dormice of the mediterranean region naturaland ... riassunto – scoiattoli e ... (waldren, 1982;...

INTRODUCTION

The Sciuridae family represents a largeand varied taxonomic group of rodentswith many forms adapted for arboreallife, including the true squirrels. Theyoccur today in all continents exceptAustralia. Sciuridae appeared as earlyas the Lower Oligocene (38-37 millionsyears ago) both in Europe (genusPalaeosciurus) and in North America(genus Protosciurus) (Vianey-Liaud,

1985) and have evolved considerably,producing flying, arboreal and terrestri-al forms, especially as regards theAmerican ground squirrels (Mein et al.,1993). In Europe, the genus Sciurus hasapparently existed since the Miocene(26-7 millions years ago), and a formsomewhat larger than Sciurus vulgarisL., 1758, described from Poland, hasbeen referred to the late Pliocenechronologies and may have survived inthe Villafranchian (cf. Kurtén, 1968).

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Hystrix It. J. Mamm. (n.s.) 16 (1) (2005): 3-26

NATURAL AND ANTHROPOCHOROUS SQUIRRELS ANDDORMICE OF THE MEDITERRANEAN REGION

MARCO MASSETI

Dipartimento di Biologia Animale e Genetica ‘Leo Pardi’, Università di Firenze.Laboratori di Antropologia, Via del Proconsolo 12, 50122 Firenze

E-mail: [email protected]

ABSTRACT – In the Mediterranean Region, squirrels and dormice of natural and anthro-pochorous occurrence are today represented by 7 taxa on each. Palaeontological evidencesuggests that Upper Pleistocene dispersal of the representatives of the Gliridae family seemsto have occurred only on the islands of the western Mediterranean basin, whereas squirrelsare not represented in insular environments. The former and present distribution of some ofthe extant species is discussed in the light of the human redefinition of the natural equilib-rium which has been taking place since very ancient times.

Key words: squirrels, Sciuridae, dormice, Gliridae, Mediterranean Region

RIASSUNTO – Scoiattoli e ghiri a diffusione naturale ed antropocora della RegioneMediterranea. Attualmente, nella Regione Mediterranea, gli scoiattoli ed i ghiri a diffusionenaturale ed antropocora sono rappresentati rispettivamente da 7 taxa. Il dato paleontologicosuggerisce che la distribuzione insulare dei Gliridi nel corso del Pleistocene superiore abbiainteressato solo il Mediterraneo occidentale, mentre gli scoiattoli non sembrano essere statirappresentati in nessuno degli ambienti insulari. Si discute la diffusione passata e presentedi alcune delle specie attuali nell’ottica della plurimillenaria ridefinizione antropica degliequilibri naturali dell’area mediterranea.

Parole chiave: scoiattoli, Sciuridae, ghiri, Gliridae, Regione Mediterranea

Various other European records canonly be determined as Sciurus sp.Remains referred to Sciurus sp. havealso been provided by the palaeontolog-ical exploration of the Neogene site ofTung-gur, in China (Qiu, 1989). Theearliest records of the Near Easternform, Sciurus anomalus Güldenstaedt,1785, are from the Late Acheulian peri-od (about 150.000 years BC)(Tchernov, 1988). In North-WestAfrica, the genus Atlantoxerus isknown since the beginning of theMiocene and it might have been adescendent of the genus Heteroxerus,known from Europe (Kowalski andRzebik-Kowalska, 1991). The extantBarbary ground squirrel, Atlantoxerusgetulus (L., 1758), is not known beforethe Pleistocene and does not seem to bea descendant of extinct species of thatgenus present in Maghreb in earlierperiods. It might have originated fromtaxa inhabiting regions south of theSahara (Kowalski and Rzebik-Kowalska, 1991).The dormice form one of the mostancient rodent families, emerging in theEocene (54-53 – 38-37 millions yearsago) (Nadachoswji and Daoud, 1994).They are small to medium sizedrodents, mostly arboreal, and arerestricted to the Old World. In fact, thegeographical range of the family (fossiland extant) is limited to Europe, Asiaand Africa. The fossil record of this tax-onomic group is relatively good, proba-bly because these animals are frequent-ly caught by owls so that their remainsfind their way into the deposits formedby owl pellets (Kurtén, 1968). The old-est record is from Europe, Eogliravuswildi Hartenberger, 1971 (Mas de

