DOES THE OCCURRENCE OF PREDATORS ON CENTRAL MEDITERRANEAN ISLANDS AFFECT THE BODY SIZE OF MICROMAMMALS ? G. Amori, M. Masseti

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G. Amori, M. Masseti. DOES THE OCCURRENCE OF PREDATORS ON CENTRAL MEDITER- RANEAN ISLANDS AFFECT THE BODY SIZE OF MICROMAMMALS ?. Vie et Milieu / Life & Environment, Observatoire Océanologique - Laboratoire Arago, 1996, pp.205-211. ￿hal-03100653￿

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DOES THE OCCURRENCE OF PREDATORS ON CENTRAL MEDITERRANEAN ISLANDS AFFECT THE BODY SIZE OF MICROMAMMALS ?

G. AMORI* M. MASSETI** *Centro Genetica Evoluzionistica, CNR, Via Lancisi 29, 00161 Roma, Italia **Istituto di Antropologia, Université di Firenze, Via Del Proconsolo, 12, 50122 Firenze, Italia

MICROMAMMALS ABSTRACT. - Many hypothèses have been formulated to explain the increase in PREDATORS micromammal body size and the decrease in macromammal body size in islands. MEDITERRANEAN ISLANDS Among thèse, the occurrence of predators is considered as a possible factor BODY SIZE influencing size. This paper analyses the current composition of the terrestrial micromammal populations of the Central Mediterranean islands, stressing the populations showing body size différences, and studying the relation of the latter with the présence of predators, such as carnivores, birds of prey and snakes. This study is based on the présent state of knowledge about the micromammals of the Central Mediterranean islands, including unpublished data. The results obtained suggest that predators have slight influence on the body size of island micromammals. Their size increase seems to be due to other factors, such as réduction of food supply, increase of intraspecific compétition and, finally, endothermic control.

MICROMAMMIFERES RÉSUMÉ. - Plusieurs hypothèses ont été formulées pour expliquer l'augmentation PRÉDATEURS de taille des micro-Mammifères et la diminution de taille des macro-Mammifères ÎLES MÉDITERRANÉENNES insulaires. Parmi ces hypothèses, la présence de prédateurs a été considérée comme TAILLE un facteur possible influençant la taille. Cette étude se propose de décrire la composition actuelle des populations de micro-Mammifères terrestres des îles de Méditerranée centrale et de mettre en évidence les populations qui montrent des différences de taille. La présence de prédateurs (Carnivores, Rapaces et Ophidiens) sur les îles est également prise en compte et discutée. Cette recherche est développée sur la base des connaissances disponibles sur les micro-Mammifères vivant sur les îles de la Méditerranée centrale et de données originales. Les résultats obtenus suggèrent que les prédateurs ne jouent qu'un rôle assez réduit en ce qui concerne l'impact sur la taille des micro-Mammifères. L'accroissement des dimensions de ces derniers peut être mis en relation avec d'autres facteurs, par exemple la réduction de la disponibilité alimentaire, la compétition intraspécifique et enfin les adaptations endothermiques.

INTRODUCTION double in body size with respect to the extant edible , Myoxus glis (Petronio, 1970). The Upper Pleistocene of yielded the giant Adler & Levins (1994) and Fons et al. (1995) withe-thooted shrew Crocidura esuae (Kotsakis, indicate that the extant représentatives of the 1986), and remains of the large shrew of the genus micromammals communities (Insectivora and Ro- Episoriculus have been provided by the Late Pleis- dentia) of the Mediterranean islands are generally tocene deposits of Corsica and (Vigne, reputed to be distinguished from their mainland 1992). Large body size Muridae species have been relatives in several characteristics, such as the identified in the Late Pleistocene of Corsica and increase in body size. Sardinia (Vigne, 1992), Crète (Mayhew, 1977) and According to paleontological évidence, also the Cyprus (Bate, 1903; Boekschoten & Sondaar, extinct Pleistocene endémie micromammals of the 1972). In its terminal phase, the Pleistocene fauna Mediterranean islands often displayed an increase of the Balearics comprised the large dormouse in (body) size. The dormouse species Leithia me- Hypmomys morpheus, and the soricid Nesiotes hi- litensis of Sicily and Malta, for example, was daldo (Alcover et al., 1981). 206 G. AMORI, M. MASSETI

Table I. - List of the species of shrews, , carnivores, birds of prey and snakes occurring on the considered islands (data also from Parlanti et al., 1988; Amori, 1993; Lo Valvo et al., 1993; Montemaggiori et al., 1993; Schembri, 1993; Yeatman-Berthelot & Jarry, 1994; Sarà, 1995; Sarà & Casamento, 1995).

