THE CENTIPEDE FAUNA (CHILOPODA) OF CRETE AND ITS SATELLITE ISLANDS (GREECE, EASTERN MEDITERRANEAN)

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1 ISRAEL JOURNAL OF ZOOLOGY, Vol. 50, 2004, pp THE CENTIPEDE FAUNA (CHILOPODA) OF CRETE AND ITS SATELLITE ISLANDS (GREECE, EASTERN MEDITERRANEAN) STYLIANOS SIMAIAKIS, a,b, * ALESSANDRO MINELLI, c AND MOYSIS MYLONAS a,b a Natural History Museum of Crete, University of Crete, P.O. Box 2208, Heraklion (Irakleio), Crete, Greece b Department of Biology, University of Crete, P.O. Box 2208, Heraklion (Irakleio), Crete, Greece c Department of Biology, University of Padova, I Padova, Italy ABSTRACT An overview of the centipede fauna of Crete is presented, with distribution maps, based on the rich centipede collection of the Natural History Museum of Crete and existing data. Centipede records from the satellite islets of Crete are presented for the first time. Distribution patterns according to latitude/ longitude and altitude data of Crete are identified. The analysis of the faunistic elements showed that 25% of the Cretan centipedes have European and South European affinities, 36% have Mediterranean, Northeast, and East Mediterranean affinities, 12% are Balkanic, 10% have Palaearctic affinities, 5% have West Asiatic affinities, and 12% are Cretan endemics. Forty-two species and subspecies are recorded from Crete and the satellite islets. Strigamia acuminata is recorded for the first time from Greece, and 6 additional species (Lithobius lucifugus, L. pamukkalensis, Haploschendyla europea, Nannophilus eximius, Tuoba poseidonis, and Insigniporus sturanyi) are recorded for the first time from Crete. Data on habitat and phenology of each species are also given. INTRODUCTION Crete is one of the biggest islands of the Mediterranean, with a surface area of 8,261 km 2. It is 300 km from the north coast of Africa, 100 km from mainland Greece, and 200 km from Asia Minor, and forms the border between the Aegean and Libyan seas. It is surrounded by more than 50 islets, the biggest of which, Gavdos (32 km 2 ), is the only one inhabited. The majority of the islets are less than 5 km 2, and are very close to the coastline of Crete. Three mountain massifs have peaks that exceed 2000 m asl: Lefka Ori in the west, Psiloreitis in the center, and Dikti in the east. The Cretan archipelago was formed at the end of the Miocene, when it was isolated from Asia Minor in the east and the area of Kyklades in the north. At the beginning of the Pliocene it was isolated from the Greek mainland. Since the beginning of the Pleis- *Author to whom correspondence should be addressed. bio475@edu.biology.uoc.gr Accepted December 2004.

2 368 S. SIMAIAKIS ET AL. Isr. J. Zool. tocene, extensive uplift of the area has joined most of the archipelago into one island: the island of Crete. Hence, a composite fauna with three different elements inhabits Crete: Mediterranean, European, and Asiatic. These three elements have different intensities, depending on the exact locality. In Crete, an isolated island since the Pliocene (4.5 mya) (Dermitzakis, 1990), the first two elements have a greater presence. The centipede (Chilopoda) fauna of Crete is relatively well known, in contrast to other animal groups. What we know comes mainly from the studies of Lucas (1853), Karsch (1888), Cecconi (1895), Attems (1902), Verhoeff (1925), Chamberlin (1956), Kanellis (1959), Dobroruka (1977), Matic (1980), Matic and Stavropoulos (1988, 1993), and Zapparoli (1993, 1994, 2002). The first record of the centipede fauna of Greece was by Gaspard Auguste Brullé (1832), who studied material collected during the historical Expedition scientifique de Morée, organized by the French government in Peloponnisos, between 1829 and 1831, covering Athens, Aigina island, Olymbia, Mykines, Kyklades, and some other islands. During the next 30 years a few reports were published by C. L. Koch ( , 1863), G. Newport (1844, 1845, 1856), P. Gervais (1847), and L. Koch (1862, 1867). Until then, general lists and taxonomic revisions constituted the history of centipedes. Among these papers, Hippolyte Pierre Lucas (1853) wrote the first report on Chilopoda from Crete, collected in 1845 during the expedition of Raulin (1856). Only two species were included in this work: Scolopendra cretica Lucas, 1853 and Scutigera coleoptrata (Linné, 1758). Several decades later, Karsch (1888) presented the results of the collecting trip of the zoologist Eberhard von Oertzen from continental Greece (Attica, Peloponnisos), the Ionian Islands (Kephalonia), Kyklades (Naxos), and Crete between 1884 and 1885 (Zapparoli, 2002). Ten years later, Cecconi (1895) published a study of the centipedes of Crete collected by him in June At the beginning of the 20th century the Austrian myriapodologist Carl Graaf Attems published a critical work concerning the centipede fauna of Greece. He presented material collected by Max Beier, Gustav Paganetti-Hummler, and a few others to the Museum of Natural History of Vienna (Zapparoli, 2002). These articles were related to the fauna of Crete, the Ionian Islands, Thessalia, and Peloponnisos (Attems, 1902, 1903a,b, 1929b, 1935). In the second half of the nineteen-fifties, the American myriapodologist Ralph V. Chamberlin published data on the cave centipede fauna of Greece (Chamberlin, 1956). This material had been collected in Crete by the Swedish zoologist K. Lindberg, between April and May Three years later, the Greek zoologist, A. Kanellis, completed the first modern list of Chilopoda from Greece (Kanellis, 1959). Apart from the presentation of the known species and the new geographical references, Kanellis presented the first fairly complete key to Chilopoda of Greece. Since the end of the nineteen-sixties, many other faunistic studies have been published that have significantly increased our knowledge on the centipede fauna of continental Greece (Matic et al., 1968; Matic, 1970; Matic and Stavropoulos, 1990a; Zapparoli, 1984, 1994) and insular Greece (Dobroruka, 1965, 1977; Matic, 1976, 1980;

3 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 369 Matic and Stavropoulos, 1988, 1990a,b, 1993; Stavropoulos and Matic, 1990; Zapparoli, 1994; see also Zapparoli, 2002). Recently, Zapparoli (2002) presented a modern catalogue of the centipede fauna of Greece, in which 29 species were recorded from Crete. This work was based on material collected from the zoological trips of several research institutes and many naturalists in Greece (Museum of Natural History of Bergamo, University of Padova, Museum of Natural History of Carmagnola, Museum of Natural History of Torino, Museum of Natural History of Brescia, and Zoological Institute of the University of Innsbruck). Zapparoli, apart from his taxonomic work, provided distribution maps and ecology notes for many species. However, knowledge concerning the distribution of centipedes on Crete is still limited. Scantier still are data concerning the centipede fauna of the small adjacent islands. Furthermore, the taxonomic status of several species groups in Bothriogaster, Pachymerium, and Henia and around Lithobius piceus needs to be revised, while information related to habitat preferences is still incomplete. In this paper we present a more detailed catalogue of the centipede species of Crete and its satellite islets based on hitherto unpublished records of the collection of the Natural History Museum of Crete (NHMC), as well as on literature data. We also provide evidence on altitudinal distribution patterns, habitat notes, and phenological information for many species. Finally, we present geographical maps for each species based on literature and unpublished records. MATERIALS AND METHODS More than 3000 centipedes were collected from 89 sites on Crete and the surrounding islands of Agria Gramvousa, Chrysi, Dia, Elasa, Gavdos, Gavdopoula, Grantes, Koufonisi, Makroulo, Marmara, Strongylo, and Trachilos. Crete is covered through its main east west axis, as well as north to south, and along altitudinal gradients up to 2420 m. The three mountain massifs of the island, Lefka Ori Mts. (White Mountains) to the west, Oros Idi (Psiloreitis Mts.) to the center, and Dikti Mts. to the east, are also covered. Centipedes were collected by means of pitfall traps (PT) or by hand (H). In the pitfall traps (12 cm height, 9.5 cm diameter) the fixing medium used was ethylene glycol, which is colorless, odorless, and not volatile. Pitfall traps were set in operation for one year, and the content of the traps was collected seasonally. Centipedes were preserved in 75% alcohol. Identification took place at the Natural History Museum of Crete (NHMC). Most of the material presented here is part of the M.Sc. thesis of the first author and is deposited at the NHMC. For each species we give the complete name, the list of literature and NHMC records, distribution on Crete, altitudinal range and highest locality on the island, overall geographic distribution, chorotype, and ecology notes. The phenological notes within each ecological paragraph were based only on the data collected by pitfall traps. The collection sites are listed in numerical order together with the number and sex