Gimel, France, Early Eocene) (Daamand de Bruijn, 1994). Palaeontologicalevidence suggests that the dormice areof European origin and that they devel-oped from the ischyromyoid branch ofthe Rodentia during a radiation that fol-lowed the opening of the North Atlanticduring the Early Eocene (Daam and deBruijn, 1994). The diversification of theGliridae which began in the EarlyEocene continued during the Oligoceneand culminated in the Late EarlyMiocene of Europe, where they appearto have occupied many ecological nich-es (Daam and de Bruijn, 1994). Thedecline of this taxonomic group, interms of diversity as well as of relativeabundance in assemblages of fossilrodents, becomes apparent during theLate Middle Miocene, that is, beforethe arrival of the first Muridae inEurope (Daam and de Bruijn, 1994).From the Late Miocene onwards theGliridae are represented in Europe, Asiaand Africa by a few genera only,excluding the insular forms; a situationthat has substantially persisted up to thepresent (Daam and de Bruijn, 1994).According to Kurtén (1968), the mod-ern subfamily Glirinae arose in theOligocene and is particularly well rep-resented in the Miocene, when its firstradiation took place. More modernizedglirines evolved in the Pliocene andQuaternary, but relicts of the Mioceneradiation survived locally in thePleistocene, for instance on several ofthe Mediterranean islands.

1. THE HOLOCENE SURVIVAL OF ENDEMIC

INSULAR FORMS

From the Mediterranean islands

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palaeontological evidence reports fossilremains of only the Gliridae taxonomicgroup, and not of squirrels. The archae-ological exploration of Sicily, for exam-ple, provided remains of one endemicspecies of Villafranchian age (faunalstage of Monte Pellegrino), taxonomi-cally attributed to Maltamys cf.gollcheri (de Bruijn, 1966) (Thaler,1972). There is also evidence of theoccurrence in the Middle/UpperPleistocene of the Siculo-Maltese archi-pelago of at least three differentdormice, such as Leithia melitensis(Adams, 1863), Leithia carthei(Adams, 1863), and Eliomys wied-incitensis (Zammit Maempel and deBruijn, 1982), yielded by the archaeo-logical investigation of the Mid-dle/Upper Pleistocene faunal stages ofSpinagallo and Maccagnone (Burgio,1997). The first two taxa were original-ly included by Adams (1863 and 1870)in the same genus Myoxus, formerlyreferred to the extant edible dormouse,whereas the latter was initiallydescribed as belonging to the peculiargenus Maltamys (Alcover et al., 1999).The giant dormouse, L. melitensis, wastwice the size of the extant edible dor-mouse, Glis glis (L., 1766) (Petronio,1970). L. carthei was instead describedas a form of smaller size, “… but notdiffering in any other respect from theMyoxus melitensis” (Adams, 1870).Among the characteristic featuresyielded by the Quaternary endemic fau-nas of the Mediterranean islands,rodents often display the tendency tobecome larger than their mainlandcounterparts (Azzaroli, 1971 and 1977;Sondaar, 1971, 1977 and 1986). Thisphenomenon is generally assumed to be

primarily a consequence of the geneticisolation from continental populations,a quantitative and qualitative reductionin food supply, an alteration of intraspe-cific competition, the absence of largecarnivores and perhaps also ofendothermic adaptations (Masseti andMazza, 1996).Palaeontological evidence suggests thatlate Pleistocene dispersal of the repre-sentatives of the Gliridae family seemsto have occurred only on the islands ofthe western Mediterranean basin (Fig.1). According to Boekschoten andSondaar (1972), glirids are in factabsent from the eastern islands wherethey were more or less replaced bydiversified murids. This difference maybe caused by faunal differences on thenearest mainland, and would lead us toassume a predominance of glirids in thewestern Mediterranean coastal areasand of murids in the easternMediterranean at the time of immigra-tion. Some taxa, such as Eliomys mor-pheus (Bate, 1918) recorded from theBalearic islands of Mallorca andMenorca, are distinctly relics from theTertiary, but survived up to latePleistocene, and even Holocenechronologies (Alcover et al., 1999).Also this was a giant dormouse, initial-ly described in the genus HypnomysBate, 1918, which was later on recog-nized as a subgenus of Eliomys byZammit Maempel and de Bruijn(1982). The extinction of the Balearicdormouse is possibly linked to theappearance of man on the two islands,perhaps not before the 3rd millenniumB.C. On Mallorca the most recent datefor remains of E. morpheus, providedby the exploration of Cova Estreta is c.

Sciuridae and Gliridae in Mediterranean Region

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6.000 years bp (Encinas and Alcover,1997). Human occupation layers datedto 4.000 years bp lack this species(Waldren, 1982; Sanders and Reumer,1984; Lewthwaite, 1985), indicatingthat the extinction of the dormouseprobably took place between these twodates (Alcover et al., 1999). Also E.wiedincitensis has been included byAlcover et al. (1999) among the endem-ic forms that survived in the Holoceneof Malta.

