The crested porcupine, Hystrix cristata L., 1758, in Italy
Marco MASSETI Laboratori di Antropologia eEtnologia Dipartimento di Biologia Evoluzionistica dell’Università di Firenze Viadel Proconsolo 12 50122 Firenze (Italy) [email protected]
Umberto ALBARELLA Department of Archaeology University of Sheffield, Northgate House, West Street Sheffield S1 4ET (United Kingdom) [email protected]
Jacopo DE GRoSSI MAZZoRIN Università del Salento, Dip. Beni Culturali via D. Birago, 64 73100 Lecce (Italy) [email protected]
Masseti M., Albarella U. & De Grossi Mazzorin J. 2010. — The crested porcupine, Hystrix cristata L., 1758, in Italy. Anthropozoologica 45 (2): 27-42.
ABsTRACT In this paper the evidence for the introduction of the crested porcupine (Hystrix cristata L. 1758) in Italy is reviewed and hypotheses concerning the timing and modalities of this event arebrought forward. The crested porcupine current distribution outside Africa is limited to Sicily and the Italian Peninsula. Palaeontological data indicate that porcupines were present in Italy and other parts of Europe in the Pleistocene (possibly up to the early Holocene), but there is now broad consensus that these belonged to different species, now extinct. Apart from some unreliable prehistoric finds, there is no KEYWORDs evidence that porcupines occurred again in Italy and the rest of Europe before Porcupine, historic times. This led many authors to suggest that the Romans may have Italy, Sicily, been responsible for the introduction of the species in Italy,but such fossils, assumption was largely speculative. The available evidence in fact indicates zooarchaeology, that the crested porcupine does not occur in Italy before late Antique or even introduction, Romans, early medieval times and that it never spread in Europe beyond the Italian late Antiquity. Peninsula (and Sicily). The long chronological gap existing between
ANTHROPOZOOLOGICA • 2010 • 45 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 27 Masseti M., Albarella U. & De Grossi Mazzorin J.
Pleistocene and late Holocene specimens strongly suggests that the presence of the species in historic times in Italy is the result of ahuman-induced introduction. Variations in the morphometric and genetic characteristics of currentporcupine populations in Italy and differences in the timing of the introduction between different regions suggest that the species was probably introduced as a consequence of multiple events.
RésUMé Le porc-épic à crête, Hystrix cristata L., 1758, en Italie Cet article met en évidence l’introduction du porc-épic à crête (Hystrix cristata L., 1758) en Italie et avance des hypothèses au sujet de la chronologie et des modalités de cette insertion. La répartition actuelle du porc-épic à crête en dehors de l’Afrique est limitée à la Sicile et à la Péninsule italienne. Les donnéespaléontologiques indiquent que les porcs-épics étaient présents en l’Italie et dans d’autres parties de l’Europe au Pléistocène (probablement jusqu’à l’Holocène récent), mais tout le monde s’accorde à dire que ces espèces sont, à présent, éteintes. Mises à part quelques découvertes préhistoriques non authentifiées, il n’existe aucune preuve que les porcs-épics furent présents en l’Italie et dans le reste de l’Europe avant les périodes historiques. Ceci a amené de nombreux d’auteurs à suggérerque les Romains puissent avoir été responsables de l’introduction de ces espèces en Italie, mais cela reste largement spéculatif. En fait, les éléments à notre disposition indiquent que le porc-épic à crête n’est pas présent en Italie avant l’Antiquité tardive voire au début de la période médiévale et qu’il ne s’est jamais étendu en Europe au- delà de la Péninsule italienne (et de la Sicile). Le grand espace chronologique MOTs CLés existant entreles spécimens du Pléistocène et de l’Holocène tardif suggère Porc-épic, fortement que la présence de l’espèce durant les périodes historiques en Italie Italie, a été le résultat d’une introduction humaine volontaire. Les variations des Sicile, fossiles, caractéristiques morphométriques et génétiques des populations actuelles de archéozoologie, porcs-épics en Italie et une chronologie différente de l’introduction de l’espèce introduction d’espèce, Romains, selon régions incitent à penser que celle-ci a probablement été introduite à la Antiquité tardive. suite d’événements multiples.
CURRENT DISTRIBUTION Masseti 2008, 2009a) (Fig. 1).Thislast occurrence is the result of very recent introductions, probably The crested porcupine Hystrix cristata L., 1758, is dating to the first half of the 1980s (De Marinis currently present as awild form in Italy,but its et al. 1996). Despite what has been suggested by statusasagenuinerepresentativeofthe native several authors (Wettstein 1942; Brink 1969; European fauna is questionable and requires inves- Corbet 1978;Honacki et al. 1982;Burton & tigation. This rodent currently occurs in most of Pearson 1987; Cheylan 1988), the species has never northern Africa and in Sub-Saharan Africa from occurred in the Ionian and Aegean islands, the Senegal to Ethiopia, up to northern Tanzania Balkans or the Iberian Peninsula in the Holocene, (Cabrera 1932; Corbet &Jones 1965; Ranck 1968; even in the very recent past. This erroneous sug- Smit &Wijngaarden 1981; Niethammer 1982). gestion has, however, become embedded in the Its European distribution is limited to peninsular literature to thepoint thatevenrecently Gisbert Italy,Sicily and the island of Elba (Lovari 1993; et al. (2002) and Woods and Kilpatrick (2005) have
28 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
suggested aEuropean distribution of the crested porcupine that extends beyond Italy,toAlbania and northern Greece. Wettstein (1942) observed that local people referred to the presence of porcu- pines also on the Eastern Aegean islands of Ikaria and Lesbos, but he added that this might havebeen the result of confusion with the hedgehog. The Greek term used to indicate this latter species is skanzohiros, which means ‘spiny pig’, which might easily havegiven rise to the confusion with the English ‘porcupine’(the Italian “porcospino” and/or the French “porc-épic”). All these reports have given rise to several unproven beliefs which still persist. Cheylan (1988), for example, still erroneously quotes theoccurrence of Hystrix cristata on the Eastern Aegean islands of Lesbos, Ikaria and
