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Biodiversity Journal, 2017, 8 (1): 249–278 MONOGRAPH

The endemic of the sicilian

Calogero Muscarella1 & Alessandro Baragona2

1Via D’Ondes Reggio 8/A Scala G, 90127 , ; e-mail: [email protected] 2Via Piersanti Mattarella 5, 90020, Sciara, Palermo, Italy; e-mail: [email protected]

ABSTRACT In this survey we propose an analysis of the endemic fauna (, Arthropoda, Chordata) present in the 14 major circum-sicilian islands and in (Strait of , ). Overall, 111 endemic taxa between and subspecies have been identified. The largest taxonomical groups are Tenebrionid and Curculionids (respectively 18 and 16 taxa) and Gastropods (20 taxa), due to their strong inclination to insular differentiation, which is inversely proportional to their vagility. The number of endemic taxa per is positively associated to the extent of the surface but not to the distance from the closest continental mass or altitude or geological origin. The most important connection is with the complex paleo- geographic history pertaining the different insular complexes.

KEY WORDS ; circum-sicilian islands; paleogeography.

Received 15.03.2016; accepted 11.11.2016; printed 30.03.2017 Proceedings of the 3rd International Congress “, Mediterranean, Society”, September 4th-6th 2015, - Vendicari (Italy)

INTRODUCTION or “chersogenic” islands; in the second one, about “talassogenic islands” (see also Troia, 2012 and cor- Small islands and archipelagoes have always responding bibliography). raised the interest of biologists, especially due to The organisms actively or passively colonizing the peculiar and fauna which inhabits them the islands undergo a severe selective pressure (, 2008; Minelli, 2012). The Mediterranean is triggered by both isolation and environmental a basin gathering one of the largest insular characteristics; as an adaptative response, they groups in the world with approximately 5000 is- develop more or less pronounced biological pecu- lands. This number raises up to approximately liarities, favouring the formation of endemic species 11,879 if we consider the numerous minor islands (cf. Zunino & Zullini, 1995) (Figs. 1–4). as well (Blondel et al., 2010). The coastline of these Several factors contribuite to the birth of these islands extends for approximately 18,000 km, in- endemic species: paleogeography (a few have re- cluding 39% of all the Mediterranean coastal areas mained isolated for a long time, others have not), (cf. ANPA, 2001). The islands of the Mediterranean distance from the nearest continental mass (source are, for the most part, strips of land above sea area), size of the island (between a few square which have latterly isolated themselves from the metres and 25,700 square km as is the case of Sicily, nearby continental masses. Other islands are of vol- for instance), height above sea level, substrate, mor- canic origin and have never had contacts with other phology. These factors, as a whole, have determined lands. In the first case we speak about continental the high rates of endemism that have been observed 250 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

in these islands (See ANPA, 2001). Endemic insular Hints on the history of the faunistic explor- species are usually divided into two main categor- ation of circum-sicilian islands ies: abandoned (paleoendemic species) or of new formation (neoendemic species). The first ones are Circum-sicilian islands have, over the centuries, formed by populations in ancient times common on been the destination of several scientific explor- ample areas, successively relegated in confined, ations, thanks to which a remarkable bulk of data separated areas detached from the original contin- has been gathered, which has constituted the basis ental populations. On the contrary, neoendemic spe- for important contributions. For a detailed summary cies are relatively recent taxa, appeared as a result of the naturalistic of Pelagie Islands of the colonisation of the islands in successive and see Baccetti et al. (1995a); for the periods (Zunino & Zullini, 1995). Lo Cascio & Navarra (2003). Par- In this note we propose an analysis of the en- ticularly relevant has been the research coordin- demic fauna, known today, present in the circum- ated by Edoardo Zavattari in 1950 in and sicilian islands. Field and bibliographical research islands, whose results have flowed for carried out have allowed us to identify as a whole the most part in the volume “Biogeografia delle 121 endemic taxa between species and subspecies isole Pelagie” (Zavattari, 1960), still considered a (see Figs. 5, 6). The taxonomical status of certain landmark for the study of biogeography of the entities, however, has not been confirmed by re- circum-sicilian islands (Baccetti et al., 1995b). In cent molecular analysis, or is nevertheless con- particular, Zavattari and his partners found 415 spe- sidered uncertain by several researchers; for this cies of in the two islands. For Pantelleria a reason we have considered appropriate to exclude significant push to entomological research has been a few taxa, considering a total of 111 entities (see given by the Museo Civico di Storia Naturale of notes to Fig. 6). It is nevertheless an extremely im- Venice with three different gathering campaigns led portant value considering that in Sicily, based on between 1983 and 1986, followed by a number of the data reported by Minelli et al. (2005), integ- contributions published by the Museo itself (see rated by the contributions published until 2015 Ratti, 1986). Particularly important, between the (Magrini et al., 2006; Aliquò et al., 2006; Magrini, seventies and the nineties, was the research carried 2007; Sparacio, 2007, 2014; Magrini et al., 2007; out jointly by Palermo University and “Consiglio Magrini et al., 2008; Švihla, 2009; Arnone & nazionale delle Ricerche”. The results to this survey Massa, 2010; Bonavita & Vigna Taglianti, 2010; have constituted the basis to the monograph Baviera, 2010; Baviera & Liberti, 2010; Baviera “Arthropoda di Lampedusa, Linosa e Pantelleria”, & Magnano, 2010; Hertach, 2011; Jordana et al., reporting a whole of 1718 species of (Lo 2011; Kapp, 2010 ; Kleukers et al., 2010; Lo Valvo & Massa, 1995). In particular, to the known Cascio & Sparacio, 2010; Malicky, 2010; Magrini 855 species, another 863 (several of which were et al., 2010a; Magrini et al., 2010b; Rapuzzi & new for science) are added to the checklists repor- Sama, 2010; Stuben, 2010; Bellò & Baviera, 2011; ted in the volume. This work makes the explorative Haitlinger, 2011, 2012; Magrini & Baviera, 2011; level of the three islands satisfactory as a whole, Magrini et al., 2011; Pagliano, 2011; Rigato, 2011; even though not thorough, also due to the fact that Colomba et al., 2012; Giannuzzi-Savelli et al., in the sole Lampedusa island, between 1995 and 2012; Liberto et al., 2012; Müller, 2012; Panta- 2012, another 71 species have been cited (Goggi, leoni & Badano, 2012; Rapuzzi & Sparacio, 2012, 2004; Li Causi et al., 2013). Between 1994 and 2015; Sabella et al., 2012; Gardini, 2013; today, 4 new species have been described for Pan- Lourenço & Rossi, 2013; Magrini et al., 2013; telleria (Pseudomeira cossyrica Pierotti et Bellò, Poggi & Baviera, 2013; Pellizzari, 2013; Arnone 1994; Probaticus cossyrensis Sparacio, 2007; Ech- et al., 2014; Magrini & Paladini, 2014; Baviera, inodera diottii Stuben, 2010; Pseudoapterogyna eu- 2015; Colonnelli & Paladini, 2015; Magrini & phytus lamantiai Sparacio, 2014), 4 for Lampedusa Degiovanni, 2015; Magrini & Paladini, 2015; (Torneuma clandestinum Magnano et Mifsud, Magrini & Uliana, 2015) an overall of 850 en- 2001; Torneuma extinguendum Magnano et Mifsud, demic species are present, 13% of which is ex- 2001; Physetopoda silviae Pagliano, 2011; Neuma- clusive of the circum-sicilian islands. tora annamariae Magrini, Abbazzi et Petrioli, The endemic fauna of the sicilian islands 251

ac- Area Geographic Distance to Endemic Number of Island ronym (km²) Origin coordinates m.s.l.m. mainland and suben- taxa for (km) demic taxa km² Eolie Lat: 38.80º N; ST 12.2 Volcanic Long: 15.25º E 924 54 11 0.9 Lat: 38.63º N; PA 3.4 Volcanic Long: 15.07º E 420 41 10 2.94 Lat: 38.42º N; VU 21 Volcanic Long: 14.98º E 499 20 12 0.57 Lat: 38.45º N; LI 37.3 Volcanic Long: 14.97º E 602 27 14 0.38 Lat: 38.57º N; Salina SA 26.4 Volcanic Long: 14.87º E 962 38 17 0.64 Lat: 38.58º N; FI 9.5 Volcanic Long: 14.58º E 773 45 10 1.05 Lat: 38.55º N; Alicudi AL 5.1 Volcanic Long: 14.37º E 675 53 6 1.18 Lat: 38.72º N; US 8.1 Volcanic Long: 13.20º E 266 51 4 0.49 Egadi Lat: 37.59° N; LE 5.6 Sedimentary Long: 12.20° E 278 13 6 1.07 Lat: 37. 55° N; Favignana FA 19.5 Sedimentary Long:12.19° E 302 8 6 0.25 Lat: 37°58° N; MA 12.3 Sedimentary Long: 12.3° E 686 35 18 1.46 Lat: 36.80º N; Pantelleria PN 83 Volcanic Long: 12.00º E 836 67 20 0.24 Pelagie Lat: 35.88º N; Linosa LN 5.4 Volcanic Long: 12.38º E 195 165 7 1.29 Lat: 35.57º N; Lampione LA 0.036 Sedimentary Long: 12.33º E 36 100 9 250 Lat: 35.52º N; Lampedusa LM 20 Sedimentary Long: 12.62º E 133 120 27 1.35

Table 1. Geographic coordinates of the circum-Sicilian islands, number of known endemic taxa and density.