2. PRESENT NATURAL AND ANTHRO-POCHORUS DISTRIBUTION OF SQUIRRELS

IN THE MEDITERRANEAN REGION

Of the tree-squirrels, the genus Sciurusis represented by two species in thePalaearctic: the European red squirrel,Sciurus vulgaris L., 1758, in Europe

and northern Asia and the Persian squir-rel, Sciurus anomalus Gueldenstaedt,1785, in southwest Asia. Together withthe Barbary ground squirrel,Atlantoxerus getulus (L., 1758), dis-persed in north-western Africa, thesquirrels of natural - non anthro-pochorous - occurrence along theMediterranean shores are today repre-sented only by these 3 taxa.

2.1 Persian squirrel or golden squirrel,Sciurus anomalus Gueldenstaedt, 1785

This is a medium-sized arboreal squir-rel, which occurs in Asia Minor,Northern Arabia, including Lebanon,Syria, Jordan and northern Israel,Transcaucasia and western Iran(Harrison and Bates, 1991; Gavish,1993; Gavish and Gurnell, 1999;

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Figure 1 - Middle and Upper Pleistocene distribution of endemic glirids on theMediterranean islands.

Mendelsshon and Yom-Tov, 1999). Thepresence of this species has also beenreported from the Mediterraneanislands of Gökçeada (Imbros), Turkey(Özkan, 1995 and 1999; Gavish andGurnell, 1999) and Lesbos, Greece(Ondrias, 1966; Hecht-Markou, 1994and 1999; Gavish and Gurnell, 1999),

in the Eastern Aegean sea (Fig. 2). Thenatural colonisation of these twoislands by the species cannot be exclud-ed.

2.2 European red squirrel, Sciurus vul-garis L., 1758

This is a medium sized arboreal squir-rel, essentially characterised by apalaearctic distribution, ranging fromthe British Isles in the west, south to theMediterranean, the Caucasus – where it

has been introduced – the southernUrals and the Altai mountains in centralMongolia, to north-east China (Gurnelland Wauters, 1999b). Red squirrels arealso found on the island of Sakhalin offthe eastern coast of Russia, and on themost northerly Japanese island ofHokkaido. Perhaps with the exception

of Euboea, the natural occurrence of thespecies on Mediterranean insular envi-ronments is practically unknown, andtheir presence on some of these islands,such as Veli Brijuni (Croatia) (Scotti,1980), is essentially regarded as a con-sequence of recent human intervention(Fig. 3).

2.3 Barbary ground squirrel,Atlantoxerus getulus (L., 1758)

This is a monotypic species native to


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Figure 2 - Present occurrence of the Persian squirrel in the Mediterranean islands.

north-western Africa, where it is natu-rally widespread in Morocco, Grandand Middle Atlas (up to 4.000 m) southto Agadir and north-west Algeria up toEl Abiod Sidi Cheikh in the east

(Kowalski and Rzebik-Kowalska,1991). From northern Maghreb thistaxon has been introduced onto theCanary archipelago (Moreno, 1992), in1965 on the island of Fuerteventura

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Figure 3 - Present occurrence of the red squirrel in the Mediterranean islands.

(Machado Carrillo, 1985; Moreno,1992; Rodriguez Luengo and GarciaCasanova, 2002), and later on in GranCanaria (Calabuig, 1999; RodriguezLuengo and Garcia Casanova, 2002).Scientific travellers and other authorsof the past have occasionally reportedthe diffusion on the Mediterraneanislands of squirrels that are today com-pletely unknown among the respectivefaunal assemblages. Werner (1928), forexample, quoted the occurrence of akind of squirrel on the island of Skyros,in the archipelago of the NorthernSporades (north-western Aegean sea,Greece), where he collected a specimenbetween the villages of Skyros andLinaria which he described as “Sciuruslilaeus”. According to Ellerman andMorrison-Scott (1951), this taxon isused to define a Greek subspecies of thered squirrel, Sciurus vulgaris lilaeusMiller, 1907, characteristic of theregion of Mount Parnassus in continen-tal Greece. Nevertheless, the occur-rence of the same species on Skyroswas subsequently also recorded byother authors such as Wettstein (1942),followed in very recent times byCheylan (1988). On the basis of theauthority of Werner, and to an evengreater extent that of Wettstein, it isvery difficult to refute the truth of thesereports, even if red squirrels are todaycompletely unknown on Skyros and theother islands of the Aegean basin, per-haps with the exception of Euboea.Furthermore, it is interesting to recallthat these rodents are also practicallyunknown in the majority of the remain-ing Mediterranean islands, where, asalready observed, representatives of thegenus Sciurus L., 1758, occur only on