4 370 S. SIMAIAKIS ET AL. Isr. J. Zool. of each specimen. Some of the samples are unsexed and a few lack a specimen number due to incomplete identification. Maps and collecting sites were drawn with Arc View GIS version 3.1. and Corel Draw 10. A description of many collection sites was provided for the first time by Chatzaki et al. (2002). The full description of the collecting sites and intervals is given in the Appendix. RESULTS SCUTIGEROMORPHA Gervais, 1837 SCUTIGERIDAE Gervais, 1837 Scutigera Lamarck, Scutigera coleoptrata (Linné, 1758) CRETE (Karsch, 1888: 220; Verhoeff, 1925: 658; Kanellis, 1959: 32; Zapparoli, 1993: 96; Matic and Stavropoulos, 1993: 106; Zapparoli, 1994: 61; Stoev, 1997: 102; Zapparoli, 2002: 9): Chania: Askifou (Dobroruka, 1977: 163; Zapparoli, 2002: 9); Chania (Lucas, 1853: 531; Raulin, 1856; Kanellis, 1959: 32; Zapparoli, 2002: 9); Gallos (type locality of Scutigera coleoptrata var. crinita Attems, 1902) (Attems, 1902: 541; Kanellis, 1959: 32; Zapparoli, 2002: 9); Kolimbari (Chamberlin, 1956: 53; Zapparoli, 2002: 9); Vizari (type locality of Scutigera coleoptrata var. crinita Attems, 1902) (Attems, 1902: 541; Kanellis, 1959: 32; Zapparoli, 2002: 9). Rethymno: Gonies to Anogeia (Zapparoli, 1994: 61; Zapparoli, 2002: 9); Rethymno (Lucas, 1853: 531; Raulin, 1856; Kanellis, 1959: 32; Zapparoli, 2002: 9). Irakleio: Dafnes (type locality of Scutigera coleoptrata var. crinita Attems, 1902) (Attems, 1902: 541; Kanellis, 1959: 32; Zapparoli, 2002: 9); Irakleio (Lucas, 1853: 531; Raulin, 1856; Kanellis, 1959: 32; Zapparoli, 2002: 9). Lasithi: Agios Nikolaos (Zapparoli, 1994: 61; 2002: 9); Gournia; Gournia, between Agios Nikolaos and Ierapetra (Zapparoli, 2002: 9). CRETE: Chania: 3 (a 15; b 9; c 4; e 1; f 4); 6 (a 2 ; e 1 1 ; f 1 1 ; g 3 ; h 1 ; j 1 ); 16 (a 1 ; b 3 ); 17 (a 3 5 ; b 4 ; c 3 ; d 4 5 ); 18 (a 7 9 ; b 1 1 ; c 10 6 ; d 3 ); 19 (a 1 2 ; c 1 ; d 1 1 ); 20 (a 3 ; b 1 ; c 5 2 ; d 1 ); 21 (b 1 ; d 1 ); 22 (a 4 8 ; b 1 ; c 14 5 ; d 11 1 ); 23 (a 1 ; c 8 1 ; d 2 1 ); 24 (c 2 1 ; d 9 ); 25 (a 3 1 ; c 1 ); 26 (c 5 1 ; d 11 2 ); 27 (a ; b 6 9 ; c ). Rethymno: 29 (a 39 5 ; b 22 7 ; c 6 ; d 2 ; f 1 1 ); 30 (a 2 ; c 1 1 ); 32 (a 1 ; b 6 ; c 2 ); 34 (a 1 ); 39 (b 1 ; d 1 1 ); 40 (c 1 1 ); 43 (b 1 ). Irakleio: 46 (a 1 ); 48 (a 3 2 ); 49 (a 1 3 ); 51 (a 4 ); 56 (b 2 ); 59 (a 12 ; b 25 6 ; c 3 5 ); 60 (b 12 3 ; c 3 2 ). Lasithi:

5 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS (a 1 ); 63 (a 3 ; b 15 4 ; c 1 1 ); 66 (a 1 ); 70 (a 1 ); 72 (b 1 ); 73 (a 1 ); 75 (a 6 2 ; b 7 14 ; c 6 9 ; e 1 ); 78 (b 7 2 ; c 2 ); 79 (a 1 3 ); 82 (a 1 ). Cretan (Fig. 1). Altitudinal range: m. Highest site: 34. Geographical distribution: Mainland and insular Greece, Albania, Algeria, Austria, Balearic Is., Bulgaria, Caucasus, Corsica, Crimea, Croatia, Egypt, France, FYR Macedonia, Italy, Libya, Montenegro, Morocco, Near and Middle East, Portugal, Romania, Sardinia, Sicily, Slovenia, South Germany, South Hungary, southern part of former Czechoslovakia, Spain, Switzerland, Tunisia, Ukraine (Zapparoli, 2002). South European Mediterranean. Scutigera coleoptrata is very common on Crete. It has been found at many sites from western to eastern Crete, mainly in the lowlands up to 800 m. The peak of activity of females is from spring to autumn while males are found in late summer and in early autumn. Quite euryoecious in habitats modified by man with Coridothymus capitatus, Sarcopoterium spinosum, Pistacia lentiscus, Juniperus macrocarpa, J. phoenicea, Pinus brutia, and Nerium oleander. Fig. 1. Scutigera coleoptrata and Eupolybothrus litoralis distribution on Crete.

6 372 S. SIMAIAKIS ET AL. Isr. J. Zool. LITHOBIOMORPHA POCOCK, 1895 LITHOBIIDAE Newport, 1844 Eupolybothrus Verhoeff, 1907 Subgenus Eupolybothrus Verhoeff, Eupolybothrus (Eupolybothrus) litoralis (L. Koch, 1867) CRETE (Verhoeff, 1925: 658; Attems, 1929b: 469; Zapparoli, 1990: 54; 1993: 96; 2002: 14; Matic and Stavropoulos, 1993: 106; Stoev, 1997: 99), Chania: Agia Roumeli (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Vizari (Attems, 1902: 548; Kanellis, 1959: 29; Zapparoli, 2002: 14); Akrotiri, Moni Gouvernetou, Koutrouli Cave (Chamberlin, 1956: 53; Zapparoli, 2002; 14); Askifou (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Georgioupoli (Zapparoli, 2002: 14); Gallos (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Chania (Koch, 1867: 899; Kanellis, 1959: 32; Zapparoli, 2002; 14); Imbros Gorge (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); between Kandanos and Floria (Zapparoli, 2002: 14); Lakki (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Mournies (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Nerokouros (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14). Rethymno: Assomatos (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Oros Idi (Dobroruka, 1977: 164; Zapparoli, 2002: 14); Rethymno (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14). Irakleio: Dafnes (Attems, 1902: 547; Kanellis, 1959: 29; Zapparoli, 2002: 14); Kamilari Cave (Chamberlin, 1956: 53; Zapparoli, 1994: 27; 2002; 14); Irakleio (Zapparoli, 2002: 14). Lasithi: Gournia (Zapparoli, 2002: 14); Lato (Zapparoli, 2002: 14); Neapoli, Moni Kremaston (Zapparoli, 2002: 14); Vrouchas (Zapparoli, 2002: 14). CRETE: Chania: 6 (a 2 ; b 2 ; c 1 4 ; d 8 ; e 5 ; f 2 ; g 1 ; i 2 1 ; j 2 ; l 4 1 ; m 1 ); 10 (c 1 ); 11 (b 1 ); 15 (a 2 2 ; b 2 2 ); 16 (a 1 7 ; b 3 3 ; c 6 1 ); 21 (d 1 ). Rethymno: 28 (a 2 7 ); 29 (a 2 2 ; f 1 1 ); 30 (a 2 ; b 1 2 ; c 2 ); 31 (a 4 1 ); 32 (a 1 ; b 2 5 ; c 2 ; d 4 2 ); 36 (c 1 1 ; d 3 ; e 1 ); 39 (b 1 6 ; d 2 1 ); 41 (a 6 ); 43 (a 4; b 6); 44 (a 1 ). Irakleio: 46 (a 2 ); 49 (a 1 ); 50 (a 1 ); 51 (a 1 ); 52 (a 3 3 ); 55 (a 4 1 ); 56 (b 4 ). Lasithi: 63 (a 2 2 ; b 1 ); 64 (a 1); 67 (a 2 1 ); 69 (a 1 ); 72 (b 3 1 ); 74 (b 1 ); 75 (b 2 ); 78 (e 1 ); 82 (a 1). Cretan (Fig. 1). Altitudinal range: m. Highest site: 11. Mainland and insular Greece, Albania, Bosnia Herzegovina, Bulgaria, Croatia, Cyprus, Montenegro, Near and Middle East, Romania, Slovenia, Turkey (Zapparoli, 2002).

7 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 373 Balkan East Mediterranean. Eupolybothrus litoralis is widespread on the island and on the surrounding islands examined. It has been found at many sites from western to eastern Crete, at altitudes up to 2000 m. Adults occur during the whole year, with a peak of activity of females in autumn and of males in late winter and in spring. Quite euryoecious, present in phrygana with Coridothymus capitatus, Sarcopoterium spinosum, in more or less arid biotopes, in riparian sites with Nerium oleander, in human-affected ecosystems, in open mountainous areas with Genista acanthoclada as well as in Pinus brutia, Quercus coccifera, Pistacia lentiscus, Juniperus macrocarpa, or J. phoenicea formations. Lithobius Leach, 1814 Subgenus Lithobius Leach, Lithobius (Lithobius) agilis C. L. Koch, 1847 CRETE (Zapparoli, 1993: 96; 2002: 20; Stoev, 1997: 99): Rethymno: Oros Idi, N slope, m (Zapparoli, 1994: 35; 2002: 14). CRETE: Chania: 1 (a 1 ); 6 (c 1 ; d 2 ; j 1 ); 11 (c 1 1 ). Rethymno: 34 (b 2 ); 36 (c 1 ). West-central Cretan (Fig. 2). Altitudinal range: m. Highest site: 36. Mainland and insular Greece, Austria, Belgium, Bosnia Herzegovina, Croatia, Denmark, former Czechoslovakia, France, Germany, Hungary, Netherlands, North Italy, Poland, Romania, Sardinia, Slovenia, South Scandinavia, Switzerland, Ukraine (Zapparoli, 2002). European. It is common in Pinus brutia forests, in plateau and open alpine formations with Satureja spinosa, Berberis cretica, and Genista acanthoclada, as well as in phryganic ecosystems with Sarcopoterium spinosum, Verbascum spinosum, Euphorbia acanthothamnos, and Drimia maritima. 4. Lithobius (Lithobius) cretaicus Matic, 1980 CRETE (Zapparoli, 1993: 96; 2002: 23; Matic and Stavropoulos, 1993: 106; Stoev, 1997: 99): Irakleio: Kamilari Cave, 200 m (type locality) (Matic, 1980: 100; Zapparoli, 1994: 36; 2002: 22; Stoev and Beron, 2000: 93).