Rhodes, and Sarà (1998) that of the same species fig.1.—Current distribution of the crested porcupine Hystrix on Samos and Rhodes. cristata Linnaeus, 1758. Modified from the IUCN Red List of threatened species (http://www.iucnredlist.org/apps/redlist/ It is also worth mentioning that the crested porcu- details/10746/0) (prepared by Angela Trentacoste). pine currently occurs also off the north-western Libyan coast, on the island of Farwa(Bashir 2009). In addition, the Indian crested porcupine, H. indica further remarked that the limited geographic dis- Kerr,1792, is found on the Tu rkish islet of Domuz, tribution of the species in Europe could be the in the gulf of Fethiye, as partofafauna almost result of an artificial introduction. Brehm (1916) entirely comprised of anthropochorous species. The also thought that the origins of this rodent had to latter is the species of porcupine occurring in Asia be found in NorthAfrica, and that the Romans Minor and the Middle East (cf. Corbet &Jones 1965; were responsible for its introduction into Italy.This Woods & Kilpatrick 2005). Since porcupines are suggestion eventually becamecommonplace in the not competent swimmers, it is fairly certain that literature, but was never fully explained or indeed they were brought to these islands by people, which disproved,and many laterauthors,suchas indicatesapropensity by humans to transportthese Ghigi (1959), Niethammer (1963, 1982), Corbet animals. An additional consideration is that both &Jones (1965), Toschi (1965), Brink (1969), islands occur within the boundaries of Islamic Kingdom (1974), Pratesi and Tassi (1974), countries, which raises the possibility of a special Corbet (1978) and Santini (1980) havedogmati- interest in the species by people with these cultural cally accepted it. Such widespread consensus is and religious backgrounds. surprising if we consider that nobody so far has been able to indicate the time and place of such hypo- thetical introduction, or attribute it to specific people STATEOFTHE ART or events (Masseti 2008). Orsomando and Pedrotti (1976) remarked at the time that there The suggestion that the porcupine was introduced was neither palaeontological or archaeozoological into Italy from Barbary(i.e. NorthAfrica) had evidence that allowed us to suggest with any cer- already been raised by the late 18th C(Minà Palumbo tainty an introduction of the crested porcupine in 1868). At about the same time Cornalia (1871-72) Italy in Roman times. In this respect it may be also claimed that the animal had been introduced worth pointing out the absence of the porcupine from Africa, and he also specified that the species in both the iconography and the archaeology of had not expanded beyond the regions of Calabria Pompeii, despite the attested occurrence of a large and Abruzzo. Early in the 20th C, Miller (1912) variety of animal species, many of them exotic
ANTHROPOZOOLOGICA • 2010 • 45 (2) 29 Masseti M., Albarella U. & De Grossi Mazzorin J.
(see King 2002). In addition, in his Naturalis certainly based on a misunderstanding of the pal- Historia (VIII, 125), Pliny the Elder claims that aeontological evidence, as thereare no reliable “Hystrices generat India et Africa” [porcupines occur European records of porcupines between the early in India and Africa], which indicates that he Holocene and early historic times (Riquelme Cantal had no knowledge of their existence in Italy. The &Morales Muñiz 1997). As Kurtèn(1968) had hypothesis that the porcupine may havebeen intro- already remarked at the time, the crested porcupine duced by the Phoenicians (as claimed by Tr ucchi currently living in Italy is unlikely to be at all related & Sbordoni 2007) is also completely unfounded. with the Pleistocene animals. To provide support This people have often — but arbitrarily — been for this hypothesis there is the consideration that made responsible for the movement of several almost all findings of Pleistocene porcupines from other animal species outside their natural distri- continental Italy and Sicily haverecently been bution (Masseti 1996, 2008). regarded as contamination from upper layers or re-identifiedasdifferent speciesofthe genus Hystrix, such as H. refossa Gervais, 1852, and/or H. vinogra- ARCHAEOLOGICAL dovi Argyropulo, 1941 (Kotsakis et al. 2003; AND PALAEONTOLOGICAL EVIDENCE – Bonfiglio et al. 2004; Salari &Sardella 2008, 2009; PREHISTORY Masseti 2008). In previous years Rustioni et al. (1999) had already identified a porcupine right mandible According to Riquelme Cantal and Morales from Monte Tignoso (Northern Apennines) as a Muñiz (1997), thereisnooccurrence of the crested likely H. vinigradovi. H. refossa is a larger species porcupine in Europe beforehistoric times, and even and has been attested in Europe since the Upper then, no evidence of its presence outside Italy, but Pliocene, but, like H. vinogradovi, was still present they are, of course, referring to the Holocene. Sev- in the Pleistocene (Weers 1994, 2005). eral archaeological sites, dated to the Middle and Upper Pleistocene and ranging from the Lessini Mountains in north-eastern Italy to Sicily,have THE SICILIAN CASE attested the occurrence of porcupines in the past of Italy.Bartolomei (1969, 1980) regards the genus As in the rest of Italy, in Sicily the genus Hystrix Hystrix to be characteristic of the Interglacial was present in the Upper Pleistocene but seems to periods in central Europe. Palaeontological data disappear in the early Holocene (Kotsakis 1979; from continental Italy indicate the occurrence of Burgio &DiPatti 1990; Burgio 1997; Burgio et al. these rodents in the Upper Pleistocene, though 2002, 2005). In Eastern Sicily porcupine remains no further chronologically definition could be havebeen found at the SanTeodoroCave(Acquedolci, provided (Esu &Kotsakis 1986). An odd discovery Messina) (Anca 1860a, 1860b). More findings is represented by the occurrence of animal bones derivefromthe early Holocene of Grotticella di that arethought to havebeen gnawed by porcu- Porto Palo (Villari 1995), and these probably rep- pines at the Neolithic site of Grotta Zachito in resent the latest known occurrence of a representa- Campania (Regalia 1903). In viewofthe identi- tive of this genus in Sicily before historic times. Of fication of other unlikely species, such as camel interest is the absence of porcupine remains from and rat, at this site, this finding must, however, the Upper Pleistocene and early Holocene levels of be treated sceptically. Uzzo Cave(Tagliacozzo1993), one of the most Amori and Angelici (1992, 1999) have suggested carefully investigated animal remain assemblages that in the Upper Pleistocene H. cristata occurred from Sicily.Ifweaccept that the genus Hystrix in the Central and Eastern Mediterranean (Italy became extinct on the island in the early Holocene, and Balkans) but periodically expanded into Central then it follows that the current occurrence of the Europe. The same authors believe that the current crested porcupine must be aconsequence of an absence of the species in the Balkans may be the introduction (Burgio et al. 2005), almost certainly result of arecent extinction. This is, however, almost caused by humans.