Lesbos and Gökçeada. As alreadyobserved, the latter two islands are,however, inhabited by another speciesof squirrel, the Persian squirrel, whosewesternmost continental distributionextends to far-eastern Europe and west-ern Anatolia. At the same time, howev-er, there is no evidence to exclude theformer diffusion of red squirrels onSkyros, where a population of thespecies could have existed up to thefirst half of the twentieth century, laterbecoming extinct. Red squirrels couldhave been imported by man into theisland from Euboea, where their pres-ence was reported by Lindermayer(1835) in the first half of the nineteenthcentury. In the light of modern ethnozo-ological enquiry, it would also appearthat squirrels figure among thosespecies of mammals which have beenthe subject of particular human atten-tion for a variety of cultural purposes,and/or for food. In some areas of thesouthern Levant (Jordan), for example,people still eat Persian squirrels andlive specimens are regularly sold in themarkets (Mendelsshon and Yom-Tov,1999).Another four species of squirrels havebeen introduced, at least since the 19th-20th centuries, in several areas of theMediterranean. These are as follows.

2.4 Grey squirrel, Sciurus carolinensisGmelin, 1788

The original distribution of this nearcticspecies was restricted to eastern NorthAmerica, where it ranged from the Gulfof Mexico north to the Great Lakes andthe St Lawrence river, and west to theprairies (Lever, 1994). It has been intro-


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duced in Australia (where it is nowextinct), South Africa, Great Britain,Ireland and Italy (Lever, 1985; Gurnell,1987). Records of the first importationsto Europe are referred to those carriedout from USA and Canada to GreatBritain several times with transloca-tions between 1876 and 1929 (Lever,1994; Gurnell and Wauters, 1999a). In Italy the species was introduced forthe first time in the site of Candiolo(Torino) in 1948 (Currado et al., 1987).From here the grey squirrel becameprogressively diffused until it occupieda range of several hundred square kilo-metres, and it is now present with apopulation of several thousand individ-uals (Bertolino et al., 2000). The expan-sion recorded in recent years has ledthis population to colonise certain areasof the Alpine foothills, and at present aprogressive expansion along the AlpineArc is considered highly probable(Genovesi and Amori, 1999; Genovesiand Bertolino, 2000). The secondItalian nucleus has developed aroundGenova Nervi following the release offive specimens which took place in1966; the population currently presentis estimated at around several hundredindividuals distributed over a rangeextending between Nervi andBogliacco (Currado et al., 1997;Bertolino et al., 2000). Finally, the thirdnucleus is of more recent origin, havingbeen created through the release ofthree pairs in Trecate (Novara) in 1993;reports from some protected areaslocated within the administrativeboundaries of the regions of Piedmontand Lombardy lead us to believe thatthis population too is spreading alongthe valley of the river Ticino (Currado

et al., 1997; Bertolino et al., 2000).

2.5 Pallas’s squirrel, Callosciurus ery-thraeus (Pallas, 1778)

The only free-living population knownin Europe is that introduced to Capd’Antibes (Alpes-Maritimes, France) inthe early 1970s (Jouanin, 1986). Theoriginal distribution of this speciesranges from Sikkim, Bhutan and Assameast through Burma to S. China, includ-ing Sichuan north of the Yangtze, andsouth to Indochina, Thailand andMalaya, as well as on Taiwan andHainan (Corbet and Hill, 1992). It ishowever absent from much of centralThailand occupied by the co-generic C.finlaysonii (Horsfield, 1824).The squirrels of Cap d’Antibes seem tobelong to the subspecies C. e. erythro-gaster Blyth, 1843, from the areas ofAssam and Myamar (Burma) betweenthe rivers of Brahmaputra andChindwin (Gurnell and Wauters,1999c).

2.6 Variable squirrel or Finlayson’ssquirrel or Thailand tree squirrel,Callosciurus finlaysonii (Horsfield,1824)

The homeland of this species comprisesThailand, Cambodia and Laos, perhapswith an isolated population in Burma,east of the lower Irrawaddy (Corbetand Hill, 1992). Until a few years ago,the only known European population ofthe taxon was reputed to be that ofAcqui Terme (Alessandria), in north-western Italy (Currado et al., 1999;Bertolino et al., 2000; Andreotti et al.,2001). This was originated by the

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release of two pairs of animals into anurban park in the early 1980s (Bertolinoet al., 2000). Recently, however, thepresence of the species was recorded onthe Tyrrhenian coast of Basilicata,where the current population is appar-ently derived from the release of sever-al specimens by private citizens in thedistric of Maratea, again in the 1980s(Aloise et al., 2003; G. Aloise and M.Cagnin, in verbis) (Fig. 4).