8 374 S. SIMAIAKIS ET AL. Isr. J. Zool. Fig. 2. Lithobius agilis, L. cretaicus, and L. creticus distribution on Crete. Central Cretan (Fig. 2). Altitudinal range: 200 m. Highest site: Kamilari Cave (type locality). Crete (Matic and Stavropoulos, 1993). Cretan endemic South European affinities. Only known from the type locality (Zapparoli, 2002). 5. Lithobius (Lithobius) creticus Dobroruka, 1977 CRETE (Stoev, 1997: 99; Zapparoli, 2002: 23): Rethymno: Oros Idi (type locality) (Dobroruka, 1977: 161; Zapparoli, 2002: 23); Oros Idi, N slope, m (Zapparoli, 1994: 37; 2002: 23). CRETE: Chania: 6 (l 1 ); 8 (d 1 ); 9 (a 1 ); 11 (a 1 ; c 1 1 ); 15 (a 2 ); 17 (c 8 5 ); 18 (c 1 ); 19 (c 2 1 ; d 2 ); 20 (c 1 2 ); 22 (c 8 ); 23 (c 1 ); 24 (a 6 6 ); 25 (a 1 ; c 1 ); 27 (a 1 1 ; c 2 1 ). Rethymno: 30 (a 2 2 ); 32 (a 1 ; b 2 1 ); 35 (a 1 ; c 1 1 ; d 1

9 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS ; e 1 ); 36 (a 1 ; c 1 ; d 1 ); 37 (a 1 ; b 1 1 ); 38 (a 1 1 ); 39 (b 3 ); 41 (a 2 ); 43 (a 1 ). Irakleio: 48 (a 2 ); 50 (a 5 4 ); 52 (a 3 2 ); 55 (a 2 2 ); 59 (b 3 6 ; c 1 ); 60 (c 1 ). Lasithi: 61 (a 1 ); 63 (a 3 5 ); 65 (c 1 ); 67 (b 2 ); 78 (e 2 ); 79 (a 1); 82 (a 1 ); 84 (a 1 ). Cretan (Fig. 2). Altitudinal range: m. Highest site: 38. Crete (Zapparoli, 2002). Cretan endemic South European affinities. Lithobius creticus is widespread on the island and on the surrounding islands examined. It is distributed from western to eastern Crete, at altitudes up to 2400 m. Females are found in spring while males occur in late spring and in early summer. Quite euryoecious, present in forests with Pinus brutia and Cupressus sempervirens, in open mountainous shrubs with Berberis cretica, Genista acanthoclada, and Phlomis sp., in ecosystems dominated by ferns and Platanus orientalis, in common mixed phryganic and maquis formations with Sarcopoterium spinosum, Phlomis sp., Pistacia lentiscus, Juniperus phoenicea, or J. macrocarpa, and in sand dunes dominated by Juniperus macrocarpa along with sparse Pistacia lentiscus and Coridothymus capitatus as well as in coastal phryganic areas dominated by Calicotome villosa, Sarcopoterium spinosum, and Urginea maritima. 6. Lithobius (Lithobius) erythrocephalus C. L. Koch, 1847 CRETE (Zapparoli, 1993: 96; 2002: 25; Matic and Stavropoulos, 1993: 106); Chania: Nykteridospilio (Afrata), 2 m (Zapparoli, 1994: 39; 2002: 25); Hellenospilio (Afrata), 15 m (type locality of Sigibius creticola Chamberlin, 1956) (Chamberlin, 1956: 52; Zapparoli, 1994: 38; 2002: 25). CRETE: Chania: 6 (b 1 ); 8 (a 1 2 ; c 3 ; i 1 ; j 3 ); 10 (a 1 ; d 1 ); 15 (a 1 ). Rethymno: 28 (a 1 ); 32 (c 1 ; e 1 ). Irakleio: 55 (a 1 ). Lasithi: 67 (a 2 1 ). Cretan (Fig. 3). Altitudinal range: m. Highest site: 10. Mainland and insular Greece, Albania, Armenia, Austria, Azerbaijan, Bosnia Herzegovina, Bulgaria, Croatia, Denmark, Finland, former Czechoslovakia, France (Alps), FYR Macedonia, Georgia, Germany, Hungary, Iran, Italy, Jordan, Latvia, Lebanon, Lithuania, Montenegro, Netherlands, Norway, Poland, Romania, Serbia, Sicily,

10 376 S. SIMAIAKIS ET AL. Isr. J. Zool. Slovenia, Sweden, Switzerland, Turkey, Ukraine, West Russian Federation (Zapparoli, 2002). European West Asiatic. Lithobius erythrocephalus is distributed from western to eastern Crete, at altitudes up to 1650 m. The peak of activity of females occurs in spring and in autumn. It has been mainly collected in forests with Pinus brutia, Cupressus sempervirens, or Quercus coccifera, in open mountainous phryganic ecosystems, and from plateaus with Satureja spinosa, Berberis cretica, Genista acanthoclada, and Euphorbia acanthothamnos as well as from old terraces with sparse phrygana surrounded by P. brutia. 7. Lithobius (Lithobius) lucifugus L. Koch, 1862 CRETE (Stoev, 1997: 100; Zapparoli, 2002: 31). According to Stoev (2003: personal communication) this citation is dubious. CRETE: Chania: 6 (1 ); 10 (2 ); 11 (8 6 ); 15 (2 ); 16 (4 5 ). Rethymno: 34 (1 1 ); 36 (1 ); 37 (1 ). Irakleio: 55 (2 ). Lasithi: 65 (2 ). Fig. 3. Lithobius erythrocephalus, L. lucifugus, L. piceus, and L. nigripalpis distribution on Crete.

11 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 377 Cretan (Fig. 3). Altitudinal range: m. Highest site: 37. Mainland Greece, Austria, Bosnia Herzegovina, Caucasus, Crimea, Croatia, former Czechoslovakia, Germany, Hungary, Montenegro, North Italy, Poland, Poland, Romania, Russia, Scandinavia, Slovenia, Southeast France, Switzerland, Turkey, Ukraine (Zapparoli, 2002). First record for Crete. European West Asiatic. Lithobius lucifugus is common in the middle to high altitudes of the mountains from western to eastern Crete. Present peak of adult activity is the dry period of summer. It is present in forests with Pinus brutia and Cupressus sempervirens, in valleys, subalpine and alpine shrubs with very sparse vegetation composed of Berberis cretica, Prunus prostrata, Astragalus angustifolius, and Satureja spinosa, and in places dominated by ferns and Platanus orientalis. 8. Lithobius (Lithobius) nigripalpis L. Koch, 1867 CRETE (Verhoeff, 1925: 658; Attems, 1929b: 300; Matic and Stavropoulos, 1988: 42; Zapparoli, 1993: 96; 2002: 34; Matic and Stavropoulos, 1993: ; Stoev, 1997: 100): Chania: Agia Roumeli (Attems, 1902: 551; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Akrotiri, Moni Gouvernetou, Koutrouli Cave (type locality of Lithobius sitianus Chamberlin, 1956, cf. Zapparoli, 1994: 45) (Chamberlin, 1956: 53; Zapparoli, 1994: 45; 2002: 34); Askifou (type locality of Lithobius peregrinus circula Attems, 1902, cf. Zapparoli, 1994: 45) (Attems, 1902: 551, 552; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Georgioupoli, hillside, phrygana, pasture, m (Zapparoli, 2002: 34); Georgioupoli, at hill, phrygana, pasture, m (Zapparoli, 2002: 34); Georgioupoli, seashore, cultivated land, olive plantation (Zapparoli, 2002: 34); near Georgioupoli, spring and muddy area at hill base (Zapparoli, 2002: 34); Gallos (Attems, 1902: 551; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Imbros Gorge (Attems, 1902: 551; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Imbros (Zapparoli, 2002: 34); between Kandanos and Floria (Zapparoli, 2002: 34); Niatos Askifou (Zapparoli, 2002: 34); Paleochora, Maniationa (Zapparoli, 2002: 34); Samaria Gorge (Attems, 1902: 552; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Kournas Lake (type locality of Lithobius lindbergi Chamberlin, 1956) (Chamberlin, 1956: 53); Mournies (Attems, 1902: 552; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Nerokouros (Attems, 1902: 552; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); Omalos (Cecconi, 1895: 193; Kanellis, 1959: 24; Zapparoli, 1994: 45; 2002: 34); Omalos, 1000 m (Zapparoli, 2002: 34). Gavdos Island (Dobroruka, 1977: 164). Rethymno: Anogia (Matic, 1980: 100; Zapparoli, 2002: 34); Assomatos (Attems, 1902: 551; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34); near Gonies (Zapparoli, 1994: 45; 2002: 34); Oros