2013) and 1 for Lampione (Otiorhynchus (- (Pseudomeira aeolica Bellò, Pesarini et Pierotti, michnus) poggii Di Marco, Osella et Zuppa, 2002). 1997; Ocys beatricis Magrini, Cecchi et Lo Cascio, We also have a good overall level of the faun- 2000; Nalassus pastai Aliquò, Leo et Lo Cascio, istic knowledge for the Aeolian Archipelago and for 2006; Anthaxia (Haplantaxia) flaviae Lo Cascio the Island of Ustica, systematically investigated et Sparacio, 2010; Firminus massai Arnone, Lo especially from the sixties with the project “Piccole Cascio et Grita, 2014). Isole” promoted by CNR (Lo Cascio & Navarra, Differently from the other circum-sicilian is- 2003). The conspicuous material gathered has been lands, the Egadi islands have been the subject for published in the conference proceedings to the deeper zoological surveys only starting from the “XVIII Convegno della Società Italiana di Biogeo- end of the sixties; these surveys have been carried grafia”, whose subject was “The and vegetal out by CNR (project “Piccole Isole”) (Reverberi & population of circum-sicilian islands” (AA.VV., Riggio, 1971). The results to these surveys have 1973). In recent times several contributions have mainly pertained some groups of invertebrates updated the knowledge on many Aeolian popula- (Matic, 1968; Focarile, 1969; Strasser, 1969; tions of invertebrates (Gridelli, 1972; Ratti, 1987; Marcuzzi, 1970; Magistretti, 1971; Alicata, 1973; Aliquò 1993, 1995; Lo Cascio & Magrini, 1998; Caruso, 1973; Tamanini, 1973), Amphibians and Cecchi & Lo Cascio, 1999, 2000; Cecchi et al., Rectiles (Bruno, 1970; Lanza, 1973). From that 1999; Arnone et al., 2001; Dapporto & Lo Cascio, moment until today other contributions have been 2001; Lo Cascio et al., 2006) and vertebrates (Lo added, which favoured a widening of the available Cascio, 1994, 2000, 2009, 2010; Deidun et al., knowledge, particularly for tenebrionid beetles 2011; Lo Cascio et al., 2001, 2005; Scalera et al., (Aliquò, 1993, 1995) and terrestrial molluscs 2004) as well as the discovery of a few new species (Beckmann, 1992; 2003; Cianfanelli et al., 2004; 252 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

Fiorentino et al., 2010; Manganelli et al., 2007; detected by the Sicilian Island Award (S.I.A.) as Liberto et al., 2012; 2015), as well as the descrip- valid islands (Islands acknowledged by IOTA regu- tion of some interesting endemic species (the or- lation -www.dxawards.com/Lists/sicilianisawd.htm topheran Acinipe galvagnii Cusimano et Massa, [accessed 24 Agosto 2015]) which as a whole con- 1977; the coleoptera Otiorhynchus (Arammich- stitute approximately 1.11% of all the regional sur- nus) aegatensis Magnano, 1992; Typhloreicheia face (about 285.4 km² on a total of 25,711 km²). For (Typhloreicheia) berninii Magrini, Bastianini et the most part they are rocks or , generally of Petrioli, 2003; Malthinus egadiensis Švihla, scarce naturalistic interest and perimetral to the 2009; Alaocyba ientilei Baviera, 2010; Danacea major islands. In this analysis the 14 biggest islands (Danacea) hierena Baviera et Liberti, 2010; have been taken into consideration (Stromboli, Entomoculia hieratica Poggi et Baviera, 2013; the Panarea, Vulcano, Lipari, Salina, Filicudi, Alicudi, Phasmatodeo Bacillus grandii maretimi Scali et Favignana, Levanzo, Marettimo, Ustica, Pantel- Mantovani, 1990; the mollusc Schileykiella bodoni leria, Lampedusa, Linosa) and the islet of Lam- Cianfanelli, Manganelli et Giusti, 2004. pione, while the Maltese Archipelago has been excluded as administratively not bond to Sicily. These islands can be gathered into three main MATERIAL AND METHODS archipelagoes: the Aeolian Islands (Volcanic), the Egadi (Sedimentary), the Pelagie Islands (both vol- Geographical placement canic and sedimentary); the remaining two islands, Ustica and Pantelleria (both volcanic) are rather The circum-sicilian islands are a totality of 105 isolated (see Fig. 5). All of these islands are charac- (between major islands, islets, rocks and sea stacks) terized by an ample latitudinal extension, running

Figure 1. Acinipe galvagnii from Marettimo; Figure 2. Siciliaria scarificata from Marettimo. Figura 3. Gryllotalpa cossyrensis from Pantelleria. Figure 4. Heliopates avarus donatellae from Pantelleria. The endemic fauna of the sicilian islands 253

from 35° 30'N of Lampedusa, in Pelagie islands, to endemic taxa. It is a complex process, influenced the 38° 50'N of , in the Aeolian by a multiplicity of factors, above all taxa vagility archipelago. The strait of (approximately and paleogeographic evolutio n of the islands. 3 km of length in the narrowest int) currently (Whittaker, 1998). Generally, since the a drop in the separates Sicily from , while the porti on dispersive capacities raises the possibilities of ge- which is closest to the African (), netic isolation, groups of with scarce dis- is about 70 km distant from the island of Pantelleria. persive capacities show higher levels of endemism in confrontation with others with higher dispersive Data gathering and elaboration capacity (Minelli et al., 2005). By observing figure 5, as a confirmation to this To propose a thorough summary we have anal- hypothesis, we can observe the absence of Diptera, yzed the ample bibliography available today on the Odonata, Neuroptera and generally of groups char- circum-sicilian islands. In particular, we have made acterized by a marked vagility among the 111 en- reference to the works included in Zavattari (1960), demic taxa of the fauna of circum-sicilian islands. Francini Corti & Lanza (1973), Massa ( 1995a), Well represented are the Gastropod Molluscs (20 Sparacio (1995, 1997, 19 99), to the management taxa, 18%), whose scarce dispers ive capacity is plans of the natura 2000 sites “Isole Egadi”, “Isole well-known. As it is legitimate to expect, of the 111 Eolie”, ”Isole Pelagie”, “Isola Di Ustica”, “Pantel- taxa taken into consideration, about 50% is made leria” (AAVV, 2009a, b, c, d), as well as to the up by Coleoptera, by far the most ecologically di- dossiers of the checklist of Fauna d’Italia (Minelli versified group on a global level, constituting alone et al., 1993–1995) and to the exquisite though in- over 20% of Italian fauna (Ruffo & Stoch, 2005). complete Ckmap 5.4.1 (Stoch, 2006) (for the list of Among Coleoptera, as in figure 6, the largest fam- the refence from which the taxa distribution has ilies are made up by Tenebrionids (18 taxa, 33%) been drawn, see Fig. 6). As long as possible, we and Curculionids (16 taxa, 30%), two groups char- have consulted the original description of the con- acterised by a strong tendency to insular differen- sidered species bearing in mind the successive chro- tiation (Massa, 1995b). nological, taxonomical and nomenclatural updates. The level of endemism in insular populations, The reference nomenclature follows the checklist furthermore, is positively correlated to surface, hab- of Fauna d’Italia (Minelli et al., 1993–1995), up- itat diversity, age and distance of the island from dated case by case. We have attempted to sum up the continent (Whittaker, 1998). data related to all the metazoans. However, only ad- visory of endemic species referable to Arthropoda, To verify how much these factors influence the Mollusca and Chordata have been taken into con- endemic contingents in the circum-sicilian islands, sideration. Research related to other groups are now the number of endemic taxa of every island has partial and incomplete. Some species present ex- been correlated to clusively in the small islets that are perimetral to the 1) the distance from the nearest continental mass major island have been included among these last (Sicily/Northern ) ones. In particular, Anaspis akaira , known solely 2) maximum height (indirect index of habitat for the Conigli islet, has been included among the diversity) species of Lampedusa; Passalozetes paucesculptus, 3) surface. On the basis of the variable as in fig- known for , among the ones of Panarea. ure 6, a regression has been carried out, using the The data has been organised in a database created Pearson index of correlation. From the results - in a Microsoft Access 2007 environment, and elab- tained, we observe that neither the distance from the orated with Microsoft Excel 2007. continent (r = 0.198) nor height (r = 0.235) are cor- related to the number of endemic taxa (the Pearson coefficient “r” is a measure of the correlation DISCUSSION between the two viariables: it can variate between +1 or -1 and it acquires these extreme levels if the All the insular population, as effectively isol- correlation - positive or negative - is perfect, while ated, show more or less pronounced character- it acquires values close to zero if the two variables istics which, in time, may lead to the formation of are independent). A weak correlation has emerged 254 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