2.7 Siberian chipmunk, Tamia sibiricus(Laxmann, 1769)

This is a taxon characteristic of thenorthern Palaearctic, where its naturaldistribution ranges from northernRussia across Asia as far as Siberia andChina and the islands of Sakhalin,Hokkaido and S. Kurile (Corbet,1978). It has been introduced by man towestern Europe, where isolated popula-tions are today present in Belgium,Germany, The Netherlands, France,Switzerland and Italy (Amori, 1999;Andreotti et al., 2001). In Italy the pop-ulation which has established itselfalong the Piave in the vicinity ofBelluno originated in 1969-1970, fol-lowing the escape of several individualsfrom a pet shop, expanding subsequent-ly within a range of at least 3 km fromthe point of the escape (Dal Farra et al.,1996). The chipmunk also proves to berecorded in certain districts ofPiedmont, Liguria, Friuli-VeneziaGiulia, Trentino and Lazio (Amori andGippoliti, 1995; Dal Farra et al., 1996).Several unsuccessful attempts at releasein urban and suburban parks and gar-dens took place in the course of the1970s in Tuscany, in the Florence-Pratoarea.

In the course of the last decade, thespecie has been sporadically reported inLombardy, for the park of the valley ofTicino (B. Valenti, pers. com.).

3. PRESENT NATURAL AND ANTHRO-POCHOROUS DISTRIBUTION OF DORMICE IN

THE MEDITERRANEAN REGION

The Gliridae are a monophyletic familythat is represented by only eight generaand some thirteen or fourteen species inthe extant fauna (Daam and de Bruijn,1994). In the Mediterranean Regionthis group comprises the following 7species.

3.1 Edible dormouse, Glis glis (L.,1766)

Also known as fat dormouse or squir-rel-tailed dormouse, it is dispersed inEurope, including several Mediter-ranean islands (in most of which it hasbeen introduced by man in ancienttimes), northern Asia Minor, theCaucasus and north-western Iran(Storch, 1978). Outside its natural con-tinental distribution it is naturalised inEngland (Lever, 1994; Yalden, 1999).

3.2 Common dormouse or hazel dor-mouse, Muscardinus avellanarius (L.,1758)

It belongs to a monospecific genusclosely allied to Glis, characterised by awestern Palaearctic distribution whichcomprises most of Europe – apart fromsouth-western France and the Iberianpeninsula – and northern Anatolia; asfar as it is presently known, theMediterranean island populationincludes only Sicily and Corfu (Greece)(Morris, 1999).


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3.3 Asian garden dormouse or black-tailed dormouse, Eliomys melanurusWagner, 1840

It occurs in North Africa from Moroccoto Egypt, western Arabia and theLevant (Harrison and Bates, 1991;

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Figure 4 - Present distribution of the variable squirrel in Italy.

Mendelsshon and Yom-Tov, 1999).

3.4 Garden dormouse, Elyomis querci-nus (L., 1766)

This species is endemic to Europewhere it is found from the westernIberian peninsula to the Urals; thespecies is today absent from the BritishIsles, where it was instead recorded insettlements of Roman age, probably asa result of human introduction(Filippucci, 1999). It is present, and hasbeen recently recorded, on severalwestern Mediterranean islands, such asMenorca, Mallorca, Formentera,Corsica, Sardinia, Capraia (Tuscanarchipelago), Sicily, Lipari (Aeolianarchipelago), and on the Croatian

islands of Krk, Bra�, Hvar, ��edro,Kor�ula, and Lastovo (Corbet, 1978;Alcover, 1988; Filippucci, 1999; Amoriand Masseti, 1996).

3.5 Forest dormouse, Dryomis nitedula(Pallas, 1778)

It is distributed from easternSwitzerland and north-eastern Italy,across eastern Europe and Asia Minorto the east as far as Mongolia and theTien Shan mountains in China(Ondrias, 1966; Corbet and Hill, 1991;Harrison and Bates, 1991). There aresome isolated populations in southernItaly (Basilicata and Calabria)(Lehmann, 1964; Toschi, 1965; Paoluc-ci et al., 1987; Cagnin and Aloise,1994). While Kry�tufek (1999) record-ed this species as not occurring onEuropean islands, Ondrias (1966) andCheylan (1988) include it among thenon-flying terrestrial mammals of

Euboea. The occurrence of this specieson the Cycladic island of Andros,recorded by Chondropoulos andFrageudakis-Tsolis (in verbis), is still tobe confirmed.