30 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
The analysis of the documentaryevidence may help in clarifyingthisissue. According to thePersian traveller Zakarîâibn Muhammad ibn Mahmûd, al-Qazwînî, in 13th C Sicily there were no animals that “… attacked humans using teeth, claws or quills” (cf. Ruta 2007). It is only centuries later, in 1743, that the erudite Antonino Mongitore(1663-1743) writes that in the Sicily of his times therewerethree different kinds of swines: “[…] domestici, che si nudriscono nella città e nelle campagne, intiere man- drie, per vendersi in nudrimento degli abitatori; selvaggi, detti cignali: e questi sono diletto dei caccia- tori, de’ quali fa menzione Fazello; e spinosi, armati di spine, che scagliano controquei, che intendono d’accostarsi per molestarli, ed ucciderli […]” [domes- tic, whose herds feed in town and country, and are then sold for food; wild, also known as wild boar: theseare theentertainment of thehunters men- tioned by Fazello; and prickly, which have quills as fig.2.—Early 16th C tile from Faenza (Ravenna, north-central weapons, which they throw against those who mean Italy), with the representation of a crested porcupine (Florence, to bother or kill them […]], finally making a clear private collection). reference to the presence of the porcupine. The occurrence of the crested porcupine in Sicily was workofAgricola, published in 1614, the introduc- later attested by other authors, such as Ghisio (1779) tion of the porcupine into Europe is suggested to and Ortolani and Rafinesque Schmaltz (1810). have occurred at the time thebookwas written (Cabrera 1932). According to Agricola, therefore, the introduction of the species represented a rela- DOCUMENTARY tively late phenomenon, probably of the 16th C or AND ICONOGRAPHIC EVIDENCE only slightly earlier. This view was later accepted by several other authors (e.g.Buffon 1766; Well beforeMinà Palumbo(1868), Cornalia Lacepède 1830; Costa 1839; Minà Palumbo 1868; (1871-72), Miller (1912) and Brehm (1916), other Cornalia 1871-1872) who also agreed that the authors had suggested that the crested porcupine porcupine was an alien species to the European was not a native species in Europe. Among these, fauna, thus contributing to the current perception the first modern author wasGiorgius Agricola of the species. The porcupine is also known from (1490-1555) — in the world Georg Bauer — who, the Italian iconography of the early 16th C(Fig. 2). following up from the passage by Pliny the Elder As acaptiveanimal, the crested porcupine was, already referred to, remarked in his De Animantibus according to literaryaccounts, kept in England, Subterraneis [OnUndeground Animals] published in the menagerie of HenryI(1068-1135) at in Basel in 1548, that: “L’ India el’Africa produce Woodstock (Plot 1705; Rybot 1972; Ververs 1976; quest’animale, epoco ha che franoi ne fu portato uno” Landsberg 1998), whereitcould havebeen sup- [India and Africa havegot this animal, and recently plied by William of Montepellier (Hahn 2003). one has been brought to us]. It is noteworthy that In France, knowledge of the species is demon- this 16th Cauthor,inline with the scientific knowl- strated by the occurrence of its image in the edge of the time, did not make any distinction device of Louis of Orléans (1372-1407), which between the species H. cristata, living in Italy and was a few decades later again used by his grandson, most of Africa, and the already mentioned Indian the king Louis XII (1462-1515) (see Hochner 2001) crested porcupine. In a posthumous edition of the (Fig. 3).
ANTHROPOZOOLOGICA • 2010 • 45 (2) 31 Masseti M., Albarella U. & De Grossi Mazzorin J.
(1590 - after 1646), which have as their subject a crested porcupine and were produced for the 16th C‘paper museum’ofCassiano dal Pozzo (Masseti 2008). Currently one of these pictures is kept at the Royal LibraryofWindsor Castle, England, while the other belongs to a private col- lection (Freedberg 2002; Attenborough et al.2007). One picture portrays an adult specimen (Fig. 4), while in the other an ear, the paws, the snout (with aspecial attention to the representation of the inci- sors) and some quills areveryfinely represented (Fig. 5). It is likely that the reproduction of these details did not happen by chance but it can rather be explained by the interest that the animal must fig.3.—The emblem of the porcupine inherited by Louis XII, king of France (1462-1515), from his grandfather,Louis of Orleans haverisen in the intellectual circles of the time. (Florence, private collection). They were clearly influenced by the careful descrip- tion of the crested porcupine that had been provided by Agricola (1548): “… horadirò de’l’Histrice… To better understand the influence that the work AlcunideGreci lo chiamanoArathochiron, per ciò of Giorgius Agricola had on western thought, it is che egli ha somiglianza, e forma d’un porcellin di due worth pointing out the existence of two polychrome mesi, & è irsuto, e pien di spine come lo spinoso: la pictures, attributed to the painter VincenzoLeonardi sua testa però ha più somiglianza con la lepre, gl’orecchi
fig.4.—Representation of an adult specimen of crested porcupine attributed to the painter Vincenzo Leonardi (1590 – after 1646), produced for the 16th C “paper museum” of Cassiano dal Pozzo (London, collection Sven Gahlin, NRC CDP 60).
32 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
fig.5.—Drawing attributed to the painter Vincenzo Leonardi, produced for the “paper museum” of Cassiano dal Pozzo, portraying the ear, the snout with a special attention to the representation of the incisors, and some quills of H. cristata (Royal Collection, RL 19438 © 2009 HM Queen Elisabeth II).
ANTHROPOZOOLOGICA • 2010 • 45 (2) 33 Masseti M., Albarella U. & De Grossi Mazzorin J.
