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Issue 19, Autumn 1993
by Bernardino Ragni,
Introduction, Method and Materials
The Eurasian Lynx lynx (Linnaeus 1758) is a species belonging to the Italian fauna.
Proof of its presence is available since the Pleistocene Ice Age in northern and Alpine
Italy, as well as in peninsular Italy, and even on the island of Elba (Malatesta,
1950).
In the proto-historic and historic period subsequent to the Ice Age (Holocene epoch),
evidence of the presence of the species in peninsular Italy disappeared around the
Neolithic-Bronze Age (Grifoni & Radmilli, 1964; Wilkens, 1988; Wilkens, pers.
comm.) but continued to be found throughout the Italian Alpine region until the
first decade of this century (Ragni, 1988).
The most recent material data of the presence of the autochtonous Italian Alpine lynx
come from the Western Alps, Valdieri (Cuneo), 01.11.1909, stuffed specimen conserved
in the Bologna Civic Museum of Natural History; Central Alps, Valtellina (Sondrio),
1830, stuffed specimen conserved in the museum of the Institute of Zoology of Pavia
University; Eastern Alps, Auronzo di Cadore (Belluno), April 1837, stuffed specimen
conserved in the Civic Museum of Natural History of Venice.
Before the above mentioned times, the presence of the lynx in the Italian Alps is
indicated by 17 other stuffed specimens and by four skeletal remains conserved in
various Italian museums (Ragni et al. 1987; Mingozzi et al. 1988).
In the other Alpine countries, evidence of the presence of the species, before the
recent re-colonization, go back to the same period (Eiberle, 1972).
Several authors have tried to prove morphological differences between the Lynx lynx
population of the Alps and those of other European regions (e.g.: Tschudi, 1853;
Lavauden, 1930; Schauenberg, 1969).
In order to compare the extinct, autochthonous, Alpine population of Lynx lynx with
other European populations, we have reviewed the available bibliographical sources,
and carried out metrical and morphological examination of 11 skeletal specimens and
26 skin specimens. The material dates from the decade before 1800 and the following
decade, and comes from the Italian, French, Swiss and Austrian Alps.
We have recorded the coat marking colour system of 46 adult individuals of Eurasian
lynx, observed, photographed, trapped or killed from 1972 to 1991, in the Italian,
French, Swiss, Austrian and Yugoslav Alps.
In order to do the biometrical comparison we selected three craniometrical characters:
the total length (TL), the condilo-basal length (CDL) and the zygomatic breadth (ZB),
to be measured on specimens with permanent dentition. Such characters "describe"
satisfactorily the size of the skull. They are among the most simple and unequivocal
measurements to take, and are among the most used in morphological works on the
species (e.g.: Vasiliu & Decei, 1964; von den Driesch, 1976; Werdelin, 1981; Miric,
1981, Wiig & Andersen, 1986).
Other biometrical comparisons between the present lynx populations and the extinct
population of the Alps are impracticable, because of the unavailability of in carne
samples, and because of the high subjectivity of the few biometrical measures found
in literature about the Alpine lynx.
Results and Discussion
Comparison of the Alpine sample has only been possible with the Scandinavian
(Mazak, 1968; n = 16), Carpathian (Vasiliu & Decei, 1964 and Mazak,
1968; n = 34) and Balkan (Miric, 1981; n = 6) samples because these
are the only individual measurements of the parameters available.
The size of the Scandinavian and Carpathian lynxes appears to be significantly
greater than that of the Alpine, but the differences with the small Balkan sample
are not significant (Student's t-test, respectively: p0.05 and p0.050.1).
This pattern suggests that the extinct, autochthonous Alpine population of Lynx
lynx was one of the smallest in size in Europe. Nevertheless, the taxonomic and
evolutionary validity of such differences is compromised by the fact that all the
linear, skeletal characters of the Felids are subject to a strong environmental
influence and the extent of its genetic component is unknown (Werdelin, 1981;
Ragni & Randi, 1986).
Within the known variation of the coat marking-colour system of the Eurasian lynx,
we can recognize three types: spotted, striped and concolor. These patterns are
related to the somatic regions: occipis-cervicalis, scapularis, dorsalis, lateralis,
omeralis and femuralis.
The marking-colour patterns of such regions are subjected to strong individual
variations, being however fixed for each animal throughout its life from sexual
maturity.
The somatic regions of the spotted type show a prevailing pattern of permanent black
spots (Ragni, 1981); in the striped type the pattern is arranged in horizontal
stripes and bars of permanent-evanescent, brownish-black; the concolor-type shows the
somatic regions without marking or with a pattern so scattered and obsolescent that
it cannot be defined.