3.6 Woolly dormouse, Dryomis lanigerFelten and Storch, 1968.

Its range is restricted to south-westernAsia Minor, from the Taurus Mts(Felten et al., 1973) to eastern Anatolia(Mursaglolu, 1973; Kry�tufek andVohralik, 2001).

3.7Mouse-tailed dormouse, Myomimusroachi (Bate, 1937).

It is distributed in the easternmostBalkan peninsula (Thrace and south-east Bulgaria) and Asia Minor(Kurtonur and Özkan, 1991; Filippucciand Peshev, 1999).

In considering the diffusion of severalof these species of glirids in theMediterranean region, and in particularon the islands, it is useful to cast furtherlight upon the way in which the erro-neous evaluation, or rather the inatten-tive reading, of the pages of authors ofthe past has given rise to cultural mod-els which are still difficult to eradicate.Let us take, for example, the supposedformer occurrence of D. nitedula.Erhard (1858) reported the diffusion ofa species of glirid, similar in name tothe forest dormouse, which he calledMyoxus nitela Schreber, 1782, on theCycladic islands of Andros, Naxos andSiphnos, where it occurred in orchardsand orange groves; it was in factbelieved that it fed on citrus fruits. This


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report gave rise to the belief in the dif-fusion of this rodent on these islands,but in reality the taxonomic classifica-tion does not correspond to that of theforest dormouse. In fact, in Ellermanand Morrison-Scott (1951) Myoxusnitela Schreber, 1782, is indicated asone of the synonyms of Eliomys querci-nus (L., 1766), that is of the gardendormouse, a species currently unknownin the Aegean area, being widespread,as already observed, in the central-western Mediterranean basin, where itis not found further east than Dalmatiaand the north-western Balkan peninsu-la. As already observed, althoughaccording to Kry�tufek (1999), D. nit-edula does not occur on Mediterraneanislands, Ondrias (1966) and Cheylan(1988) reported it from Euboea.Recently the presence of the forest dor-mouse has also been recorded from theisland of Andros (Chondropoulos andFraguedakis-Tsolis, in verbis), althoughthis has yet to be confirmed. We have,on the other hand, known for some timeof the presence of the edible dormouseon islands such as Crete (Zimmermann,1953; Kahmann, 1959; Niethammerand Krapp, 1978), Euboea (Ondrias(1966), Corfu (Niethammer, 1962;Niethammer and Krapp, 1978), andCephallonia (Niethammer and Krapp,1978; Giagia-Athanassopoulou, 1998).On Cephallonia its occurrence has beenrecently confirmed by H. Pieper (in lit-teris), whereas Dimaki (1999) providedelements for the existence of thespecies on Andros. According to H.Alivitzatos and A. Lane (in verbis), theedible dormouse is also present on theisland of Thassos, where some evidencefor the occurrence of the species was

found in the surroundings of the villageof Panaghia, on August 2000. Anotherreport of the occurrence of a type ofdormouse, possibly the forest dor-mouse, is recorded by Wettstein (1942)from the island of Rhodes where, how-ever, according to other authors thespecies was, and still is, unknown (cf.Festa, 1914; De Beaux, 1929; Zimmer-mann, 1953).

4. THE FOREST DORMICE OF PYRGY CEN-TRAL ITALY (6TH-5TH CENTURY B.C.)

Let’s now focus on the extant distribu-tion of the forest dormouse in continen-tal Italy. As we have already seen, thisspecies is characterised by a very pecu-liar distribution, markedly fragmentedand divided into two ranges, one innorth-eastern Italy which represents thesouth-western border of the continuousEuropean range (Paolucci et al., 1987;Nappi, 2001) and the other with isolat-ed nuclei located very far to the south ofthe peninsula, in Basilicata andCalabria (Lehmann, 1964; Toschi,1965; Filippucci at al., 1985; Cagninand Aloise, 1994; Capizzi and Santini,2002).However, the indication of a muchmore extensive distribution of thespecies within the Italian peninsula inthe past has been provided by therecovery of certain remains in the fossillayers of the Upper Pleistocene of theGrotta Breuil, on Monte Circeo(Latina) (Kotsakis, 1990-1991;Alhaique et al., 1995), dating to the lastglacial episode. Referring instead tomuch more recent chronologies is thesingular discovery of D. nitedula in awell of the Etruscan sanctuary of Pyrgy,

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near Santa Severa in Upper Lazio(Caloi and Palombo, 1980). Thechronological span of this material hasbeen comprised within the first half of

the 6th century BC and the first half ofthe 5th century BC. (Colonna, 1970)

(Fig. 5). More than being faced with theevidence of a broader natural distribu-tion of the species in the Italian envi-ronment, in this case what we could

actually be dealing with is evidence ofan alteration of the range attributable to