a quei de l’huomo, & i piedi a quei de l’orso […] Le ARCHAEOLOGICAL EVIDENCE – prime spine nascon dal manto del dosso,edalelatora, HISTORICAL PERIOD ma lunghissime l’ha dai lati, epiu di tutte lunghissime sonquelle chehadalaparte di sopradidettilati. The archaeological evidence confirms Agricola’s Ciascuna di dette spine in parte è nera, & in parte observations that the porcupine had already been bianca. Lunghe sono due otre,oquattropalmi, le introduced into Italy by the 16th C, and in fact quali spine quando gli piace, come il pavon la coda, points out to an earlier occurrence. Modern zooar- lieva in alto, e dovendo entrar ne la tana l’abbassa, chaeological research has documented the certain quando provato ad ira, egli si distende la cotenna, e presence of the species at Settefinestre (Grosseto) la tira ne la bocca de’ gl’abbaianti cani che l’attizano, (King et al.1985), Farnese (Viterbo) (Wilkens 1991; e non giungendo con esse i cani, con tanto impeto le Colonnelli &DeGrossi Mazzorin 2000) and tira, ch’elle si ficcano ne legni. Ha identi come la Formello (Rome) (Minniti 2005). In all three sites lepre, quattron’ha lunghi, due di sopra, edue di the earliest occurrence of the porcupine can be sotto…” [… now I will tell of the porcupine […] dated to aperiod broadly encompassing the some Greeks call it Arathochiron, for what it 15th and 16th C. At Settefinestreseveral remains resembles, it has the shape of a two month piglet, of the animal were found in destruction and it is hirsuteand full of quills:its head, however, re-occupation levels dated from the 15th Conwards, resembles more ahare, its ears ahuman, and its whereas at Farnese and Formello the bones of the feet a bear […]. The first quills come out from the porcupine occurred in urban wells used as dumps. back, but they are very long on the sides, particu- The species has also turned up in mid 16th to mid larly in the upper part. The quills areblack and 17th C levels of the urban site of San Salvo (Chieti, white. They are between two and four hands long, Abruzzo) (DeGrossi Mazzorin &DeVenuto 2006). and can be raised, like the feathers of the peacock Much earlier remains derive from the baptistery of tail, but when the animal enters its den they are Santa Cecilia (12th-13th CAD) (DeGrossi Mazzorin lowered. When it is threatened the porcupine can &Minniti2004)and, particularly,the CryptaBalbi raise its quills and throw them towards the mouth exedra (from both 7th and 8th CAD), both in Rome of barking dogs that chase it. If it misses its target (DeGrossi Mazzorin &Minniti 2001; Min- the quills arestill expelled with such energy that niti 2005). Outside Rome two medieval porcupine they thrust themselves in wood. Its teeth are like remains — one dated to the 9th-mid 13th C AD hare’s, four of them are long, two in the upper and and the other to 1350-1415 AD — were found at two in the lower jaw...]. The belief that porcupines Cencelle (Civitavecchia), in northern Latium could throw their quills can already be found in (Minniti 2009). It must also be mentioned that a the treatise Sulla Caccia [OnHunting] by Oppiano 7th CADfragment of distal humerus from the di Apamea (III, 391-406). This is an entirely fan- Crypta Balbi has clear cut marks (Fig. 6), indicating tastic belief,not at all supported by scientific obser- the exploitation of this species by humans, probably vations. Among others, Pseudo-Aristotle (623a), as food. Other porcupine remains derivefromthe Aristotle (De Natura animalium, IX, 39, 7), Pliny late Roman site of SanGiovanni di Ruoti in Basili- the Elder (Naturalis Historia,VIII, 125), Aelian cata (MacKinnon 2002) and these havebeen dated (De Nat. Anim., 31), Claudian (De hystrice,17) through cal C14 analysis to 560-720 AD (94,2% and Isidore of Seville (XII, 2, 35), all contributed probability) (Small 2005), and arethereforeapproxi- to this conviction, which is in fact still held in large mately contemporarywith the specimens from the parts of the rural world. This myth is also attested Crypta Balbi.Afew years ago we also identified in the popular English tradition of the 12th C AD. some teeth, cranial and postcranial bones (repre- William of Malmesbury(c.1080/1095-c.1143) senting at least one adult and one juvenile) in an describes the porcupine as a wild beast, which is: Iron Age context at Capanna Pelli (Cavallino, Lecce) “… hispidis setis coopertam, quas in Canes insectantes (Fig. 7) (a discovery mentioned by Riquelme Cantal naturaliter emittunt” [coveredinquills, which are & Morales Muñiz 1997). Subsequently, however, instinctively thrownagainst chasing dogs] (Plot 1705). these remains were dated to alater period,
34 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
according to acal C14 analysis which attributed them to 770-980 AD (95,4% probability,CeDAD Laboratory of the University of Salento) (Fig. 8). Their occurrence in an apparently earlier context at Capanna Pelli may be explained with the bur- rowing habits of this species. On the basis of the evidence illustrated above the remains of San Gio- vanni di Ruoti and Crypta Balbi thereforerepresent the earliest evidence of the post-glacial presence of the porcupine in Italy that is currently available (Masseti 2008). On this basis De Grossi Mazzorin and Minniti (2001) and Minniti (2005) con- cluded that the most likely hypothesis is that the species was introduced into Italy between Late Antiquity and the early Middle Ages. The evidence presented in this paper confirms such suggestion and also indicates an element of likely chronologi- cal continuity between the late Roman and late medieval animals (see Fig. 9for distribution map of sites mentioned in this section).
THE HISTORICAL CONTEXT
The introduction of the crested porcupine in this period makes sense when we look at the more gen- eral historical context.InLate Antiquity and for most of the early Middle Ages therewas an increase in trade between NorthAfrica, the Italian Peninsula fig.6.—Porcupine distal humerus with cut marks from the Crypta Balbi exedra, Rome (7th CAD) (photo by Jacopo and Sicily (Amari 1935; Brown 1989; Bresc 2003; De Grossi Mazzorin). Gioia & D’Angelo 2007), which is also attested by the abundance of potteryofAfrican origin that has been found, for instance, at the Crypta Balbi in praecipue si veneficio acciderit haec iniuria, item Rome (Saguì 2001). Exotic animals, such as the irenacei cinis cum melle aut corium combustum cum dromedary (Camelus dromedarius L.1758) are also pice liquida. Caput quidem eius ustum per se etiam commonly found in these periods (Wilkens 2003; cicatricibus pilos reddit, alopecias autem in ea cura- De Grossi Mazzorin 2006). The introduction of tione praeparari oportet novacula; ex sinapi quidam exotic species is likely to have occurred, at least in et aceto uti maluerunt. Quae de irenaceo dicentur, some cases, for pharmaceutical reasons as is for omnia tanto magis valebunt in hystrice” [The ashes instance the case for the remains of catfish of the of the head, tail and in fact the whole body of the family Clariidae, found in Rome in the area of mouse can be usedtotreat alopecia, especially when the Te nuta di Vallerano and Passaggio di Commodo theproblem was generatedbysomebody having in the Flavian Amphitheatre (De Grossi Mazzorin being bewitched; another remedy is represented by et al. 2005). The porcupine is also described by mixing hedgehog ashes with honey, or its calcined Pliny the Elder as having been used pharmaceuti- skin with liquified pitch. The burnt head of a cally. In particular he claims that it was useful in hedgehog, used on its own, also allows the re-growth dealing with diseases of the scalp: “Alopecias cinis of hair on scars, but to do so it is necessary to shave emurium capitibus caudique et totius muris emendat, the hairless areas fully with arazor; some like to
ANTHROPOZOOLOGICA • 2010 • 45 (2) 35 Masseti M., Albarella U. & De Grossi Mazzorin J.
fig.7.—Porcupine skull from Capanna Pelli (Cavallino, Lecce) (8th-9th C AD). Dorsal and ventral views of two fragments belonging to the same specimen (photo by Jacopo De Grossi Mazzorin). add mustard or vinegar. All the properties of the acidum halitum faciunt. Hocidem hystricis spina hedgehog also apply to the porcupine, and in fact fecisse ad firmitatem pertinet” [Teeth scratched with even moreso] (Nat. Hist., XXIX, 107). Another a vulture feather generate a bad breath but if the use was in taking care of the oral area (Nat. Hist., same is done with a porcupine quill, this consoli- XXX, 27): “Pinna vulturis si scalpantur dentes, dates the teeth]. Finally the porcupine was used to
36 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
prevent premature births (Nat. Hist., XXX, 123): “Partus conceptos hystricum cinis potus continent…” [porcupine ashes used in a potion keep the foetus in the uterus].
