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The Lynx in the Italian Alps

CAT NEWS
Issue 19, Autumn 1993

by Bernardino Ragni,
Mariagrazia Possenti,
Stefano Mayr *

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
Average measures (millimetres) of the parameters

    TL,  CBL,   ZB,   n,   ref 
1.Finland,149.60,134.60,101.60,17,a
2.Sweden,149.20,135.20,102.50,16,b
3.Norway,148.10,134.75,98.35,130,c
4-5.W. Siberia & Eur. Russia,143.33,130.87, 99.45,102,d
6.Rumanian Carpathians,150.80,136.90,106.00,25,e
7.Polish Carpathians,154.00,137.60,107.00,1,f
8.Slovak Carpathians,153.50,137.50,106.47,126,g
9.Balkans,139.90,126.70,97.60,6,h
10.Caucasus,146.55,131.52,104.35,38,i
11.Alps,129.80,117.63,95.47,14,l
(n, size of the sample; a, Hell (1980); b, Mazak (1968); c, Wiig & Andersen (1986); d, Miric (1981); e, Vasiliu & Decei (1964); f, Mazak (1968); g, Hell (1980), Mazak (1968); h, Miric (1981); i, Miric (1981); l, present work.)

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:

 Ts,Ta,n,r
Balkans,79,21,14,1
Carpathians,89-78,11-22,150,2-3
Caucasus,75,25,?,4-5
Scandinavia,100 (striped),0,10,6
Europe,85.75-83,14.25-17,174, 
Alps,0,100,26,7
(n = sample size; 1, Miric (1981); 2, Vasiliu & Decei (1964); 3, Stollmann (1963); 4, Harrison (1968); 5, Ognev (1935); 6, Mazak (1968); 7, present work.)

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:

 Ts,Ta,n,ref
Italy,100,0,9,1
France,100,0,1,2
Switzerland,100,0,9,3
Austria,100,0,6,4
Slovenia,100,0,21,5
Alps,100,0,46, 
(n, sample size; 1, present work; 2, Kempf. et al. (1979); 3, Breitenmoser, Haller, Schloeth, pers. comm., Ragni; 4, Molinari, Gossow, pers. comm.; 5, Cop, pers. comm., Ragni.)


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:

  • only the Italian Alps connect with all the sub-regions of the other countries, by mean of a 1,600 km border;

  • only the Italian Alps can develop simultaneous wildlife gathering and turnover, from and to all Alps of the other countries;

  • the above conditions indicate that the Italian Alpine sub-region is the key and is irreplaceable in "rebuilding" a new European population of Alpine lynx.

Summary
The Eurasian lynx (Lynx lynx Linnaeus 1758) has been part of the Italian fauna since the Pleistocene Ice Age. Proofs of its presence disappeared from the whole peninsula by around the Neolithic-Bronze Age, but remained over the entire Alpine region until the beginning of this century. We have reviewed the literature, carried out morphological and metrical analysis of 11 skeletal specimens and 26 skin-specimens, and analyzed the marking-colour patterns of 46 adult individuals of the present Eurasian lynx population in the Alps to compare the extinct autochthonous Alpine lynx population with other European populations of Lynx lynx. We found the size of the Alpine lynx to be one of the smallest of the species in Europe. Among the marking-colour patterns of the coat of Eurasian lynx we recognize three types: spotted, striped and concolor. The spotted and striped coat patterns can be considered the result of differently graded, phenotypic expression of the "spotted-striped" or "mackerel" tabby (Ts) allele, while the allele controlling the concolor coat pattern can be referred to as the "abyssinian" tabby (Ta). The frequencies of Ts and Ta types in European non-Alpine populations vs. the extinct Alpine population are inversely distributed. It can be concluded that the Alpine lynx represents a well differentiated taxon (Lynx lynx alpina?), although very likely intra-specific. This suggests that in only 50 years man has brought about the extinction of a taxon in a large geographical area, and its complete replacement with a different one. The "new" lynx is slowly but progressively colonizing the Alpine region, which, because of its extent and environmental conditions, is the only area of Central-Western Europe capable of supporting a viable and permanent population of Lynx lynx.