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Figure 5 - Former (dots) and present (grey) distribution of the forest dormouse in Italy.

human action. As has been extensivelydocumented by the investigation car-ried out by André (1981), Carpanetoand Cristaldi (1994) and Colonnelli etal. (2000), in the Mediterranean regiona remarkable human impact on the geo-graphical distribution of some dor-mouse species and their populationdensity can be documented since antiq-uity through historical and biogeo-graphical analyses, supported by pale-ontological and archaeozoological data.Furthermore, ethnozoological investi-gations document the utilisation ofdormice for food or medicine in severalareas of the Italian territory, throughtraditional captive-breeding techniques,up to very recent historical times. Onseveral Italian islands, for example, theextant occurrence of some of theserodents, such as the edible dormouse onElba, Corsica, Sardinia and Salina(Aeolian archipelago), and/or the gar-den dormouse on Corsica, Sardinia,perhaps Capraia (Tuscan archipelago),and Lipari (Aeolian islands) (cfr.:Cristaldi and Amori, 1982; Amori andMasseti, 1996), might be interpreted asthe sole objective evidence that has sur-vived of their ancient artificial diffusion(Masseti, 2002). In the specific case ofthe island of Capri, Gulf of Sorrento(south-western Italy), findings of gar-den dormice in a cave on the easternslopes of Monte Solaro, within anarchaeological context characterised byfaunal remains of recent age, raiseintriguing questions about the chronol-ogy of the appearance of this rodent onthe island. This appearance has, in fact,been considered as contemporary withthe introduction of the black rat, Rattusrattus (L., 1758), onto the island

(Barbera and Cimmino, 1990), which isbelieved to have occurred on Capri notbefore the 5th century AD. (Albarella,1992). The edibility of dormice was oneof the major preoccupations of theLatin authors of the Classical Age, suchas Varro (Res rustica, 3.15. 1-2), Plinythe Elder (Naturalis historia, 8.224),Apicius (9.1.1), and Petronius (31.10).According to King (2002), it seemshowever that small glirids, such as thecommon and the garden dormice, werenot eaten, unlike Glis glis. In any case,ancient sources were imprecise aboutdormice and could be referring to anyof the common species, although theedible dormouse probably attracted themost attention by virtue of its largersize and its consumption as a delicacy.

5. CONCLUDING REMARKS

Throughout most of the 19th and 20thcenturies, it was common practice forthe scientific explorers to bring homean excessive number of subspeciesfrom their explorations of theMediterranean islands. In fact, previousauthors often classified many of themodern non-flying terrestrial mammalsof these territories as subspecific and/orgeographic forms, almost entirely onthe basis of arbitrary criteria and theexamination of scattered materials.Based on the data given in literature,the various subspecies are distinguishedby the coat patterns, and by the size ofbody and skull. This leads us naturallyto reconsider the great taxonomic pro-liferation to which the various popula-tions of the Sciuridae and Gliridae ofthe Mediterranean islands have beensubject, for the purposes of their classi-

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fication at subspecies level. Regardingthis, we ought to consider the taxonom-ic treatment, on the part of the interna-tional scientific community, of the edi-ble dormice of the Tyrrhenian andAegean islands (Glis glis insularisBarrett-Hamilton, 1898; G. glis meloniiThomas, 1907; G. glis argenteusZimmermann, 1953), as well as the gar-den dormice of the Balearics, Corsica,Sardinia, and Lipari (Eliomys quercinusgymnesicus Thomas, 1903; E. querci-nus ophiusae Thomas, 1925; E. querci-nus sardus Barrett-Hamilton, 1901; orE. quercinus liparensis Kahmann,1960) (Tab. 1). But, in the majority ofcases, these populations originated

from continental populations intro-duced by man, since the diffusion oftheir ancestors is not documentedamong the insular Quaternary faunalhorizons. As is consequently under-standable, this led to a multiplication offorms which now, however, demandbetter taxonomic and genetic definition. The examination of fossil materialshelps to indicate the previous distribu-

tion of the different species on the basisof the location of the osteologicaldeposits which contained their remains.However, to assess the original range ofthe species, earlier chronologies prior tothe Neolithisation (end of 7th-6th mil-lennium B.C.) of the Italian territories,for instance, should be considered, afterwhich improved human sea-faringskills and the established commercialnetworks between the Mediterraneancountries enabled the artificial exporta-tion of faunistic species of culturalinterest, together with those alreadyinvolved in the process of domestica-tion (Masseti 1998; Lorenzini et al.,2002). Looking again at the former dis-

tribution of the squirrels and dormice insouthern Italy, and in particular in theterritories within the extant politicalboundaries of the regions of Basilicataand Calabria, it is almost impossible tofind, for example, information aboutthe original diffusion of D. nitedula.Data are in fact only available for theoccurrence of the red squirrel(Bulgarelli, 1972), the edible dormouse