DISCUSSION
Although we can now — on the basis of recent archaeological findings — for the first time iden- tify the approximate period of the introduction of the porcupine to Italy,itisstill difficult to attribute this introduction to aspecific historic fig.8.—Capanna Pelli (Cavallino, Lecce): calibrated C14 event.Wecannot rule outthe possibility thatthis curve, which attributes the porcupine remains to 770-980 AD was causedbymigratingtribes in Late Antiquity, (95,4% probability,CeDAD Laboratory of the University of Salento). butthese areunlikely to havebeenthe Vandals or the Arabs as literarysources, such as the already mentioned al-Qazwînî,state that porcupines were occurrence amounts to a genuine introduction of still unknown in 13th CSicily,and therefore the species in the Italian countryside with stable that the introduction into the island occurred at and self-sufficient populations. alater date. More recent genetic workbyTrucchi and Sbordoni Recent genetic analysis seems to confirm the sug- (2009) supports the morphological evidence and gestion of a relatively recent history of the Italian implies that the Italian porcupine is characterised porcupine, as indicated by alow and simplified by a more complex pattern than a simple demo- level of genetic variability of the populations living graphic exponential growth from asingle propagule. in this country(Tr ucchi&Sbordoni 2007). In The attempted correlation between the molecular addition, research on cranial morphometric variabil- clock and the archaeo-historical evidence proposed ity undertaken by Angelici et al. (2003), has raised by Tr ucchi and Sbordoni (2009) is far less con- the suggestion that the porcupines living in conti- vincing. They suggest that the porcupine was intro- nental Italy havecharacteristics that aredistinct duced sometime between the early colonisation of from the animals living in Sicily and North Africa, Sicily by the Phoenicians and the first continental which, conversely,seem to be related to eachother. sub-fossil record that, citing Angelici et al. (2003), We must, however, not ignore the possibility of a they consider to be aspecimen from Basilicata dated ‘founder effect’, which may be at the origin of the to c. 1,500 years BP.Thereare several problems loss of heterozygosis of the Sicilian population. with this assumption, one of which concerns the Nevertheless, Angelici et al.(2003) believethat the date of the early Phoenician colonisation of Sicily, morphometric differences between the Italian con- which was regarded by Tr ucchi and Sbordoni (2009), tinental porcupines on the one hand and the Sicilian citing Leighton (1999), as having occurred and African ones on the other are too large to have c. 2,500 BP,when it should in fact be dated to developed in the last few millennia. If this is the several centuries earlier,between the 13th and case we should therefore conclude that the intro- 11th C BC (Tusa 1973; Camerata Scovazzo 2000). duction of the porcupine into Italy occurred as part In addition, Tr ucchi and Sbordoni (2009) areincor- of two or moreindependent events, whose geo- rect in citing Angelici et al. (2003) in reference to graphic origins were also different, at least as far the specimen from Basilicata, which these latter Sicily and continental Italy areconcerned. Although authors do not mention at all in their work. The it has nowbeen proven that thespecies was present natureofthis find from Basilicata is therefore in continental Italy at leastfromlate Antique/early doubtful, though it seems reasonable to assume medieval times,wecannot be surethat this that they arereferring to the above mentioned
ANTHROPOZOOLOGICA • 2010 • 45 (2) 37 Masseti M., Albarella U. & De Grossi Mazzorin J.
fig.9.—Location of Italian post-glacial archaeological sites, which have produced remains of Hystrix cristata Linnaeus, 1758 (modified from Masseti 2008) (prepared by Marco Masseti). porcupine specimen from San Giovanni di Ruoti, before the late Roman Empire. Although this sug- whose dating (Small 2005) was published two years gestion is consistent with our evidence, it certainly after the work by Angelici et al. (2003), and there- cannot be based on Pliny, whose work dates back forecould not havebeen mentioned by these latter. to 79 AD, thereforemuch earlier than the late Rather odd, and inconsistent with the timeframe Roman Empire. It is also hard to understand why they suggest elsewhere in the paper, is also the sug- they regard the crested porcupine to be at the same gestion by Tr ucchi and Sbordoni (2009) that Pliny time an invasivespecies on the one hand, and a the Elder’s words (Naturalis historia VIII, 53), indi- game species, whichcould have also have been kept cate that the species was not introduced into Italy as an exotic ornament “in rich Roman country
38 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
homes” (Trucchi &Sbordoni 2009, 6) on the other. Associazione CulturaleTethys, Firenze; Daphne Hills, Definitely unlikely is also the suggestion that Department of Zoology of the Natural History the porcupine could havebeen used as acircus Museum, London; Arthur MacGregor,Royal animal. Archaeological Institute, London; ArturoMorales, In general it seems thatTr ucchi and Sbordoni (2009) Departamento de Biologıa, Facultad de Ciencias, have tried to adapt the notoriously wayward evi- Universidad Autonoma de Madrid; Paul Mazza, dence of the molecular clock to an archaeo-historical Dipartimento di Scienzedella Terradell’Università context that they haveinsufficiently researched and di Firenze; Claudia Minniti, University of Sheffield; inconsistently adopted. This seems to fit with a Armando Nappi, Associazione Vivara Amici delle general trend towards accepting the genetic evi- Piccole Isole, Napoli; Gianni Palumbo, BioPhilia dence uncritically and without sufficient carein S.a.s.; Marco Rustioni, Museo di Paleontologia di integrating it with the results of other lines of Montevarchi, Arezzo;Leonardo Salari, Dipartimento investigation (see Masseti 2009b). di Scienze della Terra dell’Università “Sapienza” di Roma; Maurizio Sarà,Dipartimento di Biologia Animale, Università di Palermo; Barbara Wilkens, CONCLUDING REMARKS Dipartimento di Storia dell’Università di Sassari; Bruno Zava,Wilderness-Studi ambientali, Palermo. In conclusion, historical and archaeological evidence Dale Serjeantson(UniversityofSouthampton) point out to an initial introduction of the crested kindly commented on an earlier draft of this paper porcupine in Italy in the Late Antique/early medi- and Angela Trentacoste (University of Sheffield) eval period. Morphological and genetic evidence provided valuable help with the preparation of indicate that theremay havebeen several introduc- Fig. 1 and the final editing and formatting. tion events, possibly originating from different geographic areas. It is also important to emphasise that, although the chronological continuity of REFERENCES porcupine findings between late Antique and medi- eval times must not be ignored, we cannot discount Agricola G. [Bauer G.]. 1548. — De animantibus the possibility that the crested porcupine only subterraneis.Froben, Basilea. [Agricola G. 1563. — De gl’animali di sottoterra.Froben, Basilea. Tr ans- became an integral partofthe Italian fauna — with lated in Tu scanian language by Michelangiolo viable and self-sufficient populations — in later Florio Fiorentino.]. times. On the basis of the evidence currently avail- Amari M. 1935. — Storia dei Musulmani di Sicilia. able this seems to be the case in Sicily wherewe Vol. II. Romeo Prampolini, Catania. Amori G. &AngeliciF. M. 1992. — Note on the have so far no evidence of the presence of the spe- status of crested porcupine Hystrix cristata in Italy. cies beforeearly modern times. Futureresearch will Lutra 35: 44-50. no doubt contribute to provide further details to Amori G. & Angelici F. M. 1999. — Hystrix cristata this intriguing story. Linnaeus, 1758, in Mitchell-JonesA.J.,AmoriG., Bogdanowicz W.,Kryštufek B.,ReijndersP.J.H., Spitzenberger F. ,Stubbe M.,Thissen J.B.M., Vohralík V. &Zima J. (eds), Atlas of European Acknowledgements Mammals. The Academic Press, London: 308-309. In the course of this project we havebeen fortunate Anca F. 1860a. — Notice on the discovery of two to have the help of many friends and colleagues. Bone Caves in Northern Sicily. Quarterly Journal of the Geological Society of London 16(4): 459-460. We areparticularly grateful to Martin Clayton, Anca F. 1860b. — Note sur deux nouvelles grottes Royal Collection, Windsor Castle; Anna M. De ossifères découvertes en Sicile. Bulletin de la Société Marinis, Istituto Superioreper la Protezione ela Géologique de France 2, 17: 684-695. Ricerca Ambientale (ISPRA), Ozzano dell’Emilia Angelici F. M., Capizzi D., Amori G. & Luiselli L. 2003. — Morphometric variation in the skulls of (Bologna); Carolina Di Patti, Museo Geologico the crested porcupine Hystrix cristata from main- “G.G. Gemmellaro”, Dipartimento di Geologia e land Italy,Sicily,and northern Africa. Mammalian Geodesia, Università di Palermo; Flaviano Fanfani, Biology 68: 165-173.