?m aS ao 1 S 3 £ R S 3 ■ B 1 I 1 î 1 £ s 76 S 5- ISLAND ARE A iS 4 codddujggs ce < 5 d d o ui ui Linosa 5,34 Levanzo 5,61 8,6 Pianosa 10,3 Marettimo 12,06 Stromboli 12,28 Capraia 19,26 Favignana 19,7 Lampedusa 20,2 Giglio 21,2 Vulcano 21,2 Salina 26,5 Lipari 37,5 Asinara 50,9 Gozo 67,1 86 Elba 223,5 Malta 245,7 Corsica 8722 Sardinia 23833 Sicily 25460

Many hypothèses have been formulated to ex- abundant on the islands (Sondaar, 1977; Azzaroli, plain the increase in micromammal body size on 1982; Weesie, 1982 and 1988). islands. The factors that might be responsible for The aim of this study is to assess the relation- this increase in size are thought to include a ships between the occurrence of predators on is- contraction of food supply, a réduction of inter- lands and the différences (increase) in micromam- specific compétition, an increase of intraspecific mal body size. compétition, and, even, a decrease in prédation The current composition of the non-flying ter- (Thaler, 1973; Heaney, 1978; Angerbjôrn, 1986; restrial micromammal populations of the Central Orsini & Cheylan, 1988; Libois & Fons, 1990). Mediterranean islands, has been analyzed with The decrease in prédation could be explained particular attention to the populations with diffé- by a relative lack of carnivores on islands, as rences in body size, and of predators, such as paleontological évidence has shown (Sondaar, carnivores, birds of prey (nocturnal and diurnal) 1977). It is generally assumed that the absence of and snakes, which are mainly specialized in fee- carnivores permits the survival of larger micro- ding on small . individuals. As far as is presently known, however, this is true only in part. In fact, although carnivores of STUDY AREA, MATERIAL AND METHODS large body size are still unknown from the Medi- terranean islands, Quaternary endémie carnivores For the présent study, we have considered the islands of médium and small size are documented on located in the Central , between the Corsica, Sardinia, Sicily, Malta, Crète and Cyprus 5° and the 26° meridians (fig. 1). On the présent state (Alcover & McMinn, 1994; Masseti, 1995). of knowledge, complète data on the faunistic composi- tions, regarding micromammals, carnivores, birds of Pleistocene island micromammals might have prey, and mammal-eating snakes, are available only for also represented the main prey item for endémie the 21 islands listed in Table I. For this reason it was raptors, such as the Cretan terrestrial owl Athene not possible to consider ail the Central Mediterranean cretensis, and for continental raptors, which were islands. PREDATORS ON MEDITERRANEAN ISLANDS 207