17

Taxon Island References

Sciurus vulgaris lilaeus Greece and Skyros island Miller, 1907Glis glis insularis E Sicily Barrett-Hamilton, 1898Glis glis melonii E Corsica and Sardinia Thomas, 1907Glis glis argenteus E Crete Zimmermann, 1953Muscardinus avellanarius pulcher Southern Italy and Sicily Barrett-Hamilton, 1898Muscardinus avellanarius zeus Greece and Corfù Chaworth-Musters, 1932Eliomys quercinus gymnesicus E Mallorca and Menorca Thomas, 1903Eliomys quercinus ophiusae E Formentera (Balearics) Thomas, 1925Eliomys quercinus sardus E Corsica and Sardinia Barrett-Hamilton, 1901Eliomys quercinus liparensis E Lipari (Aeolian islands) Kahmann, 1960

Table 1 - Subspecies of the present representatives of the families Sciuridae and Gliridae(Mammalia, Rodentia) described for the Mediterranean islands (E = endemic subspecies).

(Topa, 1933; Bulgarelli, 1972) and thecommon dormouse (Bulgarelli, 1972)(Tab. 2; Fig. 6). The forest dormousedoes not appear to be represented

among the Pleistocene fossil findings ofthese territories, giving rise to intrigu-ing questions about its origin.Given the lack of information about itsdistribution in southern Italy, in thepresent state of knowledge is it possiblethat this species was introduced forfood or for other cultural purposes inancient times? Effectively we ought tobear in mind the existence of consider-able information regarding the translo-cation of different zoological taxa thatwas performed by man between north-ern and southern Italy in antiquity, suchas, for example, the hypothesised intro-duction of the subspecies of asp viper,Vipera aspis hugyi Schinz, 1833, char-acteristic of southern Italy and Sicily,onto the island of Montecristo in theNorthern Tyrrhenian sea (Bruno, 1985;Zuffi and Bonnet, 1999; Zuffi, 2001). Or is the absence of the forest dor-mouse from the fossil horizons ofsouthern Italy instead to be directlyrelated to the evidence of palaeontolog-ical sampling biases? The invitation isto further study which, if conducted in

the correct manner, integrating theinformation to be derived from thepalaeontological, archeozoological,neontological and genetic ambits, will

be bound to make a further illuminatingcontribution to a better understandingof the origin of D. nitedula in southernItaly, within the more general frame-work of the modern definition ofMediterranean mammal fauna.

ACKNOWLEDGEMENTS

I would like to express my appreciationand gratitude to the following friendsand colleagues for their suggestions andassistance as I was preparing this paper:Umberto Albarella, Department ofArchaeology of the University ofDurham; Haralambos Alivitzatos andA. Lane, Athens; Gaetano Aloise andMara Cagnin, Dipartimento di Ecologiadell’Università della Calabria, Arcava-cata di Rende (Cosenza); Vassilis P.Chondropoulos and Stella Fraguedakis-Tsolis, Section of Animal Biology ofthe University of Patras; Maria Dimaki,The Goulandris Natural HistoryMuseum, Athens; Piero Genovesi,Istituto Nazionale per la FaunaSelvatica, Ozzano dell’Emilia (Bolo-

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Taxon Palaeontological site Age References

Sciurus vulgaris Torre Nave, Praia a Mare (CS) MUW Bulgarelli, 1972Glis glis Grotte di Cirella, Diamante (CS) LW Topa, 1933Glis glis Torre Nave, Praia a Mare (CS) MUW Bulgarelli, 1972M. avellanarius Torre Nave, Praia a Mare (CS) MUW Bulgarelli, 1972

Table 2 - Fossil remains of squirrels and dormice provided by the exploration of someWürmian sites in Calabria region (southern Italy); MUW = Middle and Upper Würm, LW =Lower Würm

gna); Armando Nappi, AssociazioneRicerca e Conservazione Ambientale,Napoli; Laura Pardi, SoprintendenzaArcheologica della Toscana; Harald

Pieper, Zoologisches Museum Kiel,and Marco A.L. Zuffi, Museum NaturalHistory and Territory of the Universityof Pisa.


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Figure 6 - Geographical location of the prehistoric sites of Calabria (southern Italy) that pro-vided Würmian remains of squirrels and dormice.

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Masseti

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Disegno di Laura Romagnoli

dormice of the mediterranean region naturaland ... riassunto – scoiattoli e ... (waldren, 1982;...

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