by relating the number of endemic species with the due to the sea depth in this coasta l area), so that area of the islands (Linear function data: y even the colonization of scarcely vagile insects, as =0.1608x + 8.9845; R = 0.3209; Pearson correlation the Tenebrionids, was made easier in these periods index r = 0.54, see figure 8). (Fattorini, 2001). This way, species belo nging to Also, no relations seem to exist between the groups with a high tendency to speciation have had number of endemic species and the geological struc- the possibility to reach the Aeolian islands and ture of the islands (volcanic or sedimentary). Alicudi rapidly differentiate (Fattorini, 2011). This might and Favignana, for instance, show the same number be the case of the disderid spiders, present in Lipari of endemic taxa (6) but they are characterised by a with two endemic taxa - Harpactea aeoliensis profoundly different lithology (see figure 6). Alicata, 1973 and Dysdera flagellifera aeoliensis The complex paleogeographic history (see fur- Alicata, 1973 (Alicata, 1973); of the Curculionid ther on) of the circum-sicilian islands, together with Beetles Otiorhynchus (Arammichnus) meligunensis the dispersive features typical of every taxa, seem Magnano, 1992 and Pseudomeira aeolica Bellò, to have been the most incisive factors determining Pesarini et Pierotti, 1997; of the Blattaria, present the endemic population, and it does not seem pos- with three exclusive specie s (Ectobius aeoliensis sible to hypothesize a unique colonisation and spe- Failla et Messina, 1974, E. filicensis Failla et Mess- ciation model. Quite certainly, the different islands ina, 1974 ed E. parvosacculatus Failla et Messina, (or at least the different archipelagoes), have had 1974) or of the gastropods of the Oxychilus type different population means, whose vicariantist and Fitzinger, 1833, O. (Hyalocornea) alicurensis (Ben- dispersalist models overlap. oit, 1857) of Alicudi and O. (Oxychilus) lagrecai Giusti, 1973 of Filicudi. A likely hypothesis to ex- Considerations on the endemic populations plain the genesis of some endemic species recon- and paleogeography of the circum-sicilian nects to the high degree of environmental instability islands typical of the Aeolian islands: continuous eruptions allegedly determined the nullification of the present Aeolian Archipelago fauna, repeatedly causing “bottleneck” effects, trig- gering and quickening the birth of many of the en- The Aeolian Islands are of relatively recent demic species present on these islands (Lo Cascio formation: the most reliable radiometric dating & Navarra, 2003; Fattorini, 2009; Lo Cascio & estimate that the archipelago formed approximately Sparacio, 2010). This might as well be the origin of 1.3 million years ago (in reference to the disap- some taxa such as Anoxia (Mesanoxia) matutinalis peared apparatuses), while the most ancient rocks moltonii Sabatinelli, 1976, exclusive of Vulcano but above sea level, present in Filicudi, date back to present with the nominal subspecies in the nearby about 600,000 years ago (De Rosa et al., 2004; Lipari and Salina, Anthaxia (Haplantaxia) flaviae Lucchi et al., 2013). They are separate from Sicily Lo Cascio et Sparacio, 2010 known for Panarea, by a sea area which is up to 2000 m. deep and have Salina and Lipari but sympatric in the latter with the always been isolated, even during the marine re- akin A. (Haplantaxia) scutellaris Genè, 1839 prob- gressions in Pleistocene. Considering the relatively ably for a process of “double invasion” (Lo Cascio young age of the archipelago, the high number of & Sparacio, 2010) and especially the LacertidPodar- endemic species found today (30) and its relative cis raffoneae (Mertens, 1952) and its subspecies. In faunistic richness are surprising (cf. Lo Cascio & particular, for P. raffoneae, it would be otherwise Navarra, 2003). Based on this interpretative model, difficult to explain the supposed “antiquity”, con- the entire Aeolian fauna should be of recent ac- sidering that the molecular clocks that have been quisition as entirely formed by propagules of high used for the datations confer it an age between 2 vagility species which rapidly differentiated on the and 13 million years, well before the formation of spot thanks to the well known phenomena of the the “present” Aeolian islands (see Lo Cascio & “bottleneck” and the “founder effect”. We should Navarra, 2003). Its current distribution, limited to as well consider that during the phases of marine a few peripheral islets and Vulcano, is interpreted regression the distance between Sicily and these is- as relictual in the field of an original area which lands was undoubtedly shorter (though not annulled probably involved the whole archipelago; the most The endemic fauna of the sicilian islands 255

part of the populations have allegedly faced local are, as a matter of fact, a fragment of Sicily, to extinctions as a result of the processes of competitive which they alternately remained connected during exclusion derived by the colonisation of the ar- the eustatic variation in Pleistocene, and from chipelago by the P. siculus (Rafinesque- which they are separated by a sea bed only 40 m. Schmaltz, 1814) (Lo Cascio, 2010). deep (Ruggieri, 1973; Agnesi et al., 1993), the last However, to the antique datation of the molecu- time durind Würmian glaciation (until about 12,000 lar clock for P. raffoneae we need to add the diffi- years ago) (Massa, 1973; Ruggieri, 1973). On the culty to explain the presence of terrestrial molluscs contrary, Marettimo originated almost exclusively with high preference for calcium. Among these from sediments that show no similarities to ones Hypnophila incerta (Bourguignat, 1858), the Sicilian as they are correlated to surfa- present in many of these islands, Oxychilus (Hyalo- cing present in northern Africa and Iberian Penin- cornea) alicurensis of Alicudi and O. (Oxychilus) sula (Ruggieri, 1973). Furthermore, it is separated lagrecai of Filicudi. As such, the genesis of Hy- from the two other islands by a channel (“Maret- gromiidae Helicotricha carusoi Giusti, Manganelli timo Channel”) whose maximum depth is 350 m, et Crisci, 1992, is hard to interpret as it belongs to enough to prevent connections with the Sicilian a that is endemic of the Aeolian and implies territory during Pleistocenic regressions (Agnesi et a very long time for its differentiation. al., 1993). These pronounced differences also Giusti (1973) hypothesizes the existence of reflect, as it is right to expect, on faunistic popula- ancient groups of above sea level which tions of the three islands. Marettimo, as pointed out left groups of paleo-endemic species to the new by several authors (Alicata, 1973; Bordoni, 1973; Aelioan islands of formation. These territories Caruso, 1973; Lanza, 1973; Magnano & Osella, might have been situated further north, and even 1973), is characterized by a pre- popula- derived by the fault of Tyrrhenian microplates in tion with predominant similarities with the western their shift towards their present position, or further Mediterranean. Favignana and Levanzo show south, in contact with the Sicilian coast line. poorer endemic populations and generally with A different biogeographical explanation was Sicilian affinities (Canzoneri, 1968). As a whole, 20 proposed for Ocys beatricis Magrini, Cecchi et Lo endemic entities are known in the archipelago, 18 Cascio, 2000: a small carabid, endemic in Lipari of which are present in Marettimo alone. Levanzo and rather isolated in the field of its genus and only has scarcer endemic contingents (6 taxa, only one akin to another specie, Ocys inguscioi Magrini et of which - gastropod Rupestrella rupestris coloba Vanni, 1992, localised in southern Puglia. Con- (Pilsbry, 1918) - exclusive of the island) and Fav- sidered that these are winged species, potentially ignana (6 taxa, all in common with the other islands capable of highly dispersive capacities, it is possible of the archipelago). Two endemic vertebrates are to suppose a climate or ecological change to have present - the Soricid Crocidura sicula aegatensis caused the disappearance in the original distribution Hutterer, 1991, present in all of the three islands, area of a common hypothetical ancestor, and the and the lacertid waglerianus marettimen- distance between the surviving population to have sis (Klemmer, 1956) in Marettimo alone - whose taxo- triggered a differentiation on a species level (Mag- nomic status, besides, is considered rather doubtful rini et al., 2000; Lo Cascio & Navarra, 2003). The (Capula, 1994; Sarà, 1995). Invertebrates make up endemic sub species of garden dormouse present in the largest endemic element under the biogeo- Lipari, Eliomys quercinus liparensis Kahamann, graphical profile. In the field of the endemic fauna 1960, differentiated, according to several authors of Marettimo we need to highlight the presence of (see Angelici et al., 2009) starting from nuclei ori- interesting paleoendemic species, such as the un- ginally introduced in the Roman age for dietary. derground Coleoptera Typhloreicheia (Typhlor- eicheia) berninii Magrini, Bastianini et Petrioli, 2003 Egadi Archipelago and Alaocyba ientilei Baviera, 2010; the Gastropods Oxychilus (Hyalofusca) denatale (Pfeiffer, 1856), The three major islands of the Egadi are of sed- Siciliaria scarificata (Pfeiffer, 1856), Marmorana in- imentary origin, different from the geological and sularis (Benoit, 1857) and Schileykiella bodoni Cian- paleogeographic history. Favignana and Levanzo fanelli, Manganelli et Giusti, 2004; the Tenebrionid 256 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