12 378 S. SIMAIAKIS ET AL. Isr. J. Zool. Idi (Dobroruka, 1977: 164; Zapparoli, 2002: 34); Oros Idi, m (Zapparoli, 1994: 45; 2002: 34); Oros Idi, slope of Anogia, 800 m (Zapparoli, 2002: 34); Oros Idi, slope of Anogia, 900 m (Zapparoli, 2002: 34); Oros Idi, slope of Anogia, 1500 m (Zapparoli, 2002: 34); environs of Oros Idi (Zapparoli, 2002: 34); Perama (Matic, 1980: 100; Zapparoli, 2002: 34); Bali (Zapparoli, 2002: 34); Rethymno (Cecconi, 1895: 193; Attems, 1902: 552; Kanellis, 1959: 24, 25; Zapparoli, 1994: 45; 2002: 34). Irakleio: Irakleio (Zapparoli, 2002: 34); karst area near Irakleio (Zapparoli, 2002: 34); Karteros (Chamberlin, 1956: 53; Zapparoli, 2002: 34); Kasteli-Viannos, Afrati, in olive plantation, 500 m (Zapparoli, 2002: 34); Knossos (Matic, 1980: 102; Zapparoli, 2002: 34); Dafnes (Attems, 1902: 551, 552; Kanellis, 1959: 25; Zapparoli, 1994: 45; 2002: 34). Lasithi: Agios Nikolaos (Eason, 1990: 261; Zapparoli, 2002: 34); Agios Nikolaos, Ammoudara, at coast, 50 m near sea shore, phrygana (Zapparoli, 2002: 34); Agios Nikolaos, Dikteon Antron, 960 m (Eason, 1990: 257; Zapparoli, 1994: 45; 2002: 34); between Agios Nikolaos and Kritsa, Katharo Plateau, oaks, karst, m (Zapparoli, 2002: 34); Exo Potami (Zapparoli, 2002: 34); Gournia (Zapparoli, 2002: 34); environs of Kritsa (Zapparoli, 2002: 34); between Kroustas and Prina (Zapparoli, 2002: 34); between Lasithi and Tzermiado, Timios Stavros, m (Zapparoli, 2002: 34); Lasithi, W, path Kato Metochi Kastamonitsa (Zapparoli, 2002: 34); Lato (Zapparoli, 2002: 34); Megalo Katofigui Cave (Chamberlin, 1956: 53; Zapparoli, 1994: 45; 2002: 35); Milatos Cave, 140 m (Chamberlin, 1956: 53; Eason, 1990: 257; Zapparoli, 1994: 45; 2002: 35); Orno Oros, Mesa Mouliana (Zapparoli, 2002: 35); Neapoli, Moni Kremaston (Zapparoli, 2002: 35); Neapoli (Zapparoli, 1994: 45; 2002: 35); Sitia (Zapparoli, 2002: 35); Tzermiado, Trapezas Cave (Eason, 1990: 257; Zapparoli, 2002: 35); Vrouchas (Zapparoli, 2002: 35). UNCERTAIN DATA Iva Gipfel Sock ; N. Alavia, Age Thopsi ; Chaui [or Chani?] Koku (Zapparoli, 1994: 45; 2002: 34). Referring to Wettstein (1942, 1952) Iva Gipfel Sock could be the highest peak of Oros Idi, Psiloreitis Mt., 2456 m; Chaui Koku is probably in the locality of Kokkini Chani near Irakleio, whilst Age Thopsi could be in the locality of Aga Topi, Omalos, 1300 m, near Chania. CRETE: Chania: 1 (a 1 ); 6 (a 5 ; c 1 2 ; d 1 1 ; e 2 ; g 2 1 ; i 1 2 ; j 1 ; l 1 ; m 1 ); 8 (a 2 1 ; c 1 3 ; d 4 1 ; j 1 2 ); 10 (i 1 ); 11 (a 1 ; b 2 1 ; c 3 6 ; d 1 ; e 1 ; g 1 ); 14 (a 1 ); 15 (a 4 7 ; b 3 2 ; c 4 ); 16 (a 1 ; b 2 ; c 1 ); 17 (b1 ; c 5 11 ; d 1 ); 18 (a 1 2 ; b 4 2 ; c ; d 1 1 ); 19 (a 1 ; b 1 ; c 2 ; d 1 2 ); 20 (a 2 1 ; b 1 ; c 5 3 ; d 2 1 ); 21 (a 2 1 ; c 2 1 ; d 1 ); 22 (a 1 1 ; b 1 ; c ; d ); 23 (a 2 ; b 1 3 ; d 1 ); 24 (a ; b 1 ); 25 (a 3 ); 26 (a 8 4 ; b 1 1 ); 27 (a 2 ; c 2 ). Rethymno: 28 (a 1 1 ); 29 (a 5 2 ; b 1 ; d 1 ; f 1 1 ); 30 (a 2 ; b 1 1 ); 31 (a 2 1 ); 32 (a 3 3 ; b 4 3 ; c 4 3 ; d 5 ; e 4 8 );

13 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS (a 1 2 ; b 4 3 ); 35 (e 1 ); 36 (a 3 1 ; c 1 1 ; d 1 5 ; e 1 1 ); 37 (a 1 ; b 1 ); 39 (b 1 6 ); 43 (a 2 ; b 1 ); 44 (b 1 ). Irakleio: 45 (a 2 ); 46 (a 1 ); 48 (a 1 ); 49 (a 1 ); 54 (a 2 1 ); 55 (a 10 5 ); 57 (a 6 1 ); 58 (a 5 3 ); 59 (a 1 5 ; b 5 11 ; c 1 2 ); 60 (b 2 ). Lasithi: 62 (a 6 4 ); 63 (a 2 ; e 2 ); 64 (b 1 1 ; d 2 2 ); 65 (a 4 2 ); 67 (a 4 6 ; b 1 ); 68 (a 1 ); 69 (a 3 ); 70 (a 1 ) 72 (a 1 1 ); 74 (b 3 ); 77 (a 3 2 ). Cretan (Fig. 3). Altitudinal range: m. Highest site: 37. Mainland and insular Greece, Bulgaria, FYR Macedonia Romania, Turkey (Zapparoli, 2002). Northeast Mediterranean. Lithobius nigripalpis is the most common species of Lithobius on Crete. It has been found at many sites from western to eastern Crete, up to 2400 m. Both females and males are found in spring, but males are also found during summer and autumn. Quite euryoecious in habitats modified by man, in agroecosystems, in typical phrygana with Coridothymus capitatus, Phlomis lanata, Satureja thymbra, Salvia fruticosa, Genista acanthoclada, Erica sp., and Cistus sp. in maquis dominated by Pistacia lentiscus, Quercus ilex, Q. coccifera, and Olea europea, and ferns, in plateaus with prostrate shrubs, mainly Juniperus oxycedrus, Berberis cretica, Prunus prostrata, and Satureja spinosa, in subalpine and alpine sites with B. cretica, P. prostrata, Astragalus angustifolius, and S. spinosa, and in Pinus brutia, Cupressus sempervirens, and Q. coccifera forests as well as on sand dunes dominated by J. macrocarpa. 9. Lithobius (Lithobius) piceus L. Koch, 1862 CRETE (Kanellis, 1959: 24; Matic and Stavropoulos, 1993: 104; Stoev, 1997: 100): Chania: Omalos (Cecconi, 1895: 193; Kanellis, 1959: 25). Rethymno: Rethymno (Cecconi, 1895: 193; Kanellis, 1959: 24). CRETE: Rethymno: 35 (d 1 1 ); 36 (d 1 ). Central Cretan (Fig. 3). Altitudinal range: m. Highest site: 36. Mainland Greece, Austria, Bosnia Herzegovina, Caucasus, Crimea, Croatia, former Czechoslovakia, France, Germany, Hungary, Italy, Montenegro, Poland, Poland, Romania, Sardinia, Slovenia, Switzerland (Zaleskaja, 1978).

14 380 S. SIMAIAKIS ET AL. Isr. J. Zool. European. Lithobius piceus has been collected at two localities only, in subalpine and alpine phrygana at the central mountain massif of Crete dominated by Berberis cretica, Astragalus angustifolius, and Satureja spinosa. 10. Lithobius (Lithobius) viriatus Sseliwanoff, 1878 CRETE: Lasithi: Oros Selena, karst, limestone rocks, 1559 m (Zapparoli, 2002: 41). A former quotation of Lithobius viriatus from Crete (Zapparoli, 1993) was later found erroneous by its author (Zapparoli, 2002). Chania: 15 (a 2 5 ). Rethymno: 34 (b 1 2 ); 36 (d 1 ). Lasithi: 72 (b 5 1 ). Cretan (Fig. 4). Altitudinal range: m. Highest site: 36. Mainland Greece, Albania, Bulgaria, Caucasus, Crimea, Israel, Jordan, Northwest Iran, Southeast Italy, Turkey (Zapparoli, 2002). Northeast Mediterranean West Asiatic. Lithobius viriatus occurs mainly at the high altitudes of Crete. For instance, it is found in the western mountain massif in open valleys with Berberis cretica, Genista acanthoclada, Sarcopoterium spinosum, and Phlomis sp. together with some Quercus coccifera and Acer sempervirens, as well as in alpine phrygana in the central mountain massif of Crete dominated by Berberis cretica, Astragalus angustifolius, and Satureja spinosa. It was also collected among riparian vegetation with Nerium oleander, Coridothymus capitatus, Calicotome villosa, and sparse Euphorbia sp. Subgenus Sigibius Chamberlin, Lithobius (Sigibius) micropodus (Matic, 1980) CRETE (Zapparoli, 1993: 96; 2002: 45, Stoev, 1997: 101): Irakleio: Knossos (Matic, 1980: 101; Zapparoli, 2002: 45). Central Cretan (Fig. 4). Altitudinal range: Highest site: Knossos.

15 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 381 Mainland and insular Greece, Albania, Bosnia Herzegovina, Croatia, France, Italy, Montenegro, Romania, Russian Plain, Sardinia, Serbia, Sicily, Slovenia, Turkey (Zapparoli, 2002). South European Northeast Mediterranean. The unique data of the species collected at Knossos does not permit accurate conclusions. Subgenus Monotarsobius Verhoeff, Lithobius (Monotarsobius) aeruginosus L. Koch, 1862 CRETE (Verhoeff, 1925: 658; Attems, 1929c: 304; Stoev, 1997: 101; Zapparoli, 2002: 48); Chania: Askifou (Attems, 1902: 553; Kanellis, 1959: 27; Zapparoli, 2002: 45). West Cretan (Fig. 4). Altitudinal range: Highest site: Askifou. Fig. 4. Lithobius aeruginosus, L. crassipes, L. micropodus, L. viriatus, and L. pamukkalensis distribution on Crete.