Such variation, continuous in appearance (Weigel, 1961; Vasiliu & Decei, 1964) is,
in effect, morphologically discrete for the presence of a constant occipis-cervicalis
and/or dorsalis marking in the first and second types and for its constant absence
in the third type.
The variations of the marking-colour system of the Felids are genetically controlled
and, in this case, it is possible to recognize a genetic homology with the multiple
allelism to the "Tabby" locus (Robinson, 1977).
In particular: the spotted coat and the striped coat can be considered a gradient of
phenotypic expression of the "spotted-striped" or "mackerel tabby" (Ts) allele, while
the allele controlling the concolor coat is referable to the "abyssinian tabby" (Ta).
Both alleles belong to the homologous genetic series well known in Felis and, to a
lesser extent, in other genera of the Felidae family: Panthera, Acinonyx, Leptailurus,
etc (Robinson, 1978; Lindburg, 1989).
Preliminary observations, conducted on captive crosses and on the frequency of the
phenotypes in samples from natural populations, support the probability that the Ta
gene was recessive to the Ts allele. Nevertheless there is the possibility that an
"incomplete dominance" relationship could exist between the two alleles (Robinson,
1977).
The percentage frequencies of the spotted-striped types (Ts) and of the concolor type
(Ta) among the samples from non-Alpine European populations and from the extinct,
autochthonous, Alpine population, were as follows:
Within the Lynx genus, the same constancy of the coat marking-colour system shown
by the late Alpine lynx, throughout such a large time-space extents, is known in
Europe for the pardel lynx Lynx pardinus (Temminck, 1824), of the Iberian
Peninsula. In fact, all known materials from this species, from the end of the
18th century to the present day, belong to the Ts-type (Weigel, 1961; Vasiliu &
Decei, 1964; Delibes, pers. comm.).
Such genetic characterization, besides the morphometric differentiation, permits the
retention of the hypothesis that the Alpine lynx was not a mere, undifferentiated
population of the Eurasian lynx. That population could be, instead, a well
differentiated taxon, even if very likely intra-specific (e.g., tentatively:
Lynx lynx alpina.)
The percentage frequencies of the spotted-stripes types (Ts) and of the concolor type
(Ta) among the samples from the present (1972-1991) Alpine population of the Eurasian
lynx, are as follows:
Conclusions
The available observations prove that the present lynx of the Alps, relative to the
coat marking-colour system, shows an inverse frequency of the alleles if compared
with the gene frequency of the extinct population.
This means that humans, within half a century, brought about the extinction of a
taxon and its complete substitution with another taxon throughout such a large
geographical area.
Notwithstanding the regret at the loss of a different form of life, the balance of
the present process of recolonization of the Italian Alps and the overall Alpine
Region by the "new lynx", can be considered active, both ecologically and culturally
(Breitenmoser & Breitenmoser-Wursten, 1990; Ragni & Possenti, 1991).
It will be very interesting to follow the variation of the coat marking-colour
frequencies throughout the time, as a de facto experiment in order to see if the
evolutionary forces of the environment, and the genetic variations, will lead the
present Alpine lynx back to a model similar to the extinct animal.
The sudden release of groups of this carnivore in large areas, where there have long
been no lynxes, offers to illuminate, almost experimentally, the most probable and
direct causes of the lynx extinction. The available data appear to support the
key-role of the availability and accessibility of the prey-species and populations to
the maintenance and development of the population of such a specialized and obligated
carnivore as Lynx lynx (Ragni & Possenti, 1991; Breitenmoser & Haller, 1993; Ragni
et al., in press). From that it is possible to deduce that the main cause of the
lynx extinction was direct destruction of the prey populations by humans. The phenomenon
appears to be consistent with the "Volterra Principle" (Volterra, 1926; Wilson & Bossert,
1974; Ragni & Possenti, 1992).
From the above exposition it arises that the first, if not unique, ecological requisite
for the development and conservation of a new Alpine population of this cat is the
presence of permanent populations of wild ungulates (Cervidae, Bovidae), and it would
be better if the populations could be diversified and at climax stage.
Because of its extension and environmental status, the Alpine Region appears to be the
only area of Central-Western Europe able to support a viable and permanent population
of Eurasian lynx, if people will allow it! The Italian Alpine sub-region (the Italian
Alps) is, indeed, the most extensive among those of the five countries sharing the
Alps because:
Summary
References
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