ANTHROPOZOOLOGICA • 2010 • 45 (2) 39 Masseti M., Albarella U. & De Grossi Mazzorin J.
Attenborough D.,OwensS., Clayton M. & Camerata Scovazzo R. 2000. — La statuetta bronzea AlexandratosR.2007. — Amazing rare things. di Reshef (Melqart). Quaderni del Museo Archeolo- Royal Collection Publications, London. gico Regionale “Antonino Salinas” 6: 9-14. Bartolomei G. 1969. — Considerazioni ecologiche Cheylan G. 1988. — Compte-rendu de la table sulle faune pleistoceniche dell’Europa contenenti ronde : répartition géographique et statut des mam- scimmie ed istrici. Scritti sul Quaternario in onore di mifèresmenacésdans les îles méditerranéennes. Angelo Pasa. Memorie del Museo Civico di Storia Bulletin d’Écologie 19(2-3): 481-484. Naturale di Verona (Fuori Serie) 3: 39-52. Colonnelli G. & De Grossi Mazzorin J. 2000. — Bartolomei G. 1980. — Micromammiferi del Plio- Nuovidati sull’alimentazione aFarnese (VT) nei Pleistocene, in Ivertebrati fossili italiani. Catalogo secoli XV eXVI, in Malerba G.,Cilli C. & della Mostra. Verona: 249-258. Giacobini G. (eds), Atti del 2° Convegno Nazionale Bashir J.T.2009. — Biodiversity of Farwa Island, in di Archeozoologia. Asti, 14-16 novembre1997. Poulakakis N. & Vardiniyannis K. (eds), Abstracts ABACO Edizioni, Forlì: 369-376. of the 11th International Congress on the Zoogeogra- Corbet G.B. 1978. — The mammals of the Palaearctic phy, Ecology and Evolution of Eastern Mediterranean Region. Ataxonomic review.British Museum (Natural (21-25 September 2009), Herakleion. Hellenic Zoo- History) &Cornell University Press, London & logical Society, Herakleion: 187. Ithaca. Bonfiglio L., Agnesi V., Masini F. & Di Maggio C. Corbet G.B. &JonesL.A. 1965. — Thespecific 2004. — Quaternary Eustatic Fluctuations and Bio- characters of the crested porcupines, subgenus chronologyofVertebrate-Bearing Deposits Correlated Hystrix. Proceedings of the Zoological Society of with Marine Te rraces in Sicily.32rd International London 144(2): 285-300. Geological Congress (Florence, Italy). APAT, Roma. Cornalia E. 1871-1872. — Fauna d’Italia. Parteprima. Brehm A. 1916. — Die Säugetiere in Brehms Tierleben. Catalogo descrittivodei Mammiferi osservati fino ad Bibliographisches Institut, Leipzig. ora in Italia. Francesco Vallardi editore, Milano. Bresc H. 2003. — Venuti dal nord. La Sicilia nor- Costa O.G. 1839. — Fauna del Regno di Napoli. Quadro manna, in Benigno F. &Giarrizzo G. (eds), delle specie indigene ed acclimatizzate della classe dei Storia della Sicilia. 1. Dalle origini al Seicento. Mammiferi. Stamp. Azzolino e Compagno, Napoli. Editori Laterza, Roma; Bari: 83-93. De Grossi Mazzorin J. 2006. — Cammelli nell’anti- Brink Van Den F. H. 1969. — AField Guide to the chità:lepresenzeinItalia, in Sala B. &Tecchiati U. Mammals of Britain and Europe. Houghton Mifflin, (eds),Studi di archeozoologia in onorediAlfredo Boston. Riedel. Provincia di Bolzano, Bolzano: 231-242. Brown P. 1989. — The world of Late Antiquity AD De Grossi Mazzorin J. & De Venuto G. 2006. — 150-750. Thames & Hudson, London. Indagini archeozoologiche presso il centro medievale Buffon Comte De Leclerc G.-L. 1766. — Histoire di S. Salvo (Chieti), in Fiore I. & Tagliacozzo A. naturelle, génerale et particulière. Paris. (eds), Riassunti-Abstracts del 5° Convegno Nazionale Burgio E. 1997. — Le attuali conoscenze sui mammi- di Archeozoologia. Rovereto,10-12 novembre2006. feri terrestri quaternari della Sicilia, in Tusa S. (ed.), Associazione Italiana di Archeozoologia-Museo Prima Sicilia, Alle origini della società siciliana.Regione Civico di Rovereto, Rovereto: 73. Siciliana, Assessorato dei beni Culturali, Ambientali e De Grossi Mazzorin J. &Minniti C. 2001. — della Pubblica Istruzione-Ediprint, Palermo: 54-71. L’ allevamento e l’approvvigionamento alimentare di Burgio E. & Di Patti C. 1990. — I vertebrati fossili una comunità urbana. La gestione degli animali a della Grotta di S. Teodoro. Naturalista Siciliano 14: Roma tra il VII ed il Xsecolo, in Arena M.S., 1-19. Delogu P. ,Paroli L.,Ricci M.,Saguì L. & Burgio E.,Costanza M. &DiPatti C. 2002. — Vendittelli L. (eds), Roma dall’antichità al Idepositi avertebrati continentali del Pleistocene Medioevo. Archeologia estoria del Museo Nazionale della Sicilia occidentale. Naturalista Siciliano 26(3-4): Romano, Cripta Balbi. Roma: 69-78 & 328-330. 229-282. De Grossi Mazzorin J. & Minniti C. 2004. — Lo Burgio E.,Costanza M.,DiPatti C. &Mannino G. studio dei resti animali: un contributo alla storia 2005. — Attuali conoscenzesulle faune dei siti del consumo alimentareaRoma tra il XII eil preistorici della Sicilia occidentale, in Fiore I., XIII secolo, in Parmegiani N. &ProntiA. (eds), Malerba G. &Chilardi S. (eds), Atti del S. Cecilia in Trastevere. Nuoviscavi ericerche, Monu- 3° Convegno Nazionale di Archeozoologia. Siracusa, menti di Antichità Cristiana.Pontificio Istituto di 3-5 novembre 2000. Istituto Poligrafico dello Stato, Archeologia Cristiana II serie, XVI. Città del Vaticano: Roma: 145-171. 283-306. Burton J.A. & Pearson B. 1987. — Collins Guide to De Grossi Mazzorin J., Minniti C. & Rea R. 2005. the Rare Mammals of the World. Collins, London. — De ossibus in anphitheatroFlavio effossis: Cabrera A. 1932. — Los mamiferos de Marruecos. 110 anni dopo irinvenimenti di Francesco Luzj, Trabajos del Museo Nacional de Ciencias Naturales. in Malerba G. &Visentini P. (eds), Atti del Serie Zoologica 57: 5-361. 4° Convegno Nazionale di Archeozoologia, Pordenone,