16 382 S. SIMAIAKIS ET AL. Isr. J. Zool. Mainland and insular Greece, Albania, Austria, Bosnia Herzegovina, Caucasus, Crimea, Croatia, former Czechoslovakia, Germany, Netherlands, North Italy, Romania, Slovenia, Switzerland, Turkey, Ukraine (Zapparoli, 2002). European Northeast Mediterranean. No habitat data available (Zapparoli, 2002). 13. Lithobius (Monotarsobius) crassipes L. Koch, CRETE (Zapparoli, 1993: 96; 2002: 50; Matic and Stavropoulos, 1993: 106; Stoev, 1997: 101); Chania: Agia Sofia Cave, Kissamos (Chamberlin, 1956: 52; Zapparoli, 2002: 50). Rethymno: Perama (Matic, 1980: 101; Zapparoli, 2002: 50). Irakleio: Kamilari Cave, 200 m (Matic, 1980: 101; Zapparoli, 1994: 58; 2002: 50); Agia Triada Cave (Chamberlin, 1956: 52; Zapparoli, 2002: 50). West Central Cretan (Fig. 4). Altitudinal range: m. Highest site: Agia Sofia Cave. Mainland and insular Greece, Algeria, Austria, Bosnia Herzegovina, Bulgaria, Canary Is., Caucasus, Central Asia, Crimea, Croatia, former Czechoslovakia, France, FYR Macedonia, Germany, Iberia, Ireland, Italy, Jordan, Madeira, Montenegro, Netherlands, Poland, Romania, Russia, Sardinia, Scandinavia, Serbia, Sicily, Slovenia, Switzerland, Syria, Tunisia, Turkey, Ukraine, United Kingdom (Zapparoli, 2002). West Palearctic. On Crete Lithobius crassipes has been mainly collected in caves (Chamberlin, 1956; Matic, 1980; Zapparoli, 1994). Subgenus Porobius Porat, Lithobius (Porobius) pamukkalensis Matic, 1980 CRETE: Lasithi: 72 (a 1 12 ); 75 (a 1 ; d 1 ; e 2 ); 78 (c 1 ; d 3 ; e 1 ); 82 (a 2 1 ). East Cretan (Fig. 4). Altitudinal range: m. Highest site: 78.

17 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 383 Insular Greece, South Turkey (Zapparoli, 2002). First record for Crete. Northeast Mediterranean. Lithobius pamukkalensis occurs mainly in the lowlands of eastern Crete. Females are found in spring while males are found in late autumn and in winter. It has been collected among riparian vegetation with Nerium oleander, Coridothymus capitatus, Calicotome villosa, and sparse Euphorbia sp., as well as in sparse phrygana dominated by Phlomis lanata, Satureja thymbra, Salvia fruticosa, Genista acanthoclada, Erica sp., and Cistus sp. SCOLOPENDROMORPHA Pocock, 1895 SCOLOPENDRIDAE Newport, 1894 Scolopendra Linné, Scolopendra cretica Lucas, 1853 CRETE (Kraepelin, 1904: 318; Verhoeff, 1925: 659; Attems, 1930: 37; Würmli, 1980: 350; Lewis, 1985: map 2; Zapparoli, 1993: 95; 2002: 63; Matic and Stavropoulos, 1993: 104; Stoev, 1997: 102; Zapparoli, ): Chania: Akrotiri, Moni Gouvernetou, Katholiko Cave (Chamberlin, 1956: 51; Zapparoli, 2002; 63); Akrotiri, Moni Gouvernetou, Koutrouli Cave (Chamberlin, 1956: 51; Zapparoli, 2002; 63); Askifou (type locality of Scolopendra oraniensis lusitanica var. cretica Attems, 1902) (Attems, 1902: 559; Kanellis, 1959: 19; Dobroruka, 1977: 164; Zapparoli, 2002: 63); Askifou (Zapparoli, 2002: 63); Niatos Askifou (Zapparoli, 2002: 63); Georgioupoli, m (Zapparoli, 2002; 63); near Georgioupoli, m (Zapparoli, 2002; 63); Gallos (type locality of Scolopendra oraniensis lusitanica var. cretica Attems, 1902) (Attems, 1902: 559; Kanellis, 1959: 19; Zapparoli, 2002: 63); Chania (type locality of Scolopendra cretica Lucas, 1853; type locality of Scolopendra oraniensis lusitanica var. cretica Attems, 1902) (Lucas, 1853: 529; Raulin, 1856; Karsch, 1888: 222; Attems, 1902: 559; Kanellis, 1959: 19; Zapparoli, 2002: 63); Kissamos (type locality of Scolopendra cretica Lucas, 1853) (Lucas, 1853: 529; Raulin, 1856; Kanellis, 1959: 19; Zapparoli, 2002: 63); Kolimbari (Chamberlin, 1956: 51; Zapparoli, 2002: 63); environs of Kournas Lake (Zapparoli, 2002: 63); Omalos (type locality of Scolopendra oraniensis lusitanica var. cretica Attems, 1902) (Cecconi, 1895: 193; Attems, 1902: 559; Kanellis, 1959: 19; Zapparoli, 1994: 26; 2002: 63); Omalos, 1000 m (Zapparoli, 2002: 63); Paleochora, Maniationa (Zapparoli, 2002: 63); Stomio (Matic and Stavropoulos, 1988: 34; Zapparoli, 2002: 63); Gavdos Island (Dobroruka, 1977: 164; Zapparoli, 2002: 63). Rethymno: road Gonies Anogia, m (Zapparoli, 1994: 26; 2002: 34); Melidoni (Karsch, 1888: 222; Kanellis, 1959: 19; Matic, 1980: 99; Zapparoli, 2002: 63); Oros Idi (Dobroruka, 1977: 164; Zapparoli, 2002: 34); Oros Idi, m (Zapparoli, 1994:

18 384 S. SIMAIAKIS ET AL. Isr. J. Zool. 26; 2002: 63); Oros Idi, 1600 m (Zapparoli, 1994: 26; 2002: 63); Oros Idi, slope of Anogia, 900 m (Zapparoli, 2002: 63); Oros Idi, slope of Anogia, 1500 m (Zapparoli, 2002: 63); Rethymno (Cecconi, 1895: 193; Kanellis, 1959: 19; Zapparoli, 2002: 63); 8 km W of Rethymno (Zapparoli, 2002: 63). Irakleio: Karteros (Chamberlin, 1956: 51; Zapparoli, 2002: 63); Dafnes (type locality of Scolopendra oraniensis lusitanica var. cretica Attems, 1902) (Attems, 1902: 559; Kanellis, 1959: 19; Zapparoli, 2002: 63); Gonies, 300 m (Zapparoli, 1994: 26; 2002: 63); beach E Irakleio, 0 m (Zapparoli, 1994: 26; 2002: 63); karst area 15 km between Irakleio and Chania (Zapparoli, 2002: 63); 10 km S of Irakleio (Zapparoli, 2002: 63); Kamilari Cave (near Koubbedes) (Zapparoli, 1994: 26; 2002: 63); Knossos (Matic, 1980: 99; Zapparoli, 2002: 63); beach of Timbaki (Zapparoli, 1994: 26; 2002: 63); Viannos, Kato Simi (Zapparoli, 2002: 34). Lasithi: Agios Nikolaos (Zapparoli, 1994: 21; 2002: 63); Agios Nikolaos, Ammoudara, 50 m from the seashore, phrygana (Zapparoli, 2002: 63); near Agios Nikolaos, Flamouriana (Zapparoli, 2002: 63); Agios Nikolaos, Itanos (Zapparoli, 2002: 63); Gournia (Zapparoli, 2002: 63); between Kroustas and Prina (Zapparoli, 2002: 63); Ierapetra, Males, Sarakina Gorge, 200 m (Zapparoli, 2002: 63); track to Kato Metochi Kastamonitsa, m (Zapparoli, 2002: 63); Kritsa (Zapparoli, 2002: 63); environs of Kritsa (Zapparoli, 2002: 63); Lato (Zapparoli, 2002: 63); between Marathospilia and Psihro (Chamberlin, 1956: 51; Zapparoli, 2002: 63); Milatos (Chamberlin, 1956: 51; Zapparoli, 2002: 63); Psihro (Chamberlin, 1956: 51; Zapparoli, 2002: 63); Pirgos (Milatos) (Chamberlin, 1956: 51; Zapparoli, 2002: 63); Selinari (Zapparoli, 2002: 63); Sitia (type locality of Scolopendra cretica Lucas, 1853) (Lucas, 1853: 529; Raulin, 1856; Kanellis, 1959: 19; Zapparoli, 2002: 63); above Tzermiado, m (Zapparoli, 2002: 63); Vrouchas (Zapparoli, 2002: 63); UNCERTAIN DATA Chaui [or Chani?] Koku (Zapparoli, 2002: 63); Vivari (Attems, 1902: 559; Kanellis, 1959: 19; Zapparoli, 2002: 63). Chaui Koku is the locality Kokkini Chani next to Irakleio whilst the locality Vivari possibly referred to the village Vizari at Rethymno. CRETE: Chania: 1 (a 2); 5 (a 3 ); 6 (a 1 ; b 2 ; c 2 ; d 2 2 ; e 2 1 ; f 5 ; g 2 1 ; h 1 3 ; i 2 ); 8 (b 1 ; c 1 1 ; d 3 1 ; e 1 ; f 4 2 ; g 4 2 ; h 2 1 ); 14 (a 5 4 ); 15 (a 19); 16 (a 2 ; c 1 ); 17 (a 1 ; c 1 1 ; d 1 1 ; 18 (d 1 ); 19 (a 2 ); 20 (a 2; c 3; d 3); 21 (c 2; d 1); 22 (d 2); 23 (a 1; c 1; d 2); 24 (d 4); 25 (a 1 1 ; c 1 ); 27 (b 1 2 ; c 1 ). Rethymno: 29 (a 8 3 ; b 3 3 ; c 1 ; d 1 ; f 1 1 ); 30 (c 4 2 ; d 1 ; e 2 ); 31 (a 1 ); 32 (b 2 ); 34 (a ; b ); 35 (a 1 ; e 3 2 ); 39 (b 1 ); 40 (c 9); 42 (a 1 4 ). Irakleio: 46 (a 3); 47 (a 3 3 ); 48 (a 21); 49 (a 6); 51 (a 1 1 ); 52 (a 3 3 ); 54 (a 6); 55 (a 1 3 ); 56 (b 6 2 ); 57 (c 1 1 ); 58 (a 1 1 ); 59 (a 5 1 ; b 18 6 ; c 8 4 ); 60 (b 4 4 ; c 6 3 ). Lasithi: 61 (a 1 ); 62 (a 6 9 ); 63 (c 1 1 ); 64 (d 1 ); 66 (a 3 ); 67