Coleoptera Odocnemis ruffoi ruffoi (Canzoneri, parently disassociated distribution of the Issid Ho- 1970). This last species belongs to a sub family of moptera Conosimus malfanus Dlabola, 1987, until Tenebrionidae, the Elopinae, which in the Mediter- now known only for Marettimo and Salina. A future ranean area enumerate several elements with a cir- deeper look of the research might reveal the pres- cumscribed geonemy and numerous endemic ence of this species also alongside the northern Si- species. Odocnemis ruffoi has an exclusively insular cilian coastlines. Tyrrhenian distribution: nominal form is found in Marettimo, while another subspecies (ssp. osellai Island of Ustica Gardini, 1979) was described for the Island of , in the . This fact, The Island of Ustica, as the Aeolian Island, is a together with the peculiar, systematic position of talassogenic island. It is the highest tip of a vast un- the species, inspires the hypothesis of an area of dersea volcanic apparatus, whose base is over 2000 relictual distribution, which might have shrunk in meters below the sea level. The intense volcanic comparison to the original one for unspecified explosive activity that took place starting from (maybe ecological) reasons. Both the islands were Pliocene is accountable for the continuous accumu- allegedly refuge-posts for O. ruffoi, while the isol- lation of igneous and pyroclastic material on the sea ation might have determined a successive differen- bed which, in medium Pleistocene (approximately tiation in the two sub-specific forms presently 350 million years ago) led to the emersion of the is- known (AA.VV. 2009a; Aliquò & Soldati, 2010). land (AA.VV., 2009a; Bonomo & Ricci, 2010). The case of Allophylax costatipennis godenigoi Ustica is separated from Sicily by a wide and deep Canzoneri, 1970 is different. The species has a sea area and it has probably always remained isol- northern-african type of distribution and it is present ated, even during pleistocenic regressions. The on the island of Lampedusa (see Aliquò& Soldati, young geological age of Ustica, together with its 2010) while it seems to be lacking in Sicily. The dif- isolation, are the main conditions making its faun- ferentiation of the population of Marettimo at an in- istic populations (prevalently of a Sicilian or south- fraspecific level seems to be due to its geographical ern Italian type) not particularly relevant. (cf. isolation in comparison to the ones of the rest of the Francino Corti & Lanza, 1973). Only 5 endemic area of distribution of the species (AA.VV., 2009a). species, probably all of new formation, are pointed Other important paleoendemic species of Maret- out for the island. The most interesting element is timo are the Isopod Crustaceans Bathytropa ruffoi the Cave Isopod Spelaeoniscus vandeli Caruso, Caruso, 1973 and Spelaeoniscus lagrecai Caruso, 1974, apparently well differentiated by the congen- 1973. The genus Bathytropa Budde-Lund, 1885 en- eric species (Caruso & , 1995). Inter- compasses 8 species diffused in different point esting is the presence of Oxychilus (Hyalocornea) areas in the Mediterranean water basin: a distribu- nortoni (Calcara, 1843), a specie belonging to the tion which, according to Caruso (1973), suggests a subgenus Hyalocornea Monterosato, 1892, in a par- pre-Pliocenic origin. To the hypothetical fragment- ticolar distribution with O. (H.) alicurensis typical ation of the Tyrrhenid of the tertiary period is to be of Alicudi, O. (H.) canini (Benoit, 1843) of north- connected the origin of Spelaeoniscidae Racovitza, western Sicily, O. (H.) egadiensis Riedel, 1973 of 1907 (Caruso, 1973), a family present, for Italy in Favignana and Levanzo and probably also O. (H.?) the sole Sicily with 5 endemic species, 3 of which pomelianus Bourguignat, 1867 of NW- and in the circum-sicilian islands (Argano et al., 1995; the Galite Island in Tunisia (Riedel, 1980). The Caruso & Lombardo, 1995). Other entities which Blattaria Ectobius usticaensis Failla et Messina, are exclusive of the archipelago belong to genera 1974, according to Failla et al. (1973) and Failla & which are rich in point-schizo-endemic species, Messina (1974) who minutely studied the anatomy such as the Coleoptera Otiorhynchus (Aram- of the glandular dimples, allegedly belongs to a dif- michnus) aegatensis Magnano, 1992 (present in all ferent evolutionary line if compared to Ectobius of the three islands), Entomoculia hieratica Poggi Stephens, 1835 Sicilian and Aeolian. Opatrum et Baviera, 2013 (Marettimo), the Chrysomelidae (Colpophorus) validum marcuzzii Canzoneri, 1972 Coleoptera Pachybrachis osellai Daccordi et Ruffo, is part of a Northern African chorotype present, 1975 (present in Levanzo and Marettimo). The ap- other than in insular Sicily with the nominal sub- The endemic fauna of the sicilian islands 257

species, in the islet of Lampione (ssp. rottembergi until the last glaciation they have been connected Canzoneri, 1972) and in Tunisia, Pantelleria and to continental Africa. They therefore own a rich Sardegna (ssp. schlicki Gebien, 1906). amount of species revolving around northern Africa, arrived via land during this period which, Pelagie Islands with the , have successively remained trapped in the two islands. Here they have under- The Pelagie Islands are connected to each other gone more or less marked speciations. Linosa, only from a geographical (and not geological) point moreover, has never had contact with other land, of view. Lampedusa and Lampione are two contin- therefore the origin of its population is to be ental carbonate (Agnesi & Federico, 1995). Differ- searched only on active or passive colonisations ently, Linosa formed between one million and that happened during some hundreds of thousands 500,000 years ago during three different stages of of years. This partially explains the scarcity of spe- volcanic activity (see Tranne, 2002). The difference cies in Linosa in confrontation to Lampedusa (1021 in their birth reflects also on the size of their popu- species, between Molluscs e Arthropods, were lations and on the level of endemism. While Linosa found in Lampedusa, only 349 in Linosa: AA.VV., has only 8 endemic species, Lampedusa has 25; 9 2009d). This faunistic poverty is attributable not are found on Lampione. Lampedusa and Lampione only to the different origin of the islands, but also are the last emerged outpost of the African plate and to their different extension (Lampedusa is 20.2