considerably from the Quaternary ones. Most of the latter were endémie and became extinct after the advent of the human Neolithic colonization (Azzaroli, 1962; Alcover, 1982; Weesie, 1988; Vigne, 1992). The extant mammalian communities consist only of species that are of continental origin, with non évidence of endemization, the diffusion of which is essentially related to human intervention beginning in prehistorical times and destined to continue throughout the historical âges (Masseti, 1993; 1995; Masseti & Vianello, 1993; Vigne & Alcover, 1985). The most widespread micromammal species are the Crocidurinae and Muridae (Table I, fig. 2A,B). Often, only a few species (two, or at most three) are présent on the same islands. Oc- casionaly, the micromammalian communities dis- play a very similar composition on small islands, such as Capraia (19.26 km2, Crucidura suaveo- lens, Mus domesticus, and Rattus rattus), and Le- vanzo (5.61 km2, C. sicula, M. domesticus and R. rattus). In several cases micromammals on island show Fig. 1. - Central Mediterranean islands considered in genotypic and phenotypic modification such as the présent work. 1 = Linosa ; 2 = Levanzo ; 3 = Usti- the tendency to produce larger forms than those ca; 4 = Pianosa; 5 = Marettimo; 6 = Stromboli; 7 = Capraia; 8 = Favignana; 9 = Lampedusa; 10 = Giglio; found on the nearest continental areas (Case, 11 = Vulcano; 12 = Salina; 13 = Lipari; 14 = Asinara; 1978). Large taxa have been reported from Cor- 15 = Gozo; 16 = Pantelleria; 17 = Elba; 18 = Malta; sica, Crocidura suaveolens (Kahmann & Kah- 19 = Corsica; 20 = Sardinia; 21 = Sicily. Filled circle mann, 1954; Vlasâk & Niethammer, 1990), Sar- indicate large body size taxa. dinia, Suncus etruscus pachiurus (Sans-Coma et •5 = Apodemus sylvaticus; • 13 = Rattus rattus, al., 1985), Lipari, Eliomys quercinus liparensis Eliomys quercinus liparensis; • 16 = Crocidura russula (Kahmann, 1960), Elba, Apodemus sylvaticus il- cossyrensis, Apodemus sylvaticus hermani; • 17 = Apo- vanus (Kahmann & Niethammer, 1971), Maret- demus sylvaticus ilvanus; • 19 = Crocidura suaveolens; timo, A. sylvaticus (Sarà & Casamento, 1995), and • 20 = Suncus etruscus pachiurus. Pantelleria, C. russula cossyrensis (Contoli & Amori, 1986) and A. sylvaticus hermani (Felten & Storch, 1970). Populations of large-sized black This research is based on the existing literature on rats (Rattus rattus) have been reported from Lipari the Mediterranean island faunistic composition, as well (Cristaldi et al., 1985) (fig. 1). as unpublished data. Another phenotypic modification is the ten- Ail the small mammal species (Insectivora, Roden- dency to produce dark colored forms such as the tia) présent on thèse islands have been concerned, with the exception of large-size species, such as Hystrix garden dormouse {E. quercinus sardus) and the cristata, which occurs on Elba and Sicily (Amori & (Myoxus glis melonii) of Corsica Angelici, 1992; Lovari, 1993), and the recently intro- and Sardinia, which are also smaller than their duced species Myocastor coypus recorded from Sicily continental counterparts (Toschi, 1965). (Reggiani et al., 1993). Ail the carnivores présent on Reduced pressure of competitors and predators the considered Central Mediterranean islands are listed is found on both the smallest islands and on Sicily, in Table I. On the other islands only the species of birds of prey and snakes characterized by food habits Sardinia and Corsica (fig. 3A). based mainly on the consumption of micromammals are At présent the carnivores that occur on the concerned. The accumulated data do not support stati- Central Mediterranean islands are restricted to a stical treatment. For this reason, our remarks can only few species possibly of anthropochorous origin, be descriptive, and are based almost exclusively on including Vulpes vulpes, Mustela nivalis, Martes observations. martes, and Felis silvestris on Sardinia (Schenk, 1976; Vigna Taglianti, 1988) and Sicily (Vigna Taglianti, 1988; Catalisano et al, 1991b) RESULTS AND DISCUSSION (fig. 3B). The same species are also présent on Corsica (Salotti, 1992), where, however, the oc- It should be noted that the extant mammalian currence of the pine marten, previously recorded assemblages of the Mediterranean islands differ by Forsyth Major (1882), Verbeek (1974) and 208 G. AMORI, M. MASSETI

12 10 8 6

CO O <0 (C o (0 (0 nj o o S g S S I E ? g _ to co to m >. co N o « c o -s g m = - i S O "S g- > ç 5 » c t c n C TJ s j>DH(l)k«01(DO 1 « 3 ïiïofï □Erinaceidae OCrocidurinae ■ Myoxid ■ Muridae i iMcrotidae [□insectivores M rodents HP carnivores □birdsofprey EHsnakes

16 14 12 10 nnnn nn nnnn n n -cooaînJcC(On3> S >=■ N C 1 J O F ~ ; « o .b .a E c3 D m 3... Œ m 10 J M c p c a c TJ ".5 s •3 CD 0) R CO O) 0) U -= CO ■I11 1 ■g

□micromammals ■carnivores

Fig. 2. - A, number of shrew and species of the Fig. 3. - A, number of species of shrews, rodents, considered islands; B, number of micromammal and carnivores, birds of prey and snakes of the considered carnivores species of the considered islands. islands; B, number of micomammal and predator spe- cies of the considered islands