References

  • Breitenmoser, u., C. Breitenmoser-Wursten. 1990. Status, conservation needs and reintroduction of the lynx (Lynx lynx) in Europe. Council of Europe, Nature and Environment Series, 45, Strasbourg, 47 pp.
  • Breitenmoser, u., H. Haller. 1993. Patterns of predation by reintroduced lynx in the Swiss Alps. J. Wildlife Manage. 57 (1): 135-144.
  • Driesch, A., von den. 1976. A guide to the measurement of animal bones from archaeological sites. Peabody Museum Bulletin, 1: 136 pp.
  • Eiberle, K. 1972. Lebensweise und Bedeutung des Lucheses in der Kulturlandschaft. Mammalia Depicta, Verlag P. Parey, Hamburg, 65 pp.
  • Grifoni, R., A. Radmilli. 1964. La Grotta Maritza e il Fucino prima dell'Età Romana. Rivista di Scienze Preistoriche (19), Firenze.
  • Harrison, D.L. 1968. The mammals of Arabia. Vol. 2: Carnivora, Artiodactyla, Hyracoidea. London.
  • Hell, P. 1980. Die situation des luchses in der Tschechoslowakei. In: Festetics, A. (Ed.), Der Luchs in Europa. Kilda-Verlag: 187-195.
  • Kempf, C., A. Balestrieri, U. Wotschikowsky, m. Fernex. 1979. Chez nous le lynx? Mythes et réalité. Les Guides Gesta, Paris: 152 pp.
  • Lavauden, L. 1930. Essai sur l'Histoire naturelle du Lynx. Aller & Fils, Grenoble: 108 pp.
  • Lindburg, D. 1989. When cheetahs are kings. Zoonooz, LXII, (3): 1 and 5-10.
  • Malatesta, A. 1950. Sulla grotta di Reale a Porto Azzurro (Elba). Rivista di Scienze Preistoriche (5), Firenze.
  • Mazak, V. 1968. Quelques donnees sur le Lynx de l'Europe du nord, Lynx lynx lynx (Linnaeus, 1758). Mammalia, 32, 3: 326-340.
  • Mingozzi, T., F. Guidali, G. Tosi. 1988. Dati storici sulla presenza della lince Lynx lynx (L.) nell'ia nord-occidentale.
  • Atti del I Convegno Nazionale dei Biologi della Selvaggina. Suppl. Ric. Biol. Selvaggina, XIV: 479-500.
  • Miric, D. 1981. The Lynx populations of the Balkan peninsula. Serbian Academy of Sciences and Arts, Vol. DXXXIX, (55): 150 pp.
  • Ognev, S.L. 1935. Mammals of USSR and adjacent countries. Vol. 3, Carnivora. Moscow.
  • Ragni. B. 1981. Gatto selvatico. In: Distribuzione e biologia di 22 specie di Mammiferi in ia. C.N.R., Roma: 105-113.
  • Ragni, B. 1988. Status e problemi di conservazione dei Felidi (Felidae) in ia. Atti del I Convegno Nazionale dei Biologi della Selvaggina. Suppl. Ric. Biol. Selvaggina, XIV: 455-478.
  • Ragni, B., L. Lapini, Fr. Perco. 1987. Situazione attuale del Gatto selvatico Felis s. silvestris e della Lince Lynx lynx nell'area delle Alpi Sud-Orientali. "Biogeografia delle Alpi Sud-Orientali", Biogeographia, XIII: 867-901.
  • Ragni, B., M. Possenti. 1991. Il ritorno della Lince nelle Alpi. Le Scienze (Scientific American), Milano (273): 54-63.
  • Ragni, B., M. Possenti. 1992. Predatori, prede e gestione faunistica. 54$ Congresso dell'U.Z.I., Perugia, 28 settembre-3 ottobre 1992, Riassunti, Relazioni e Posters, Perugia: 52-53.
  • Ragni, B., E. Randi. 1986. Multivariate analysis of craniometric characters in European wild cat, Domestic cat and African wild cat (genus Felis). Z. Saugetierkunde, 51: 243-251.
  • Ragni, B., M. Possenti, S. Mayr. In press. La lince nel Trentino.
  • Robinson, R. 1977. Genetics for Cat Breeders. Pergamon Press. London, 202 pp.
  • Robinson, R. 1978. Homologous coat color variation in Felids. Carnivore, 1, (1): 68-71.
  • Schauenburg, P. 1969. Le lynx Lynx lynx (L.) en Suisse et dans les pays voisins. Rev. Suisse de Zool., Genève, 76: 257-287.
  • Stollmann, A. 1963. Beitrag zur Kenntnies des Luchses, Lynx lynx (L.) in den tschechoslowakischen Karpaten. Folia Zoologica, Brno, 12: 301-316.
  • Tschudi, F. von. 1853. Das Thierleben der Alpenwelt. Leipzig: 590 pp.
  • Vasiliu, G.D., P. Decei. 1964. Uber den Luchs (Lynx lynx) der rumanischen Karpathen. Saugetierkundl. Mitt. Munchen, 12: 155-183.
  • Volterra, V. 1926. Variazioni e fluttuazioni del numero d'individui in specie animali conviventi. Memorie della R. Acc. Naz. dei Lincei, Vol. II, Fasc. III: 30-111.
  • Weigel, I. 1961. Das Fellmuster der wildlebenden Katzenarten und der Hauskatze in vergleichender und stammesgeschichtlicher Hinsicht. Saugetierkundl. Mitt., Munchen. 9, Sonderheft: 1-120.
  • Werdelin, L. 1981. The evolution of lynxes. Ann. Zool. Fennici, 18: 37-71.
  • Wiig, O., T. Anderson. 1986. Sexual Size Dimorphism in the Skull of Norwegian Lynx. Acta Theriologica, Vol. 31, 12: 147-155.
  • Wilkens, B. 1988. La fauna di Coppetella (Iesi, Marche). Atti della Soc. Toscana di Sc. Naturali (95), Firenze.
  • Wilson, E.O., W.H. Bissert. 1974. A primer of population biology. Sinauer Associates, Publ. 197 pp.




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