Figure 5. Number of endemic taxa per island. Figura 6. Number of endemic species Some taxa are present in more than one island. per Coleoptera family

Figure 7. Number of endemic taxa per island. Figure 8. Regression curve for the relationship Island area Some taxa are present in more than one island. in km2 (A)-number of endemic taxa present (E) (see text). 258 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

square km wide, while Linosa is only 5.43) whose ancient connection between Lampedusa and North- calcareous nature seems to offer higher colonization ern Africa, too, derive the Brachyptera possibilities to several species of Arthropods Omocestus lopadusae (La Greca, 1973) and (Massa, 1995b) and land Molluscs. This last point Pamphagus ortolaniae Cusimano & Massa, 1977 is evident if we consider that on 5 endemic taxa (Massa, 1995b; Massa, 2011) and the Buprestid present on the Italian island in the - Julodis onopordi lampedusanus Tassi, 1966. Lampedusa lopadusae lopadusae (Calcara, 1846) Particularly interesting under the biogeographical [Lampedusa], L. lopadusae nodulosa Monterosato, profile is Leptotyphlopsis lopadusae Bordoni, 1973, 1892 [Lampione], Oxychilus (Oxychilus) diductus an underground Staphylinidae belonging to a group (Westerlund, 1886) [Lampedusa], Trochoidea cumiae revolving around the Northern Mediterranean but (Calcara, 1847) [Lampedusa and Linosa], Cernuella also present is Tunisia, and well differentiated both metabola (Westerlund, 1889) [Lampedusa] - none from African and Italian congeners. According to is present in Linosa and Pantelleria. The paleogeo- Bordoni (1973) it is a species belonging to a very graphic vicissitudes lead to the inevitable conclu- old phyletic lineage, which differentiated after the sion that the endemic fauna of Lampedusa and climate changes of the Quaternary. Lampione is prevalently relictual, while that of For the terrestrial molluscs, it is of particular Linosa is invasive. In both cases, they are neo- biogeographical importance the presence of the endemic species of recent formation, evolved from genus Lampedusa Boettger, 1877 including L. species characterized by a high colonizing capa- lopadusae (Calcara, 1846) endemic of Lampedusa city and a marked evolutionary speed. This phe- island, L. lopadusae nodulosa Monterosato, 1892 nomenon is observed, as formerly said, in Tenebri- endemic of Lampione island, L. imitatrix Boettger, onids, present in the islands with 9 endemic species 1877 e L. melitensis (Caruana-Gatto, 1892) en- (see figure 5) on 37 known ones, with a rate of demic of Maltese Island. Lampedusa is akin to endemism of 24.3% (Lo Cascio, 2002). In particu- Muticaria Lindhol, 1925 of South-eastern Sicily lar, the presence in Lampione of 4 endemic taxa, and Maltese Islands. Both these genera have affin- with 2 exclusive subspecies Opatrum (Colpo- ities with species of groups which are originated in phorus) validum rottenbergi Canzoneri, 1972, the and in north-, Alphasida puncticollis moltonii Canzoneri, 1972 - and colonized these more western territories prob- and a species being described (Tentyria n. sp., see ably during Messinian Age (Giusti et al., 1995). Lo Cascio & Pasta, 2012), is symptomatic both of Here, the only endemic vertebrate is Podarcis the differentiation speed of some species of this filfolensis laurentiimulleri (Bedriaga, 1876) which, family from the founding population and of their according to recent molecular and biochemical capacity to colonise isolated and insular territories survey, differentiated from stocks of populations of (Aliquò, 1995). The insular differentiation is Podarcis sicula which colonised Pelagie Islands certainly a rather quick phenomenon also among and Maltese Islands during pleistocenic regressions Curculionoidea if, as Osella & Riti (1995) have ob- (see La Mantia & Lo Cascio, 2008; Sciberras & served, they are present in Pelagie with 9 endemic Schembri, 2008 - see also note 16 of figure 6) species. Of these, 1 species from Lampedusa (Torneuma clandestinum) and 1 of Linosa (Otio- Island of Pantelleria rhynchus (Arammichnus) linussae A. Solari et F. Solari, 1922) have connections with Tyrrhenian Pantelleria is a volcanic island, emerged approx- species, while 4 of Lampedusa (Alaocyba lampedu- imately 324,000 years ago, and since then it has sae Dodero, 1916; Neumatora annamariae (Mag- never had contacts with emerged land (Agnesi & rini et al., 2013; Torneuma extinguendum; Federico, 1995). The most relevant event, which de- Otiorhynchus (Arammichnus) lopadusae A. Solari termined the actual faunistic composition, was the et F. Solari, 1922), 1 of Linosa (Chiloneus (Chi- eruption that about 45 thousand years ago entirely loneus) solarii Pesarini, 1970) and 1 of Lampione, covered the island with a layer of stone 5 metres Otiorhynchus (Arammichnus) poggii Di Marco, thick (“ ignimbrite”). It is highly likely that Osella et Zuppa, 2002, show northern African af- this destroyed the most part of the existent flora and finities. From relictual populations witnessing the fauna. Most of the endemic species of Pantelleria The endemic fauna of the sicilian islands 259

(20 taxa pointed out) are then neoendemic species, mic of Pantelleria, Apterola (Apterola) kuenckeli which differentiated in relatively recent times start- focarilei Tamanini, 1964. The biogeographical in- ing from some founding propagules that re-coloni- terpretation of the Oedemerid Coleoptera Sten- zed the island following two main lines: from Sicily ostoma cossyrense , 1995. This species and Northern Africa (cf. Francini Corti & Lanza, belongs to a genus that includes 3 endemic species 1973; Massa, 1995b). Endemic species akin to found, other than in Pantelleria, in Madera and Northern African species are allegedly the Isopod Maltese Islands, as well as a very diffused Western- Spelaeoniscus vandeli Caruso, 1974, the Orthoptera Mediterranean-Atlantic species. It is likely, then, Gryllotalpa cossyrensis Baccetti et Capra, 1978 that it belongs to a very ancient genus, with paleo- (Baccetti et al., 1995a), the Buprestid Beetle mediterranean distribution, whose original area Acmaeodera bipunctata romanoi Sparacio, 1992 fragmented more recently in the present islands (Sparacio, 1992; Sparacio & Ratti, 1995), the where it rapidly evolved for the founder principle Curculionid Beetle Alaocyba separanda Dodero, (Bologna, 1995; Massa, 1995b). Hardly explainable 1916 (cf. Massa, 1995b) and Echinodera diottii is also the presence of Leptanilla poggii Mei, 1995, (Stuben, 2010), and Melolonthid Coleoptera a hymenoptera formicidae included in a genus with Pseudoapterogyna euphytus lamantiai (Sparacio, scarce dispersive capacities. The most likely hypo- 2014). The Curculionid Beetles Otiorhynchus thesis is, according to Mei (1995), the introduction (Arammichnus) cossyrensis Magnano, 1992 e for anthropic cause from Northern Africa in an un- Pseudomeira cossyrica (Osella & Riti, 1995) have specified moment of the recent history of the island. uniquely Tyrrhenian affinities. The Pselaphid To anthropic reasons, too, is connected the presence Tychomorphus cossyrensis (Dodero, 1919) is part of the Muridae mammal Apodemus sylvaticus of a genus of strictly West-Mediterranean diffusion hermani Felten et Storch, 1970 and of Crocidura (Poggi, 1995), while to a species with Mediter- pachyura cossyrensis Contoli, 1990 (Sarà & Zanca, ranean Geonemy belongs the only heteropter ende- 2008; Angelici et al., 2009).

Figure 1. Aeolian Archipelago: Panarea Island. 260 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

Figure 2. Aeolian Archipelago: Salina Island, Pollara.

Figure 3. Egadi Archipelago: Favignana Island with Levanzo Island in the background. The endemic fauna of the sicilian islands 261

Figure 4. Egadi Archipelago: Levanzo Island, Cala Minnola.

Figure 5. Island of Ustica, Mount Guardia dei Turchi. 262 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

Figure 6. Island of Ustica, Cala Sidoti and Punta Spalmatore.

Figure 7. Island of Pantelleria, Montagna Grande. The endemic fauna of the sicilian islands 263

Figure 8. Island of Pantelleria, Lake of Venus.