Cholley (1982) has not been recently confirmée! (Vigne, 1992; Salotti, 1992). On small islands, in on Pantelleria, where carnivores are not documen- contrast, carnivores are limited to the weasel on ted either on paleontological or on historical Asinara (Cossu et al, 1994), and Malta (Schem- grounds. bri, 1993), and the pine marten on Elba (De Ma- Carnivores may occur on islands similar or rinis & Masseti, 1993a and 1993b). larger than Elba (223.5 km2) and Malta (245.7 Birds of prey are represented by several species km2) (fig. 2B). One of the few exceptions is rep- (Table I) that also occur on the closest continental resented by the small island of Asinara areas (Brichetti et al, 1992). The most common (50.9 km2), where the occurrence of M. nivalis is island résident species (9 species) are considered perhaps related to the présence of a higher number in the présent study. The most fréquent are the of micromammalian species than that occurring barn owl (Tyto alba) and the kestrel (Falco tin- on other islands of similar dimentions (Table I). nunculus). Snakes that prey mainly upon micro- Other predators, such as birds of prey and mammals are limited to 8 species (Arnold & Bur- snakes, occur more or less on ail the islands with ton, 1978; Obst et al, 1984). On Sicily, the the same number of species. Thus, their ecological occurrence of Elaphe quatuorlineata, was perhaps pressure on prey can be considered roughly iden- erroniously reported by Arnold & Burton (1978), tical on ail the islands. Bruno (1988), Parlanti et al (1988) and Amori et al. (1993), but it was not quoted in Catalisano et Among rodents, E. q. sardus and M. glis me- al. (1991a). According to M. Sarà and G.F. Turrisi lonii are smaller than their continental relatives, (pers. comm.) the présence of the species on the in an ecological context with the highest number island is still uncertain. of predatory and compétitive species of ail the Mediterranean islands. As mentioned above, with the exception of Sicily, Sardinia and Corsica, single species of In contrast, it should be noted that with the carnivores occur on few Central Mediterranean decrease in potentially compétitive species and in islands. Thus, the occurrence of carnivores does restricted environments, micromammal show a not seem to have any relation with micromammal tendency to increase body size. In fact the micro- size. For example, the large-sized A. sylvaticus mammals of Lipari comprise only the large E. q. ilvanus lives on Elba, together with M. martes, liparensis, M. domesticus, and R. rattus, whereas while the large-sized A. sylvaticus hermani lives on Corsica and Sardinia the small E. quercinus PREDATORS ON MEDITERRANEAN ISLANDS 209 and Myoxus glis coexist with Erinaceus euro- AMORI G., ANGELICI F.M., FRUGIS S., GANDOLFI paeus, Crocidura suaveolens (Corsica), Crocidura G., GROPPALI R., LANZA B., RELINI G., G. VI- russula (Sardinia), Suncus etruscus, Mus domesti- CINI, 1993. Vertebrata. In Minelli A., Ruffo S. and cus, Rattus rattus, Rattus norvegicus and Apode- S. La Posta (eds.). Checklist délie specie délia fauna italiana, 110, Calderini Bologna, 83 p. mus sylvaticus. Also a unique spécimen of E. quercinus collected on the island of Capraia in ANGERBJÔRN A., 1986. Gigantism in island popula- 1885, and now kept at the Natural History Mu- tions of wood mice (Apodemus) in Europe. Oikos 47 : 47-56. séum "La Specola" of the University of Florence (MZUF n° 6403) is characterized by a large body ARNOLD E.N. and J.A. BURTON, 1978. A field guide size. Its skull display a condylobasal length of to the reptiles and amphibians of Britain and Europe. Collins Ltd. London, 275 p. mm 33.8, and a greatest zygomatic width of mm 21.5. This spécimen is however, an exceptional AZZAROLI A., 1962. Il nanismo dei cervi insulari. record, probably referable to an accidentai intro- Paleont. It. 56 : 1-31. duction. Today, in fact, the garden dormouse is a AZZAROLI A., 1982. Insularity and its effects on ter- species unknown among the micro-mammalian restrial vertebrates : evolutionary and biogeographic fauna of Capraia (Amori, 1993 and Table I). aspects. In : Palaeontology : Essential of Historical Geology, Edited by E. Montanaro Gallitelli, S.T.E.M. The size increase of island micromammals does Mucchi, Modena : 193-215. not seem to be directly related to the occurrence BATE' D.M.A., 1903. The Mammals of Cyprus. Proc. of predators as observed by Sarà & Casamento Zool. Soc. London 2 : 341-348. (1995) for A. sylvaticus. The explanation is per- BOEKSCHOTEN GJ. and P.Y. SONDAAR, 1972. On haps to be found in other factors such as réduction the fossil mammals of Cyprus. Proc. Kon. Ned. of the food supply, interspecific and intraspecific Akad. Wetensch. 75 : 306-338. compétition and also endothermic adaptations. BRICHETTI P., P. DE FRANCESCHI and N. BAC- The results of this research are preliminar and CETTI (eds.) 1992. Aves I. Gaviidae - Phasianidae. may be conditioned by the extention of the survey. Fauna d'Italia, Edizioni Calderini Bologna, 964 p. More research is needed to better understand the BRUNO S., 1988. Considerazioni sull'erpetofauna délia ecological rôle of the Holocene anthropochorous Sicilia. Bull. 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