more important potential food resource to the vulture, in the last
century at least. In the Aosta valley alone the official census of
1881 revealed 44,488 cattle, 35,030 sheep and 21,648 goats (Janin,
1991).
Analysis of the historical evidence has shown that direct human
persecution affected the lammergeier populations in the western Alps,
at least since the beginning of the 19th century.
It
appears that the killing pressure in the early 19th century was not as
deliberate and powerful as in the following decades. At that time the
commonest hunting technique used in the Piedmont Alps was a simple
ditch in the ground with a carcass inside, where birds jumped in to
feed and were beaten to death with sticks (de Tschudi, 1859).
Accordingly, the rarefaction process claimed by authors for the first
half of that century was probably due to other natural or indirect
human causes. However, not enough is known about environmental changes
and land use transformations. The early decline of the vulture in the
central and eastern Alps, where extirpation took place 30-40 years
before that of the western Alps (see Glutz von Blotzheim et al.,
1971), might also have played a part in the process, reducing any
chance of immigration and thus insularizing the western alpine
population.
From the middle of the 19th century direct human persecution achieved
great prominence and effectiveness for at least two reasons:
collection for taxidermy became highly popular (see Fatio & Studer,
1889, in Richard, 1914) and pest control became a widespread method of
game management; in the House of Savoy royal preserves, bounties of 50
lire, or 200 to 500 francs (equivalent to a monthly wage of a game
warden) were paid for each bird shot (Rérolle, 1900; Couturier, 1962;
Videsott, 1971).
The
shotgun appears to be the means most commonly used everywhere in the
western Alps; nearly all the 65-67 birds listed in Appendix 1 were
certainly or likely killed in this way. Seemingly, no technical
improvement in firearms favoured such a destruction; the breech loader
shotgun came into common usage in the royal preserves only at the
beginning of this century (photographic archives of GPNP, Turin).
Rather, as pointed out by Berlie (1898), meat baits poisoned with
strychnine - already widely used for fox Vulpes vulpes control
- perhaps played a more important destructive role than first
appeared; although only one bird (perhaps two) in our list (No. 28 in
Appendix 1; Salvadori, 1895) was killed by strychnine, obviously birds
that died for this reason were seldom found.
Persecution affected mostly adult birds in the crucial phase of the
breeding season (see above). As a consequence, the effect on the
populations would have been especially severe (see Hiraldo et al.,
1979; Newton, 1979).
CONCLUSIONS
Assessment of the former range and analysis of the causes of
extinction are the first two management problems that must be
addressed in any reintroduction programme (see Boitani, 1976).
The
present study has ascertained the former distribution, the range
reduction and the most likely date of disappearance of the bearded
vulture in the western Alps. As far as the causes of extirpation are
concerned, two clear considerations emerged from the analysis of the
historical data: the species was subject to intense human persecution
from the middle of the 19th century at least and there is no evidence
that, in the same period, natural causes or indirect human activities
had a significant effect on population decline.
From a historical point of view, these general conclusions would
justify the current international project of reintroduction of the
species into the western Alps, in particular to the
Argentera-Mercantour Massif, one of the best documented and previously
occupied sites.
The
destructive effects of human persecution on raptor populations have
been demonstrated in various case studies (e.g. Love, 1983).
Nevertheless, the present analysis is still insufficient to confirm
persecution as the major cause of extirpation, even if the evidence
suggests this hypothesis. The real extent of persecution endured by
the species is not ascertainable; as a consequence, it is not possible
to assess whether the human persecution shown by the data would have
been sufficiently intense to result in extermination of the bearded
vulture population in the space of 30-40 years.
To
this purpose, the rapid population decline should be compared with the
biological vulnerability of the species; by making assumptions about
population size at various survival rates, it should be possible to
assess what rate of additional adult mortality caused by people would
have led to the historically observed rate of extirpation. A
Population Vulnerability Analysis (PVA) (Gilpin & Soulé, 1986) to
examine human impact is discussed elsewhere (Mingozzi & Balletto, in
press).
ACKNOWLEDGMENTS
The
authors would like to thank all museum curators for help with the
historical research, namely: J. Clary, A. Delcourt, M. Dunand,
G. Gory, C.
Nicolai, F. Vire. We are
indebted to M. Bocca, J.
Carriat, P. Fasce, F.
Genero, R. Janavel, C.
Joulot and A. Pazzuconi for their collaboration. Special thanks are
due to Dr V. Barello for his irreplaceable willingness to help in
bibliographical research.
We gratefully acknowledge Dr M. Ferrer and one anonymous referee for
their constructive criticism of a first version of this manuscript.
For this research T. Mingozzi was supported by a financial
contribution from: Associazione Italiana per il World Wildlife Fund,
WWF ITALIA, via Salaria 290, 00199 Rome, Italy.
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