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TABLE 1. Geographical variables for the circum-Sicilian islands. S – number of species; A – Area (km2); Ds – Distance to Sicily
(km); Da – Distance to North Africa (km); Di – Distance to the nearest island (km); SDs – Sea depth to Sicily (m below sea level);
SDa – Sea depth to Africa (m below sea level); SDi – Sea depth to the nearest island (m below sea level). X1, X2 and X3 are the three
dimensions extracted by Non-metric Multidimensional Scaling from a matrix of inter-island faunal dissimilarity (Kulczinski 2
index).

Island                SA       Ds   Da Di SDs                            SDa   SDi     X1      X2      X3
                                                                         1500   10   –0.306  –0.049  0.075
1. Lipari             32 37.29 28   370 1 780                            1500  366   –0.392  0.169   –0.178
                                                                         1500   10   –0.181  0.248   0.073
2. Salina             24 26.38 39   365 4 780                            1500  1281  –0.255  0.326   –0.330
                                                                         1500  1281  –0.629  –0.038  0.416
3. Vulcano            22 20.87 21   375 1 780                            1500  1281  –0.562  0.040   0.220
                                                                         1500  600   –0.582  –0.033  –0.010
4. Stromboli          25 12.19 58   405 18 1200                          1500   80   –1.004  0.603   0.339
                                                                         1500   50   –0.917  0.290   –0.152
5.Filicudi            15 9.49 46    345 15 1500                          1500   50   –0.831  –0.032  –0.102
                                                                         1500   10   –0.737  –0.036  –0.628
6. Alicudi            18 5.1   53   330 15 1500                          1500   20   –0.283  –0.089  –0.596
                                                                         1500  1500  –0.334  –0.416  0.326
7. Panarea            22 3.34 42    390 14 780                           400    42   –0.108  –0.468  0.484
                                                                         400    42   0.001   –0.410  0.326
8. Basiluzzo          3 0.29 43.5   394 3.5 780                          400   145   0.314   –0.322  1.046
                                                                         500   500   0.292   –1.255  0.066
9. Lisca Bianca       4 0.0413 42   390 3 780                            400   400   0.247   –1.021  –0.675
                                                                         100   100   0.234   1.762   –0.285
10. Bottaro           7 0.0073 42   390 2.75 780                         100   400   0.646   –0.527  –0.879
                                                                         500   100   0.871   0.018   0.144
11. Scoglio Faraglione 4 0.0049 39  365 0.27 780                         500   100   1.121   0.015   0.033
                                                                         500   100   0.961   0.417   –0.441
12. Pietra del Bagno  3 0.0021 28   370 0.45 780                         500   100   1.389   0.378   –0.413
                                                                         500   100   1.047   0.432   1.141
13. Ustica            26 8.6   53   255 100 1500

14. Levanzo           18 5.61 12    152 4                            42

15. Favignana         28 19.7  8    140 4                            42

16. Marettimo         16 12.06 35   130 15 145

17. Pantelleria       23 86    95   67 115 500

18. Linosa            19 5.34 162   163 40 500

19. Lampione          4 0.025 220   130 18 500

20. Lampedusa         28 20.2 195   120 18 500

21. Malta             43 246   93   292 6 200

22. Gozo              26 67    82   285 6 200

23. Comino            10 3.5   85   295 2 200

24. Cominotto         2 0.25 85     295 0.1 200

25. Filfla            3 0.06 100    292 5 200

urban fabric, Discontinuous urban fabric, Industrial or commer-        There are indications that various insular populations of tene-
cial units, Port areas, Airports, Mineral extraction sites, Dump     brionid beetles described as subspecies differ profoundly geneti-
Sites, Green urban areas, and Sports and Leisure facilities), Cul-   cally (cf. Chatzimanolis et al., 2003; Ferrer, 2008). The current
tivation (including Vineyards, Non-irrigated arable land, Natural    taxonomic dividing line between species and subspecies, as
grassland, Annual crops associated with permanent crops, Com-        applied to the tenebrionids of the circum-Sicilian islands, is
plex cultivation patterns, and Land principally occupied by agri-    arguably arbitrary and the exclusion of subspecies could result
culture, with significant areas of natural vegetation), Coniferous   in a significant underestimate of endemic island tenebrionid
forest, Broad-leaved and Mixed forests, Sclerophyllous vegeta-       diversity. Thus, I considered both species and subspecies as
tion, Bare rock and Sparsely vegetated areas, and Wet areas          “evolutionarily significant units” (Ryder, 1986) and included
(including Salt marshes, Salines, and Water bodies). Even if         both categories in all analyses. The validity of the endemic sub-
these broad landscape units are coarse in comparison to the          species Heliopathes avarus donatellae (Canzoneri, 1970) is dis-
scale at which insects perceive small-scale environmental het-       puted and the identity of the endemic species Opatrum
erogeneity, they correspond well to distinct keystone structures     melitense Küster, 1849 is uncertain. Both taxa have been provi-
(Tews et al., 2004) for tenebrionid species. In particular, each of  sionally considered as valid. On the basis of the original
the seven classes used here corresponded to different microcli-      description, the differentiation of Heliopathes avarus dwejrensis
mate conditions, food resources, and soil characteristics, which     Scupola & Mifsud, 2001 from Gozo seems very subtle. The
are among the most important factors shaping tenebrionid com-        same is the case for Probaticus cossyrensis Sparacio, 2007,
munities in the Mediterranean (Fattorini, 2008b).                    which was recently separated from Probaticus anthrax Seidlitz,
                                                                     1896. All analyses were performed including and excluding
  Geographical distances and land cover categories were              these two taxa. Only results with all taxa included are presented,
obtained from the GIS of the Italian Ministry of the Environ-        because exclusion of these two taxa produced virtually identical
ment and Protection of the Territory and Sea (2009) and from         results.
Malta Environment and Planning Authority (2009). Sea depths
were obtained from bathymetric maps mainly from the Istituto           Finally, cosmopolitan species strictly associated with human
Idrografico della Marina (1997).                                     food, such as Palorus subdepressus (Wollaston, 1864), Gnato-
                                                                     cerus cornutus (Fabricius, 1798), Alphitobius diaperinus (Pan-
Biological data                                                      zer, 1796), Tenebrio spp. and Tribolium spp. were excluded
                                                                     from all analyses to reduce the risk of confounding natural pat-
  Presence/absence data of individual species on each island         terns with effects of man. A list of 45 references used to com-
(Appendix 1) are based on extensive field surveys made by spe-       pile presence/absence data is given in Appendix 1.
cialists with similar sampling efforts among islands; the faunal
inventories are fairly complete. A total of 107 native taxa are
known from these islands.

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