19 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 385 (a 8 3 ); 68 (a 2 1 ); 69 (a 16); 70 (a ); 72 (a 10 8 ; b 15 4 ); 73 (a 1); 74 (a 1 ); 75 (a 4 ; b 6 1 ; c 3 2 ); 76 (a ); 77 (a 1 2 ); 78 (a 1 ; e 6 1 ); 79 (a 13 9 ); 80 (a 14 8 ); 81 (a 6 2 ); 82 (a 10 4 ); 83 (a 6 2 ); 84 (a 5 7 ); 85 (a 4 2 ); 86 (a 8 4 ); 87 (a 6 5 ). Cretan (Fig. 5). Altitudinal range: m. Highest site: 35. Crete. It is also recorded from Cyrenaica, Turkey, West Libya (Zapparoli, 2002). The specimens from Cyrenaica, Turkey, West Libya are possibly Scolopendra canidens Newport, Cretan endemic East Mediterranean affinities. Scolopendra cretica is the most common scolopendromorph species on Crete. It has been caught at many sites from western to eastern Crete, up to the timberline (1650 m). Both males and females are found during the whole year. The activity peak of males is in Fig. 5. Cryptops anomalans, C. beroni, C. kosswigi, C. hortensis, C. trisulcatus, and Scolopendra cretica distribution on Crete.

20 386 S. SIMAIAKIS ET AL. Isr. J. Zool. spring and early summer while females are found in late spring and early summer. It is widespread in environments modified by man, in agroecosystems, in typical phrygana with Coridothymus capitatus, Phlomis lanata, Satureja thymbra, Salvia fruticosa, Genista acanthoclada, Erica sp. and Cistus sp., in mixed phrygana and maquis ecosystems or maquis dominated by Pistacia lentiscus, Quercus ilex, Q. coccifera, Olea europea, and ferns, in plateaus consisting of prostrate shrubs, mainly Juniperus oxycedrus, Berberis cretica, Prunus prostrata, and Satureja spinosa, in subalpine and alpine places with Berberis cretica, Prunus prostrata, Astragalus angustifolius, and Satureja spinosa, and in Pinus brutia, Cupressus sempervirens, and Q. coccifera forests as well as on sand dunes dominated by J. macrocarpa. The habitat preferences of Scolopendra cretica are quite similar to those of Lithobius nigripalpis. CRYPTOPIDAE Kohlrausch, 1881 Cryptops Leach, Cryptops anomalans Newport, 1881 CRETE (Attems, 1903a; Verhoeff, 1925: 659; Zapparoli, 1993: 95; 2002: 66; Stoev, 1997: 102); Chania: Akrotiri, Moni Gouvernetou, Katholiko Cave (Chamberlin, 1956: 51; Zapparoli, 2002; 66); Aselakia (Attems, 1902: 571; Kanellis, 1959: 20; Zapparoli, 2002: 66); Askifou (Zapparoli, 2002: 65); Kolimbari (Chamberlin, 1956: 51; Zapparoli, 2002: 66); Mournies (Attems, 1902: 571; Kanellis, 1959: 20; Zapparoli, 2002: 66); Nerokouros (Attems, 1902: 571; Kanellis, 1959: 20; Zapparoli, 2002: 66). Rethymno: Rethymno (Attems, 1902: 571; Kanellis, 1959: 20; Zapparoli, 2002: 66). Irakleio: Dafnes (Attems, 1902: 571; Kanellis, 1959: 20; Zapparoli, 2002: 66); Labyrinthes (type locality of Cryptops punctatus labyrinthiaca Attems, 1902) (Attems, 1902: 571; 1930: 223; Kanellis, 1959: 20; Zapparoli, 2002: 66). CRETE: Chania: 6 (3 ); 11; 15. Rethymno: 30; 35; 43. Irakleio: 56. West Central Cretan (Fig. 5). Altitudinal range: m. Highest site: 11. Geographical distribution: Mainland and insular Greece, Albania, Algeria, Austria, Belgium, Bosnia Herzegovina, Bulgaria, Crimea, Croatia, European Turkey, former Czechoslovakia, France, FYR Macedonia, Germany, Hungary, Italy, Montenegro, Morocco, Netherlands, Romania, Serbia, Slovenia, South Ukraine, Spain, Switzerland, Tunisia (Zapparoli, 2002). Southwest Palearctic. Cryptops anomalans was collected on mountainous, subalpine, and alpine zones from western to central Crete, up to 2000 m. It is not very common in Crete. It has been found in a forest of Pinus brutia with very little undergrowth, consisting mainly of

21 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 387 Quercus coccifera, and in subalpine and alpine valleys with very sparse vegetation of Berberis cretica, Prunus prostrata, Astragalus angustifolius, and Satureja spinosa, as well as in phrygana with Coridothymus capitatus, Sarcopoterium spinosum, Phlomis fruticosa, and Urginea maritima. 17. Cryptops beroni Matic and Stavropoulos, 1988 CRETE (Zapparoli, 1993: 95; 2002: 66; Matic and Stavropoulos, 1993: 104; Stoev, 1997: 102): Chania: Akrotiri, Moni Gouvernetou, Katholiko Cave, 60 m (type locality) (Chamberlin, 1956: 51; Matic and Stavropoulos, 1988: 34; Zapparoli, 1994: 23; 2002: 66; Stoev and Beron, 2000: 95). West Cretan (Fig. 5). Altitudinal range: m. Highest site: Gouverneto Cave. Crete (Zapparoli, 2002). Cretan endemic South European affinities. Cryptops beroni is only known from the type locality (Katholiko Cave), therefore its habitat preferences are not so certain. 18. Cryptops hortensis (Donovan, 1810) CRETE: Chania: Georgioupoli, in olive grove (Zapparoli, 2002: 70). Lasithi: Agios Nikolaos (Zapparoli, 1994: 23; 2002: 70). CRETE: Irakleio: 55 (a 1). Cretan (Fig. 5). Altitudinal range: m. Highest site: 55. Mainland and insular Greece, Albania, Armenia, Austria, Azerbaijan, Azores, Bosnia Herzegovina, Bulgaria, Canary Is., Corsica, Croatia, Denmark, European Russia, former Czechoslovakia, France, FYR Macedonia, Georgia, Germany, Hungary, Iberia, Ireland, Italy, Madeira, Montenegro, Morocco, Netherlands, North Turkey, Norway, Poland, Romania, Sardinia, Sicily, Slovenia, Switzerland, Tadzikistan, Turkmenistan, Ukraine, United Kingdom, Uzbekistan (Zapparoli, 2002). Central and West Palearctic. This species has only been recorded from three places on Crete, without any specific habitat notes.

22 388 S. SIMAIAKIS ET AL. Isr. J. Zool. 19. Cryptops kosswigi Chamberlin, 1952 CRETE (Zapparoli, 1993: 95; 2002: 71; Stoev, 1997: 102): Chania: Georgioupoli, in phrygana, pasture (Zapparoli, 2002; 71); Georgioupoli, in olive grove (Zapparoli, 2002; 71); near Georgioupoli, phrygana, pasture (Zapparoli, 2002; 71). Rethymno: Oros Idi, m (Zapparoli, 1994: 24; 2002: 71); Oros Idi, m (Zapparoli, 1994: 24; 2002: 71). Lasithi: Agios Nikolaos (Zapparoli, 1994: 24; 2002: 71). CRETE: Chania: 6 (j 1). Cretan (Fig. 5). Altitudinal range: m. Highest site: Oros Idi. Geographical distribution: Insular Greece, Palestine, Southwest Turkey (Zapparoli, 2002). East Mediterranean. Cryptops kosswigi has been collected on lowlands and mountainous areas from western to central Crete. It is not very common in Crete. It was found in agroecosystems (olive groves), in typical mediterranean phrygana, and in forests of Pinus brutia with very little undergrowth mainly of Quercus coccifera. 20. Cryptops trisulcatus Brölemann, 1902 CRETE: Lasithi: between Agios Nikolaos and Ammoudara, in phrygana, 50 m from sea shore (Zapparoli, 2002: 73). CRETE: Chania: 16 (c 1). Irakleio: 50 (a 1); 55 (a 1). Lasithi: 79 (a 1); 82 (a 1). Cretan (Fig. 5). Altitudinal range: m. Highest site: 16. Insular Greece, Algeria, Balearic Is., Canary Is., Corsica, Italy, Portugal, Romania, Sardinia, Sicily, South France, Southeast Turkey, Spain (Zapparoli, 2002). Mediterranean. Cryptops trisulcatus is common from western to eastern Crete, up to 700 m. It was collected in mountainous places dominated by ferns and Platanus orientalis, and in Pinus brutia forests as well as in cultivated lands.