40 ANTHROPOZOOLOGICA • 2010 • 45 (2) The crested porcupine, Hystrix cristata L., 1758, in Italy
13-15 novembre 2003. Quaderni del Museo Archeo- Lacepède C. 1830. — Storia Naturale di Buffon. logico del Friuli Occidentale 6. Museo delle Scienze, Tomo XVI. Tipografia Vignozzi, Livorno. Comune di Pordenone: 337-348. LandsbergS.1998. — The medieval garden.The De MarinisA.M.,Masseti M. &Sforzi A. 1996. British Museum Press, London. — Note on the non-flying Terrestrial Mammals of Leighton R. 1999. — Sicily before history: an archaeo- the Tu scan Archipelago, Northern Tyrrhenian Sea logical surveyfromthe Palaeolithictothe Iron Age. (Italy). Museo Regionale di ScienzeNaturali Bollet- Cornell University Press, Ithaca. tino (Torino) 14(1): 275-281. Lovari S. 1993. — Evoluzione recente delle popolazi- Esu D. &KotsakisT.1986. — Ivertebrati eimol- onidigrandi mammiferi della fauna d’Italia. Acca- luschi continentali fossili dell’Italia meridionale: prob- demia Nazionale dei Lincei. Contributi del Centro lemi biogeografici. Biogeographia (N.S.) 13: 57-81. Linceo Interdisciplinare “Beniamino Segre” 86: 21-37. FreedbergD. 2002. — The eye of the lynx. Galileo, his MacKinnon M.R. 2002. — The Excavations of San friends, and the beginnings of the modern natural his- Giovanni di Ruoti. Vol. III.The Faunal and Plant tory. The Univeristy of Chicago Press, Chicago. Remains. University Press of Toronto, Toronto. Ghigi A. (ed.). 1959. — La Fauna. Conosci l’Italia, III. Masseti M. 1996. — The Postglacial diffusion of the Touring Club Italiano, Milano. genus Dama Frisch, 1775, in the Mediterranean Ghisio G.B. 1779. — Mappa della Sicilia. Illustrazi- region. Supplemento alle Ricerche di Biologia della oni laterali con vegetazione, animali, piante di città Selvaggina 25: 7-29. monumenti. S.e., Palermo. Masseti M. 2008. — Erinaceomorfi, Soricomorfi, Gioia C. &D’Angelo F. 2007. — Analisi minero- Lagomorfi eRoditori endemici ed antropocori petrografiche sui reperti dello scarico di fornaci di dell’Italia continentale e peninsulare, in Amori G., PalazzoLungarini aPalermo (fine XI-inizio Contoli L. &Nappi A. (eds), Fauna d’Italia. XII secolo). Archeologia Medievale 34: 337-343. Mammalia II. Erinaceomorpha, Soricomorpha, Lago- Gisbert J.,García-Perea R. &Martínez- morpha, Rodentia.Edizioni Calderini de Il Sole 24 Medina F. J. 2002. — Hystrix cristata Linnaeus, ORE Business Media, Milano: 33-65. 1758 — Puercoespín, in Palomo L.J. & Gisbert J. Masseti M. 2009a. — Holocene mammals of Libya: (eds), Atlas de los Mamiferos terrestres de España. A biogeographical, historical and archaeozoological Direcció General de Conservación de la Naturaleza- approach. Journal of Arid Environments, SECEM-SECEMU, Madrid: 440-443. doi:10.1016/j.jaridenv.2009.07.008. Hahn D. 2003. — The TowerMenagerie.Simon & Masseti M. 2009b. — Mammals of the Mediterra- Schuster, London; New York. nean islands: homogenisation and the loss of biodi- Hochner N. 2001. — Louis XII and the porcupine: versity. Mammalia 73: 169-202. transformations of a royal emblem. Renaissance Stu- Miller G.S. 1912. — Catalogue of the mammals of dies 15(1): 17-36. Western Europe (Europe exclusive of Russia) in the col- Honacki H.H., Kinman K.E. & Koeppl J.W.1982. lection of the British Museum.British Museum — Mammal species of the world. Ataxonomic and (Natural History), London. geographic reference. Allen Press Inc. & The Associa- Minà Palumbo F. 1868. — Catalogo dei mammiferi tion of Systematic Collections, Lawrence. delle Sicilia. Stamperia Tamburello & C., Palermo. KingA.2002. — Mammals. Evidence from wall Minniti C. 2005. — L’approvvigionamento alimen- paintings, sculpture, mosaics, faunal remains, and tare a Roma nel Medioevo: analisi dei resti faunistici ancient literary sources, in Feemster Jashemski W. proveneienti dalle aree di scavo della Crypta Balbi e & Meyer F. G. (eds), The natural history of Pompeii. di Santa Cecilia, in Fiore I.,MalerbaG. & Cambridge University Press, Cambridge: 401-450. Chilardi S. (eds), Atti del 3° Convegno Nazionale KingA.C., RhodesP.A., Rielley K. & ThomasK.D. di Archeozoologia. Siracusa, 3-5 novembre 2000. Isti- 1985. — Iresti animali, in Ricci A. (ed.), Settefinestre. tuito Poligrafico dello Stato, Roma: 469-492. Una villa schiavista nell’Etruria romana. La villa e i