Figure 9. Island of Pantelleria. 264 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES A R T H R O P O D A ARACHNIDA ARANEAE DYSDERIDAE Dysdera flagellata Grasshoff, 1959*; Grasshoff, 1959 X Pesarini, 1995; Pantini & Isaia, 2015 Dysdera flagellifera X Alicata, 1973*; aeoliensis Alicata, 1973 Pantini & Isaia, 2015 Harpactea aeoliensis X Alicata, 1973*; Alicata, 1973 Pantini & Isaia, 2015 SALTICIDAE

Aelurillus lopadusae X Cantarella T., 1983*; Cantarella, 1983* Azarkina & Loguov, 2006; Pantini & Isaia, 2015 ORIBATIDA PASSALOZETIDAE Passalozetes paucesculptus X Bernini, 1973* Bernini, 1973 PSEUDOSCORPIONIDA CHTHONIDAE Chthonius Callaini, 1989*; (Ephippiochthonius) X Stoch, 2006; aegatensis Callaini, 1989* Gardini, 2013 HEXAPODA BLATTARIA ECTOBIDAE Ectobius aeoliensis Failla et al., 1973*; Failla et Messina, 1974 X X X Failla & Messina, 1974; Stoch, 2006 Ectobius filicensis X Failla & Messina, 1974*; Failla et Messina, 1974 Stoch, 2006 Ectobius parvosacculatus Failla et al., 1973*; Failla et Messina, 1974 X Failla & Messina, 1974; Stoch, 2006 Ectobius usticaensis Failla et al., 1973*; Failla et Messina, 1974 X Failla & Messina, 1974; Stoch, 2006 COLEOPTERA BUPRESTIDAE Acmaeodera bipunctata X Sparacio & Ratti*, 1995; romanoi Sparacio, 1992 Stoch, 2006 Anthaxia (Haplantaxia) Lo Cascio et al., 2006*; flaviae Lo Cascio et X X X Lo Cascio & Sparacio, Sparacio, 2010 2010 Julodis onopordi X Tassi, 1966*; Sparacio & lampedusanus Tassi, 1966 Ratti, 1995; Stoch, 2006 CANTHARIDAE

Malthinus egadiensis X Švihla, 2009* Švihla, 2009 CARABIDAE

Carabus morbillosus X Rapuzzi & Sparacio, lampedusae Born, 1925 2015 Ocys beatricis Magrini, X Magrini et al., 2000*; Lo Cecchi et Lo Cascio, 2000 Cascio & Navarra, 2003 Typhloreicheia berninii Magrini et al., 2003* Magrini, Bastianini et X Petrioli, 2003

Table 1/1. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. The endemic fauna of the sicilian islands 265

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES CHRYSOMELIDAE

Pachybrachis osellai X X Daccordi & Ruffo, Daccordi et Ruffo, 1975 1975*; Stoch, 2006

Scymnus () caprai X Canepari, 1983*, 1995 Canepari, 1983

Alaocyba ientilei X Baviera, 2010* Baviera, 2010 Alaocyba lampedusae X Dodero, 1916*; Osella & Dodero, 1916 Riti, 1995; Stoch, 2006; Alaocyba separanda X Dodero 1916*; Osella & Dodero, 1916 Riti, 1995; Stoch, 2006 Chiloneus (Chiloneus) X Pesarini, 1970a, b*; solarii Pesarini, 1970 Osella & Riti, 1995 Echinodera diottii X Stuben, 2010* Stuben, 2010 Neumatora annamariae Ma- X Magrini et al., 2013* grini, Abbazzi et Petrioli, 2013 Otiorhynchus (Arammichnus) X X X Magnano, 1992*; aegatensis Magnano, 1992 Baviera & Magnano, 2010; Stoch, 2006 Otiorhynchus (Arammichnus) Solari & Solari, 1922a*; cossyrensis Magnano, 1992 X Magnano, 1992; Stoch, 2006 Otiorhynchus (Arammichnus) Solari & Solari, 1922b*; linussae Solari et Solari, 1922 X Magnano 1992; Baviera & Magnano, 2010 Otiorhynchus (Arammichnus) Solari & Solari, 1922a*; lopadusae Solari et Solari, X Magnano, 1992; 1922 Stoch, 2006 Otiorhynchus (Arammichnus) Solari & Solari, 1922a*; meligunensis Magnano, 1992 X X X X X X X Magnano, 1992; Stoch, 2006 Otiorhynchus (Arammichnus) Di Marco et al., 2002*; poggii Di Marco, Osella et X Lo Cascio & Pasta, 2012 Zuppa, 2002 Pseudomeira cossyrica Pierotti & Bellò, 1994*; Pierotti et Bellò, 1994 X Osella & Riti, 1995; Stoch, 2006 Pseudomeira aeolica Bellò, Bellò et al., 1997*; Pesarini et Pierotti, 1997 X X X X X Stoch, 2006; Bellò & Baviera, 2011 Torneuma clandestinum Osella & Riti, 1995; Magnano et Mifsud, 2001 X Magnano & Mifsud, 2001; Stoch, 2006 Torneuma extinguendum Osella & Riti, 1995; Magnano et Mifsud, 2001 X Magnano & Mifsud, 2001; Stoch, 2006 MELOLONTHIDAE

Anoxia (Mesanoxia) matutina- X Sabatinelli, 1976; Lo lis moltonii Sabatinelli, 1976 Cascio & Navarra, 2003 Firminus massai Arnone, X X X X Arnone et al., 2014* Lo Cascio et Grita 2014 Pseudoapterogyna euphytus X Ragusa, 1875*; lamantiai Sparacio, 2014 Sparacio, 2014

Table 1/2. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. 266 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES

MELYRIDAE

Danacea (Allodanacaea) X Liberti, 1995 caneparii Liberti, 1985 Danacea (Danacea) hierena X X X Baviera & Liberti, 2010* Baviera et Liberti, 2010 MORDELLIDAE

Mordellistena (Mordellistena) X Franciscolo, 1991*; irritans Franciscolo, 1991 Massa, 1995 SCRAPTIIDAE Anaspis (Larisia) akaira Franciscolo, 1991*; Franciscolo, 1991 X Lo Cascio et al., 2002; Massa, 1995a, b SCYDMAENIDAE

Pseudoeudesis sulcipennis X Massa, 1995a, b; lampedusae Binaghi, 1948 Sparacio, 1995 STAPHYLINIDAE

Entomoculia hieratica X Poggi & Baviera, 2013* Poggi et Baviera, 2013 Leptotyphlopsis lopadusae X Bordoni, 1973*; Massa, Bordoni, 1973 1995a, b; Sparacio, 1995 Tychomorphus cossyrensis X Sparacio, 1995 (Dodero, 1919) TENEBRIONIDAE Alphasida (Glabrasida) Canzoneri, 1972*; puncticollis moltonii X Aliquò & Soldati, 2010; Canzoneri, 1972 Stoch, 2006 Alphasida (Glabrasida) Canzoneri, 1972; puncticollis tirellii X Aliquò & Soldati, 2010; Leoni, 1929 Stoch, 2006 Allophylax costatipennis X Canzoneri, 1970*; godenigoi Canzoneri, 1970 Aliquò & Soldati, 2010 (Asida) minima X Aliquò & Aliquò 2000; Reitter, 1917 Stoch, 2006 Erodius (Erodius) audouini X Aliquò & Soldati, 2010 destefanii Failla Tedaldi, 1887 Heliopathes avarus X Ragusa, 1897; Canzoneri, donatellae Canzoneri, 1970 1968; Aliquò & Soldati, 2010 Machlopsis doderoi Gridelli, 1960; Gridelli, 1930 X X Aliquò & Aliquò, 2000; Osella & Riti, 1995 Nalassus pastai Aliquò, X Aliquò et al., 2006* Leo et Lo Cascio, 2006 Odocnemis ruffoi ruffoi X Canzoneri, 1970*; (Canzoneri, 1970) Aliquò, 2010 Opatrum (Colpophorus) Riggio, 1885*; Gridelli, validum marcuzzii X 1960, Aliquò & Soldati, Canzoneri, 1972 2010 Opatrum (Colpophorus) Canzoneri, 1972; validum rottenbergi X Goggi, 2004 Canzoneri, 1972 Pachychila (Pachychilina) Canzoneri, 1972; Aliquò, dejeani doderoi Peyerimhoff, X 2010; Stoch, 2006 1927

Table 1/3. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. The endemic fauna of the sicilian islands 267