23 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 389 GEOPHILOMORPHA Leach, 1815 HIMANTARIIDAE Cook, 1895 Bothriogaster Sseliwanoff, 1879 Bothriogaster signata (Kessler, 1874) CRETE (Attems, 1903b: 181; 1926: 239; Verhoeff, 1925: 629; Matic and Stavropoulos, 1988: 33; 1993: 103; Zapparoli, 1993: 95; 2002: 79); Chania: Akrotiri, Moni Gouvernetou, Arkoudas Cave (Chamberlin, 1956: 52; Zapparoli, 2002: 81); Akrotiri, Moni Gouvernetou, Katholiko Cave (Chamberlin, 1956: 52; Zapparoli, 2002: 81); Askifou (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); Gallos (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); Georgioupoli, at hill, phrygana, pasture, m (Zapparoli, 2002: 81); Georgioupoli, (Zapparoli, 2002: 81); Imbros Gorge (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); near Kournas Lake (Zapparoli, 2002: 81); Lakki (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); Mournies (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); Nerokouros (type locality of Bothriogaster Thesei Attems, 1902) (Attems, 1902: 579; 1929b: 49; Kanellis, 1959: 10; Zapparoli, 2002: 81); Niatos Askifou (Zapparoli, 2002: 81). Gavdos Island (Dobroruka, 1977: 164). Rethymno: road Gonies Anogia, m (Zapparoli, 1994: 13; 2002: 81); Melidoni (Karsch, 1888: 221); Oros Idi (Dobroruka, 1977: 164; Zapparoli, 2002: 81); Oros Idi, slope of Anogia, 900 m (Zapparoli, 2002: 81). Irakleio: Karteros (Chamberlin, 1956: 52; Zapparoli, 2002: 81); karst area near Irakleio (Zapparoli, 2002: 81); Kasteli-Viannos, Afrati, in olive grove, 500 m (Zapparoli, 2002: 81). Lasithi: Agios Nikolaos, Ammoudara, 50 m near sea shore, phrygana (Zapparoli, 2002: 81); Agios Nikolaos, Ammoudara, 50 m, phrygana (Zapparoli, 2002: 81); Agios Nikolaos, Flamouriana (Zapparoli, 2002: 81); Agios Nikolaos, Katharo Plateau, direction Kritsa, karstic terrain, m (Zapparoli, 2002: 81); Gournia (Zapparoli, 2002: 81); near Kritsa (Zapparoli, 2002: 81); Kritsa (Zapparoli, 2002: 81); Kronio Cave (Zapparoli, 2002: 81); between Marathospilia and Psihro (Chamberlin, 1956: 52; Zapparoli, 2002: 81); Lasithi, direction Tzermiado, m (Zapparoli, 2002: 81); Lasithi, direction Tzermiado, Agios Timios Stavros, m (Zapparoli, 2002: 81); Lato (Zapparoli, 2002: 81); Selinari (Zapparoli, 2002: 81); Sitia (Zapparoli, 2002: 81); Neapoli, Moni Kremaston (Zapparoli, 2002: 81); Neapoli (Zapparoli, 1994: 45; 2002: 81); Sitia (Zapparoli, 2002: 81); Tzermiado, Trapezas Cave (Eason, 1990: 257; Zapparoli, 2002: 81); Vrouchas (Zapparoli, 2002: 81). UNCERTAIN DATA Chaui [or Chani?] Koku (Zapparoli, 2002: 81). Chaui Koku is the locality Kokkini Chani near Irakleio.

24 390 S. SIMAIAKIS ET AL. Isr. J. Zool. Mainland and insular Greece, Albania, Bulgaria, Caucasus, Cyprus, Cyrenaica, Egypt, FYR Macedonia, Iran, Iraq, Israel, Jordan, Palestine, Saudi Arabia, Syria, Tripolitania, Tunisia, Turkestan, Turkey, Uzbekistan (Zapparoli, 2002). Balkan East Mediterranean West Asiatic. REMARKS The true status of the forms treated here as subspecies, following tradition, requires further study. In Crete, indeed, a taxonomic distinction between them could be justified, but a sound assessment of the question will require a revision of the genus in the whole Eastern Mediterranean area. 21. Bothriogaster signata graeca Verhoeff, 1901 CRETE: Chania: 1 (a 1 ); 3 (a 1 1 ); 5 (a 1 ). Rethymno: 31 (a 1 ); 34 (a 4 3 ; b 2 ); 35 (f 2 1 ); 39 (b 2 1 ; d 1 1 ); 42 (a 1 ). Irakleio: 45 (a 1 ); 48 (a 1 ) 52 (a 1 ); 55 (a 3 ). Lasithi: 62 (a 1 2 ); 66 (a 2 2 ); 70 (a 7 ); 72 (a 1 ); 76 (a 1 1 ); 77 (a 1 ); 79 (a 2 1 ); 80 (a 1 ); 82 (a 1 ); 83 (a 1 1 ); 84 (a 1 1 ); 86 (a 2 2 ); 87 (a 4 3 ). Cretan (Fig. 6). Altitudinal range: m. Highest site: 35. Northeast Mediterranean. Bothriogaster signata graeca was found mainly from central to eastern Crete, up to 1650 m, while it was almost absent from the westernmost areas. Adults are found in autumn and in early winter. It has been caught in mountainous places with Pinus brutia, in cultivated lands, and in mediterranean phrygana dominated by Phlomis lanata, Sarcopoterium spinosum, Calicotome villosa, and Satureja thymbra as well as in subalpine and alpine phrygana. 22. Bothriogaster signata thesei Attems, 1902 CRETE: Chania: 2 (a 1 ); 14 (a 1 ); 16 (a 1 ; c 2 8 ). Rethymno: 36 (f 1 ); 43 (b 2 4 ). Irakleio: 55 (a 1 ). Lasithi: 61 (a 1 ); 69 (a 1 3 ). Cretan (Fig. 6). Altitudinal range: m. Highest site: 36. Cretan endemic East Mediterranean affinities.

25 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 391 Fig. 6. Bothriogaster s. graeca and Bothriogaster s. thesei distribution on Crete. Bothriogaster signata thesei is widespread from western areas to central places of Crete but rarely recorded from easternmost areas. It was collected in mountainous sites with Cupressus sempervirens, in coastal phryganic ecosystems, in phrygana dominated by Coridothymus capitatus, Sarcopoterium spinosum, Calicotome villosa, and Satureja thymbra, and in alpine shrubs with Berberis cretica, Astragalus angustifolius, and Satureja spinosa as well as at sites with Platanus orientalis. DIGNATHODONTIDAE Cook, 1895 Dignathodon Meinert, Dignathodon microcephalus (Lucas, 1846) CRETE (Zapparoli, 1993: 95; Stoev, 1997: 103; Zapparoli, 2002: 82): Chania: Akrotiri, Moni Gouvernetou, Arkoudas Cave (Chamberlin, 1956: 52; Zapparoli, 2002: 82); Akrotiri, Moni Gouvernetou, Koutrouli Cave (Chamberlin, 1956: 52; Zapparoli, 2002: 82). Rethymno: Oros Idi, slope of Anogia, 900 m (Zapparoli, 2002: 82). Lasithi: Sitia (Zapparoli, 2002: 82). CRETE: Chania: 3 (a 1 ; d 1 ); 10 (e 1 ); 11 (a 1 ); 18 (c 1 ); 20 (b 1 ); 27 (b 1 ). Rethymno: 30 (c 1 ); 40 (c 1 ). Irakleio: 52 (a 1 ); 55 (a 1 ). Lasithi: 63 (b 2 ; e 1 ); 64 (d 1 ); 65 (a 1 1 ); 68 (a 1 ); 72 (a 2 1 ); 74 (b 1 ); 75 (b 1 ; d 1 ); 78 (c 1 ); 79 (a 3 ); 83 (a 1 ); 86 (a 1 2 ).

26 392 S. SIMAIAKIS ET AL. Isr. J. Zool. Cretan (Fig. 9). Altitudinal range: m. Highest site: 11. Mainland and insular Greece, Albania, Algeria, Austria, Balearic Is., Bosnia Herzegovina, Bulgaria, Corsica, Crimea, Croatia, former Czechoslovakia, Italy, Montenegro, Morocco, Near and Middle East, Portugal, Romania, Sardinia, Serbia, Sicily, South France, Spain, Tunisia (Zapparoli, 2002). Balkan Mediterranean. The peak activity of males occurs during the wet period of autumn. It is common in Pinus brutia forests, in plateau and open alpine formations with Satureja spinosa, Berberis cretica, and Genista acanthoclada, as well as in phryganic ecosystems with Sarcopoterium spinosum, Verbascum spinosum, Euphorbia acanthothamnos, and Drimia maritima in mixed phrygana and maquis dominated by Coridothymus capitatus, Pistacia lentiscus, Pinus brutia, Juniperus phoenicea, and J. macrocarpa. Henia C. L. Koch, 1847 Subgenus Henia C. L. Koch, Henia (Henia) athenarum Pocock, 1891 CRETE (Verhoeff, 1925: 659; Minelli, 1982: 261; Zapparoli, 1993: 95; 2002: 83; Matic and Stavropoulos, 1993: 103; Stoev, 1997: 103): Chania: Omalos (type locality of Henia Idomenei Attems, 1902) (Attems, 1902: 575; 1903b: 275; 1929a: 235; Kanellis, 1959: 15; Minelli, 1982: 261; Zapparoli, 2002: 83). West Cretan (Fig. 7). Altitudinal range: 1100 m. Highest site: Omalos. Mainland Greece, Crimea, Caucasus, North Turkey (Zapparoli, 2002). Northeast Mediterranean. Based on literature data Henia athenarum should be present in Cupressus sempervirens, Pinus brutia, Quercus coccifera, or Acer sempervirens forests. 25. Henia (Henia) devia C. L. Koch, 1847 CRETE (Verhoeff, 1925: 629, 659; Minelli, 1982: 261; Zapparoli, 1993: 95; 2002: 84; Stoev, 1997: 104): Chania: Omalos, 1000 m (Attems, 1902: 574; 1903b: 275; 1929a: 233; Kanellis, 1959: 14; Zapparoli, 2002: 84).

27 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 393 CRETE: Lasithi: 82 (a 1 ). West East Cretan (Fig. 7). Altitudinal range: m. Highest site: Omalos. Mainland and insular Greece, Albania (Zapparoli, 2002). Balkan. Habitat requirements probably similar to Henia athenarum. Henia devia was also collected in a cave. Subgenus Scotophilus Meinert, Henia (Scotophilus) bicarinata Meinert, 1870 CRETE (Zapparoli, 1993a: 95; 2002: 88): Rethymno: Oros Idi, slopes of Anogia, 900 m (Zapparoli, 2002: 88). Lasithi: Sitia (Zapparoli, 2002: 88). Fig. 7. Henia devia, H. pulchella, H. bicarinata, and H. athenarum distribution on Crete.