Minniti C. 2009. — Economia ealimentazione nel suoi reperti.Edizioni Panini s.p.a., Modena: 278-309. Lazio medievale: nuovi dati dalle evidenze archeozo- Kingdom J. 1974. — East African Mammals. An Atlas ologiche. Archeologia Medievale 36: 275-283. of Evolution in Africa. Vol. II. Part B. Academic Mongitore D.A. 1743. — Della Sicilia ricercata nelle Press, London. cose più mirabili. Francesco Valenza, Palermo. KotsakisT.1979. — Sulle mammalofaune quater- Niethammer G. 1963. — Die Einbürgerung von Säu- narie siciliane. Bollettino del Servizio Geologico getieren und Vögeln in Europe. Parey, Hamburg. d’Italia 99: 263-276. Niethammer J. 1982. — Hystrix cristata Linnaeus, KotsakisT., Abbazzi L., Angelone C., Argenti P. , 1758 — Stachelschwein, in Niethammer J. & Barisone G.,Fanfani F. ,Marcolini F. &Masini F. Krapp F. (eds), Handbuch der SäugetiereEuropas. 2003. — Plio-Pleistocene biogeography of Italian Band 2/I. Rodentia I(Cricetidae, Arvicolidae, mainland micromammals. Deinsea 10: 313-342. Zapodidae, Spalacidae, Hystricidae, Capromyidae). Kurtèn B. 1968. — Pleistocene Mammals of Europe. Akademische Verlagsgesellschaft, Wiesbaden: Weidenfeld & Nicolson, London. 588-605.
ANTHROPOZOOLOGICA • 2010 • 45 (2) 41 Masseti M., Albarella U. & De Grossi Mazzorin J.
Orsomando E. & Pedrotti F. 1976. — Notizie sulla in Italia meridionale. Foggia, 12-14 febbraio 2004. presenza esull’habitat dell’istrice nelle Marche e STAIM 1. Edipuglia, Bari: 127-131. nell’Umbria, in Pedrotti F. (ed.), S.O.S. Fauna. Smit C.J. &VanWijngaarden A. 1981. — Threat- Animali in pericolo in Italia.Edizione W.W.F. ened mammals in Europe.Akademische Verlagsge- Tipografia Succ. Savini-Mercuri, Camerino: 249-263. sellschaft, Wiesbaden. Ortolani G.E. &Rafinesque Schmaltz C.S. 1810. — Tagliacozzo A. 1993. — Archeozoologia della Grotta Statistica generale di Sicilia. Reale Stamperia, Palermo. dell’Uzzo, Sicilia. Supplemento al Bullettino di Palet- Plot R. 1705. — The Natural History of Oxford-shire: nologia Italiana nuova serie II, 84: 1-278. Being an Essay towards the Natural HistoryofEngland. Toschi A. (ed.). 1965. — Mammalia. Lagomorpha, Westminster City Libraries, London. Rodentia, Carnivora, Ungulata, Cetacea. Collana Pratesi F. & Tassi F. 1974. — Guida alla natura della “Fauna d’Italia”. Vol. VII. Calderini, Bologna. Sicilia. Arnoldo Mondadori Editore, Milano. Trucchi E. &Sbordoni V. 2007. — Dati preliminari Ranck G.L. 1968. — The Rodents of Libya. Ta xonomy, sulla variabilità elastruttura genetica delle popolazioni Ecologyand Zoogeographical Relationships.Smithso- italiane di Hystrix cristata.Valutazioni circa l’origine e nian Institution Press, Washington D.C. lo stato del popolamento italiano, in Angelici F. M., Regalia E. 1903. — Sulla fauna delle grotte di Grola e Petrozzi F. &Galli A. (eds), Atti del Convegno Inter- Zachito. Archivio per l’Antropologia el’Etnologia nazionale “Fauna Problematica: Conservazione e 33(21): 7-75. Gestione”.Montefiascone (VT), 8-9 giugno 2007.Sta- Riquelme Cantal J.A. & Morales Muñiz A. 1997. bilimento tipolitografico Agnesotti, Viterbo: 20. — APorcupine find from Roman Africa with a Trucchi E. &Sbordoni V. 2009. — Unveiling an reviewofarchaeological data from circummediter- ancient biological invasion: molecular analysis of an ranean sites. Archeofauna 6: 91-95. old European alien, the crested porcupine (Hystrix Rustioni M., Mazza P. & Balestri M. 1999. — The cristata). BMC Evolutionary Biology 9(1): 109. Middle-Late Pleistocene mammal fauna from Mon- Tusa V. 1973. — La statuetta fenicia del Museo Nazio- tignoso (Leghorn, Central Italy). Il Quaternario. nale di Palermo. Rivista di Studi Fenici 1-2: 173-177. Italian Journal of QuaternarySciences 12(1): 105-111. Ververs G. 1976. — London’s Zoo. The Bodley Head, Ruta C. (ed.). 2007. — La Sicilia nelle geografie arabe London; Sydney. del medioevo. Edi.bi.si., Messina. Villari P. 1995. — Le faune della tarda preistoria nella Rybot D. 1972. — It began beforeNoah.Michael Sicilia orientale. Ente Fauna Siciliana, Siracusa. Joseph, London. Van WeersD.J. 1994. — The porcupine Hystrix refossa Saguì L. 2001. — Roma eilMediterraneo: la circolazi- Gervais, 1852 from the Plio-Pleistocene of Europe, one delle merci, in Arena M.S.,DeloguP. ,Paroli L., with notesonother fossil and extant species of the Ricci M.,Saguì L. &Vendittelli L. (eds), Roma genus Hystrix. Scripta Geologica 106: 35-52. dall’antichità al Medioevo. Archeologia estoria del Museo Van Weers D.J. 2005. — A taxonomic revision of the Nazionale Romano,Cripta Balbi.Roma: 62-68. Pleistocene Hystrix (Hystricidae, Rodentia) from Salari L. & Sardella R. 2008. — Hystrix vinogradovi Eurasia with notes on the evolution of the family. Argyropulo, 1941 nel Pleistocene italiano in Abstract Contribution to Zoology 74 (3/4):
Submitted on 26 January 2010; accepted on 18 May 2010
42 ANTHROPOZOOLOGICA • 2010 • 45 (2)