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES

TENEBRIONIDAE

Phaleria (Phaleria) bimacu- X X X X Aliquò, 1993; Marcuzzi, lata marcuzzii Aliquò, 1993 1996; Aliquò, 2010 Probaticus (Pelorinus) X Sparacio, 2007 cossyrensis Sparacio, 2007 Stenosis brignonei X X Aliquò & Soldati, 2010; Koch, 1935 Stoch, 2006; Tentyria grossa X X Canzoneri, 1972; Aliquò, sommieri Baudi, 1874 2010; Stoch, 2006; Tentyria grossa X Canzoneri, 1972; angustata (Kraatz, 1896) Aliquò, 2010 Trachyscelis aphodioides X Failla, 1886*; Luigioni, lopadusae Koch, 1935 1929; Goggi, 2004 COLLEMBOLA ENTOBRYIDAE Pseudosinella aeolica X Dallai, 1973* Dallai, 1973 Seira dagamae Dallai, 1973 X X X Dallai, 1973* ISOTOMIDAE

Folsomides meridionalis X X X X X X X Dallai, 1973* Dallai, 1973 NEANURIDAE Friesea lagrecai Dallai, 1973 X X X X X X Dallai, 1973* ONYCHIURIDAE

Onychiurus lampedusae X Dallai, 1973* Dallai, 1978 HETEROPTERA LYGEIDAE Apterola (Apterola) kuenckeli X Tamanini, 1964*; focarilei Tamanini, 1964 Carapezza, 1995 Plinthisus (Isioscytus) Carapezza, 1995 minutissimus meridionalis X Mancini, 1935* MIRIDAE Tuponia (Chlorotuponia) Tamanini, 1973*; hippophaes liparensis X X Ippolito, 1986 Tamanini, 1973* Phytocoris (Ktenocoris) X X Carapezza, 1995* cossyrensis Carapezza, 1995 HOMOPTERA CICADELLIDAE Adarrus aeolianus X X D’Urso, 1984*; D’Urso, 1984 Stoch, 2006 ISSIDAE

Conosimus malfanus X X Lo Cascio & Pasta, 2004 Dlabola, 1987 HYMENOPTERA FORMICIDAE Leptanilla poggii Mei, 1995 X Mei, 1995*

Tetramorium pelagium X Mei, 1995* Poldi in Mei, 1995

Table 1/4. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. 268 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES MUTILLIDAE Physetopoda silviae X Pagliano, 2003*, 2011 Pagliano, 2011 LEPIDOPTERA SATYRIDAE Hipparchia leighebi Kudrna, 1976*; Kudrna, 1976 X X X X X Kudrna & Leigheb, 1988; Stoch, 2006 ORTHOPTERA ACRIDIDAE Omocestus lopadusae X Baccetti et al., 1995a, b; (La Greca, 1973) Stoch, 2006; Massa, 2011 GRYLLOTALPIDAE Gryllotalpa cossyrensis X Baccetti & Capra, 1978; Baccetti et Capra, 1978 Baccetti et al., 1995a, b; Stoch, 2006; Massa, 2011 Acinipe galvagnii Cusimano & Massa, Cusimano et Massa, 1977 X X X 1977*; Stoch, 2006; Massa, 2011 Pamphagus ortolaniae Cusimano & Massa, Cusimano et Massa, 1977 X 1977; Stoch, 2006; Massa, 2011 PHASMIDA BACILLIDAE Bacillus grandii maretimi X Berni, 1996 Scali et Mantovani, 1990 MALACOSTRACA ISO- PODA ARMADILLIDAE Armadillidium hirtum X Caruso & Lombardo, pelagicum Arcangeli, 1955 1995 BATHYTROPIDAE Bathytropa ruffoi X Schmalfuss, 2003 Caruso, 1973 SPELAEONISCIDAE Spelaeoniscus costai X Caruso & Lombardo, Caruso et Lombardo, 1976 1995 Spelaeoniscus lagrecai X Caruso & Lombardo, Caruso, 1973 1995 Spelaeoniscus vandeli X Caruso & Lombardo, Caruso, 1974 1995 C H O R D A T A MAMMALIA INSECTI- VORA SORICIDAE Crocidura sicula aegatensis Hutterer, 1991; Angelici Hutterer, 1991* X X X et al., 2009; Sarà, 1995; Stoch, 2006 Crocidura pachyura X Angelici et al., 2009; cossyrensis Contoli, 1990 Stoch, 2006 RODENTIA GLIRIDAE Eliomys quercinus liparensis X Angelici et al., 2009; Kahamann, 1960 Stoch, 2006 MURIDAE Apodemus sylvaticus hermani X Angelici et al., 2009; Felten et Storch, 1970 Stoch, 2006

Table 1/5. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. The endemic fauna of the sicilian islands 269

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES REPTILIA Podarcis filfolensis laurent- X X X Capula,1994; Sindaco mulleri Fejervari, 1924 et al., 2006; Stoch, 2006 Podarcis raffoneae alvearioi X X X Capula, 2006; Sindaco (Mertens, 1955) et al., 2006; Stoch, 2006 Podarcis raffoneae X Capula, 2006; Sindaco antoninoi (Mertens, 1952)* et al., 2006; Stoch, 2006 Podarcis raffoneae X Capula, 2006; Sindaco raffoneae (Mertens, 1952) et al., 2006; Stoch, 2006 Podarcis raffoneae X Capula, 2006; Sindaco cucchiarai Di Palma, 1980* et al., 2006; Stoch, 2006 Podarcis sicula liscabiancae Corti & Lo Cascio, 1999; (Mertens, 1952)* X Sindaco et al., 2006; Stoch, 2006 Podarcis sicula trischittai Corti & Lo Cascio, 1999; (Mertens, 1952)* X Sindaco et al., 2006; Stoch, 2006 Podarcis waglerianus maret- Lo Cascio & Pasta, 2008; timensis (Klemmer, 1956)* X Sindaco et al., 2006; Stoch, 2006 SQUAMATA SCINCIDAE Chalcides ocellatus Corti & Lo Cascio, 2002; linosae Boulenger, 1920* X Sindaco et al., 2006; Stoch, 2006 Chalcides ocellatus Corti & Lo Cascio, 2002; zavattarii Lanza, 1954* X Sindaco et al., 2006; Stoch, 2006 M O L L U S C A ARCHITAENIOGLOSSA COCHLOSTOMATIDAE Cochlostoma paladilhianum X Manganelli et al., pirajnaea (Benoit, 1878) 1995; , 2011 CLAUSIILIDAE Lampedusa lopadusae X Calcara, 1846*; lopadusae (Calcara, 1846) Liberto et al., 2010 Lampedusa lopadusae X Liberto et al., 2010; nodulosa Monterosato, 1892 Lo Cascio & Pasta, 2012 Siciliaria (Siciliaria) X Liberto et al., 2015 scarificata (Pfeiffer, 1857) CHONDRINIDAE

Rupestrella rupestris coloba X Beckmann, 2002 (Pilsbry, 1918)

Hypnophila emiliana X X Liberto et al., 2010 (Bourguignat, 1858) Hypnophila incerta X X X X X Giusti, 1973; (Bourguignat, 1858) Liberto et al., 2010

Marmorana (Murella) muralis X Calcara, 1846 frivaldszkyi (Calcara, 1846) Marmorana (Murella) muralis X Fiorentino et al., 2008a, b insularis (Benoit, 1857)

Table 1/6. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables. 270 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

TAXON FA LE MA US LI VU ST SA AL FI PA LM LN LA PN REFERENCES HYGROMIIDAE Cernuella metabola Bank, 2011; Manganelli (Westerlund, 1889) X et al., 1995 Helicotricha carusoi Giusti, Giusti et al., 1992 Manganelli et Crisci, 1992 X X X X X X X Schileykiella bodoni Cianfanelli et al, 2004 Cianfanelli, Manganelli X et Giusti, 2004 Trochoidea cumiae Cianfanelli, 2002 (Calcara, 1847) X X LIMACIDAE Limax aeolianus Giusti, 1973 Giusti, 1973; Lo Cascio X X & Navarra, 2003 ZONITIDAE Oxychilus (Hyalocornea) Benoit, 1857-1862*; alicurensis (Benoit, 1857) X Giusti, 1973 Oxychilus (Hyalocornea) Manganelli et al., 1995 egadiensis Riedel, 1973 X X Oxychilus (Hyalocornea) Calcara, 1843; nortoni (Calcara, 1843) X Liberto et al., 2010 Oxychilus (Hyalofusca) Manganelli et al., 2007 denatale (Pfeiffer, 1856) X Oxychilus (Oxychilus) Giusti, 1973; diductus (Westerlund, 1886) X Corti et al., 2002 Oxychilus (Oxychilus) Giusti, 1973* lagrecai Giusti, 1973 X

Table 1/7. Endemic taxa of circum-sicilian island listed by alphabetic order and relative distribution. For the abbreviation of the islands see Table 2. The species followed by * have not been taken into consideration for the elaboration of the Tables.

CONCLUSIONS posed by Fattorini (2011) who took into considera- tion only Tenebrionids: The study of the populations of the circum- - in the case of the Aeolian Islands, the new en- sicilian islands, as we have seen, is particularly demic species might have originated by propagules complex (see also Francini Corti & Lanza, 1973 arrived from Sicily especially during the periods of and Massa et al., 2011), as these islands vary sub- marine regression, when the distance between these stantially for their origin (volcanic or sedimentary), islands and Sicily reduced but not annulled; these paleogeography (some have been connected to propagules might have rapidly differentiated due to Sicily or the African continent during pleistocenic a marked “bottleneck”, accentuated by the volcanic regressions, other have remained isolated), distance instability of the area. The origin of paleoendemic from the main source of colonisation (Sicily or species is more complex: their genesis is allegedly Africa), surface (, the biggest island, has a to be found in the complex geological history or surface of 245.7 km2 but most of the islands are “paleo-Aeolian islands” smaller than 30 km2) and environmental conditions. - for the Egads we can suppose a substantial col- Besides, their position at the border between onisation via land for Levanzo and Favignana, and Africa makes their faunistic composition a mo- while the populations of Marettimo have a preval- saic of European and African elements (Francini ently relictual connotation. Corti & Lanza, 1973; Massa 1995b, 2011) with im- - Ustica, Linosa and Pantelleria, of volcanic ori- portant implications of preservation (Fattorini, gin, are very distant from continental areas, with 2008, 2011). which they would never have gotten into contact, The conclusions that we draw by analysing as a which can explain the fact that they show, almost whole the endemic contingents of circum-sicilian exclusively, endemic species of new formation. islands and the main factors that have determined -Lampedusa and Lampione are very isolated and the insular differentiation are similar to those pro- of ancient origin; we can therefore presume that The endemic fauna of the sicilian islands 271

their endemic contingent derives substantially from Alicata P., 1973. I Dysderidae (Araneae) delle Eolie, a relictual population, above all for the endemic delle Egadi e di Ustica. Lavori della Società italiana genera (as in Lampedusa), and the oldest species to di Biogeografia, n.s., 3 [1972]: 341–353. which other ones (arrived during quaternary Aliquò V., 1993. Dati nuovi e riassuntivi sui Coleotteri contacts with Northern Africa and differentiated Tenebrionidi delle isole circumsiciliane (Coleoptera: during more recent times) have added. Tenebrionidae). Il Naturalista siciliano, 17: 111–125. Lastly, we need to consider that all of the cir- Aliquò V., 1995. Nuovi reperti di Tenebrionidae delle cum-sicilian islands have, more or less intensely, isole circumsiciliane (Insecta Coleoptera). Il Natur- alista siciliano, 19: 131. undergone profound changes in their natural asset Aliquò V. & Aliquò A., 2000. Terzo contributo alla revi- due to anthropic impact, particularly with the de- sione della collezione coleotterologica di Enrico struction of most of their original woods. This, pre- Ragusa: Tenebrionidae (Coleoptera). Il Naturalista sumably, has led to the extinction of some taxa and siciliano, 24: 103–144. to the high rarefaction of others. Aliquò V. & Soldati F., 2010. Coleotteri Tenebrionidi di The ecological and biogeographical importance Sicilia (Insecta: Coleoptera Tenebrionidae). Mono- of point endemic species is proportional to their fra- grafie Naturalistiche, 1. Edizioni Danaus, Palermo, gility, therefore we believe that it is particularly im- 176 pp. portant and urgent to adopt specific protection Aliquò V., Leo P. & Lo Cascio P., 2006. I Tenebrionidi measures such as the already mentioned Direttiva dell’Arcipelago Eoliano: nuovi dati faunistici e linea- 92/43 CEE (“Direttiva habitat”): “they are endemic menti zoogeografici con descrizione di una nuova and need particular attention, considered the spe- specie di Nalassus Mulsant, 1854 (Coleoptera, cificity of their habitat and/or the potential incid- Tenebrionidae). Il Naturalista siciliano, 30: 69–90. Amori G., Angelici F.M., Frugis S., Gandolfi G., ence of its exploitment on their state of Groppali R., Lanza B., Relini G. & Vicini G., 1993. conservation”. Vertebrata. In: Minelli A., Ruffo S. & La Posta S. (Eds.). Checklist delle specie della fauna italiana, 110 fasc. Ed. Calderini, Bologna. REFERENCES Angelici F.M., Laurenti A. & Nappi A., 2009. A checklist of the Mammals of small italian islands. Hystrix AA.VV., 1973. Il popolamento animale e vegetale delle Italian Journal of Mammology (n.s.), 20: 3–27. isole circumsiciliane. Lavori della Società italiana di ANPA, 2001. La Biodiversità nella regione biogeografica Biogeografia, n.s., 3 [1972], 920 pp. mediterranea. Agenzia Nazionale per la Protezione AA.VV., 2009a. Indagini faunistiche. Pg 27-30. In: dell’Ambiente, Roma, 144 pp. Cavallaro F. (coord.), Piano di gestione “Isole Eolie”. Argano R., F., Guglielmo L., Riggio S. & Ruffo Provincia regionale di Messina, 143 pp. S., 1995. Checklist delle specie della fauna italiana. AA.VV., 2009b. Descrizione biologica del sito. Pp. 24– 30. Crustacea Malacostraca II. Calderini, Bologna, 97. In: Bagliani P. (coord.), Piano di gestione del 52 pp. SIC/ZPS “Isola di Ustica”. Provincia regionale di Arnone M., Lo Cascio P. & Nistri A., 2001. I popola- Palermo, 435 pp. menti a delle Isole Eolie (Mar Tirreno) AA.VV., 2009c. Descrizione biologica del sito - Aspetti (Insecta Coleoptera). Il Naturalista siciliano, 25: 109– botanici, ornitologici e zoologici. Pg. 37–145. In: 138. Russo D. G. (coord.), Piano di gestione del sito NATURA 2000 “Isole Egadi”. Provincia regionale di Arnone M., Lo Cascio P. & Grita F. 2014. Un nuovo , 605 pp. Firminus delle Isole Eolie (Coleoptera Melolonthidae AA.VV., 2009d. Descrizione faunistica dei siti. Pp. 156– Rhizotroginae). Il Naturalista Siciliano, 28: 339–354. 248. In: Nicolini G, Dimarca A., Casamento G. & Arnone M. & Massa B., 2010. A new species of Bol- Livreri Console S. (coord.), Piano di gestione “Isole belasmus Boucomont, 1911 (Insecta Coleoptera Geo- Pelagie”, Parte I - Fase conoscitiva. Provincia re- trupidae) from Sicily (Italy). Il Naturalista siciliano, gionale di , 486 pp. 24: 401–414. Agnesi V., Macaluso T., Orrù P. & Ulzega A., 1993. Azarkina G.N. & Logunov D.V., 2006. Taxonomic notes Paleogeografia dell’Arcipelago delle Egadi (Sicilia) on Aelurillus species of the western Mediter- nel Pleistocene superiore - Olocene. Il Naturalista ranean (Araneae: Salticidae). Bulletin of the British siciliano, 17: 3–22. Arachnological Society, 13: 233–248. Agnesi V. & Federico C., 1995. Aspetti geografico-fisici Baccetti B. & Capra F. 1978. Notulae Orthopterologicae e geologici di Pantelleria e delle Isole Pelagie (Canale XXXIV: Le specie italiane del genere Gryllotalpa. di Sicilia). Il Naturalista siciliano, 19 (Suppl.): 1–22. Redia, 61: 401–464. 272 CALOGERO MUSCARELLA & ALESSANDRO BARAGONA

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