28 394 S. SIMAIAKIS ET AL. Isr. J. Zool. CRETE: Rethymno: 29b (a 2 1 ). Cretan (Fig. 7). Altitudinal range: m. Highest site: Anogia. Mainland and insular Greece, Bosnia Herzegovina, Bulgaria, Caucasus, Corsica, Croatia, France, Hungary, Iberia, Italy, Macaronesia, Maghreb, Sardinia, Sicily, Turkey (Zapparoli, 2002). Central European Mediterranean. It is quite common on lowlands of Crete from western to easternmost areas reaching altitudes of about 900 m. It is recorded from sparse phrygana dominated by Coridothymus capitatus, Phlomis lanata, Satureja thymbra, Salvia fruticosa, Genista acanthoclada, Erica sp., and Cistus sp. 27. Henia (Scotophilus) pulchella Meinert, 1870 CRETE (Verhoeff, 1925: 659; Attems, 1929a: 234; 1929c: 298): Chania: Askifou (Attems, 1902: 573; 1903b: 275; Kanellis, 1959: 14). CRETE: Lasithi: 71 (a 1 ); 72 (a 1 ); 76 (a 1 1 ); 84 (a 2 ); 86 (a 2 ). Cretan (Fig. 7). Altitudinal range: m. Highest site: Askifou. Insular Greece, former Yugoslavia, Italy (Kanellis, 1959). Balkan Central-North Mediterranean. REMARKS The relationships between this species and H. bicarinata require further study. In Crete, indeed, a taxonomic distinction between them seems to be justified, but a sound assessment of the question will require a revision of the genus in the whole Mediterranean area. It is collected on lowlands of western Crete as well as on eastern islets reaching altitudes of about 800 m. It is recorded from sparse coastal, mountainous phrygana and sand dunes.

29 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 395 SCHENDYLIDAE Verhoeff, 1908 Haploschendyla Verhoeff, Haploschendyla europaea (Attems, 1903) CRETE: Chania: 17 (b 1 ). Rethymno: 28 (a 1 ). Irakleio: 48 (a 1 ); 49 (a 1 ); 52 (a 1 ). Lasithi: 82 (a 1 ). South Cretan (Fig. 8). Altitudinal range: m. Highest site: 49. Insular Greece (Corfu) (Zapparoli, 2002). First record for Crete. South Balkan. Both females and males are found in spring and in autumn. It occurs on south coastal phryganic ecosystems, in sparse phrygana dominated by Calicotome villosa, Sarcopoterium spinosum, and Urginea maritima, as well as in dense mixed phrygana and maquis dominated by Calicotome villosa, Sarcopoterium spinosum, Urginea maritima, Olea europea, and Pistacea lentiscus. Fig. 8. Haploschendyla europaea, Nannophilus ariadnae, N. eximius, and Schendyla nemorensis distribution on Crete.

30 396 S. SIMAIAKIS ET AL. Isr. J. Zool. Nannophilus Cook, Nannophilus ariadnae Attems, 1902 CRETE (Attems, 1903b: 194; Verhoeff, 1925: 659; Attems, 1929a: 70; Zapparoli, 1993: 95; 2002: 91; Matic and Stavropoulos, 1993: 103): Chania: Mournies (type locality) (Attems, 1902: 578; Kanellis, 1959: 11; Zapparoli, 2002: 91). CRETE: Chania: 3 (c 1 ). Lasithi: 80 (a 1 ); 88 (a 1 ); 89 (a 1 ). West and East Cretan (Fig. 8). Altitudinal range: 0 95 m. Highest site: Elafonisi. Mainland and insular Greece (Zapparoli, 2002). South Balkan. It was collected on southwest coastal phryganic ecosystems with Juniperus oxycedrus, Pistacea lentiscus, Coridothymus capitatus, and Ceratonia siliqua as well as on typical sand dunes. CRETE: Lasithi: 89 (a 1 ). 30. Nannophilus eximius (Meinert, 1870) East Cretan (Fig. 8). Altitudinal range: m. Highest site: 89. Mainland Greece, Macaronesia, North Africa, Sicily, South Italy (Zapparoli, 2002). First record for Crete. Mediterranean. No habitat data available. Schendyla Bergsoe and Meinert, Schendyla nemorensis (C. L. Koch, 1836) CRETE: Rethymno: Oros Idi, slope of Anogia, 1000 m (Zapparoli, 2002: 93). CRETE: Chania: 10 (j 1 ; k 1 ). Rethymno: 36 (c 5 3 ; d 1 ); 37 (a ; b 6 4 ). Lasithi: 65 (c 2 3 ; d 3 ).

31 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 397 Cretan (Fig. 8). Altitudinal range: m. Highest site: 37. Insular Greece, Algeria, Austria, Azores, Balearic Is., Bosnia Herzegovina, Bulgaria, Croatia, Finland, former Czechoslovakia, France, Germany, Ireland, Italy, Morocco, Netherlands, Norway, Poland, Romania, Sardinia, Sicily, Slovenia, Sweden, Tunisia, United Kingdom (Zapparoli, 2002). European Central and West Mediterranean. Adults, males and females, occur during the wet period of autumn. It was found on the three highest mountain massifs of Crete, in subalpine ecosystems with prostrate shrubs, mainly Juniperus oxycedrus oxycedrus, Berberis cretica, Prunus prostrata, Satureja spinosa, as well as in alpine valleys with very sparse vegetation of B. cretica, P. prostrata, Astragalus angustifolius, and S. spinosa. GEOPHILIDAE Cook, 1895 Pachymerium C. L. Koch, 1847 Pachymerium ferrugineum (C. L. Koch, 1835) CRETE (Verhoeff, 1925: 629, 658; Zapparoli, 1993: 95; 2002: 96; Stoev, 1997: 103); Chania: Chania (Karsch, 1888: 220; Kanellis, 1959: 17; Matic, 1980: 99; Zapparoli, 2002: 96); Georgioupoli, Kournas Lake, 100 m (Zapparoli, 2002: 96); Georgioupoli, in pasture, olive grove, at shore (Zapparoli, 2002: 96); Georgioupoli, m, phrygana, pasture at hill (Zapparoli, 2002: 96); Kournas Lake (Chamberlin, 1956: 52; Zapparoli, 2002: 96); Rethymno: Oros Idi, N slope, m (Zapparoli, 1994: 21; 2002: 96); Oros Idi, slope of Anogia, 1500 m (Zapparoli, 2002: 96). Irakleio: Karteros (Chamberlin, 1956: 52; Zapparoli, 2002: 96). Lasithi: Agios Nikolaos, Itanos (Zapparoli, 2002: 96). Mainland and insular Greece, Turkey, Alaska, Albania, Algeria, Austria, Azores, Balearic Is., Bosnia Herzegovina, Bulgaria, Canary Is., Caucasus, Central Sahara, Corsica, Croatia, Cyprus, Cyrenaica, European Russia, Finland, former Czechoslovakia, France, FYR Macedonia, Hungary, Iran, Italy, Latvia, Madeira, Morocco, Netherlands, Norway, Palestine, Poland, Portugal, Pribilof Is., Romania, Sardinia, Sicily, Slovenia, Spain, Tripolitania, Tunisia, Turkestan, United Kingdom (Zapparoli, 2002). Central and West Palearctic.

32 398 S. SIMAIAKIS ET AL. Isr. J. Zool. REMARKS The true status of the forms treated here as subspecies, following tradition, requires further study. In Crete, indeed, a taxonomic distinction between them could be justified, but a sound assessment of the question will require a revision of the whole complex. 32. Pachymerium ferrugineum ferrugineum (C. Koch, 1835) CRETE: Chania: 8 (j 1 ); 16 (a 1 ). Rethymno: 29 (d 1 ); 32 (a 2 ; e 1 ); 35 (c 1 ); 36 (a 1 ; c 1 2 ); 41 (a 2 ); 43 (a 1 ). Irakleio: 45 (a 3 2 ); 50 (a 1 2 ); 54 (a 1 ); 57 (a 1 ; b 2 ; c 1 ); 58 (a 2 ). Lasithi: 64 (d 1 2 ); 65 (c 1 ; d 2 1 ); 67 (a 1 ); 72 (a 1 ; b 1 ); 76 (a 4 1 ); 78 (c 2 ); 79 (a 2 3 ). Cretan (Fig. 9). Altitudinal range: m. Highest site: 36. Central and West Palearctic. Pachymerium f. ferrugineum is very common on Crete. It has been found at many sites from western to eastern Crete, up to 2000 m, but mainly between 1000 and 2000 m. Females occur during the period from late autumn to early spring while males are found Fig. 9. Pachymerium f. ferrugineum and Pachymerium f. insularum distribution on Crete.

33 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 399 in late autumn and in early winter. It is quite common in habitats modified by man, in Pinus brutia forests, in phryganic and maquis ecosystems with Coridothymus capitatus, Sarcopoterium spinosum, Pistacia lentiscus, and Nerium oleander as well as in mountainous areas dominated by Quercus coccifera, in subalpine and alpine phrygana. 33. Pachymerium ferrugineum insularum Verhoeff, 1902 CRETE: Chania: 3 (a 1 ; c 1 ; d 1 ; e 1 ); 15 (b 7 ; c 2 ); 17 (b 1 2 ; c 1 ); 22 (a 2 ; b 4 ; c 4 2 ); 26 (c 1 ). Lasithi: 80 (a 2 ); 83 (a 1 2 ); 85 (a 2 4 ); 86 (a 10 1 ); 87 (a 4 ). South Cretan (Fig. 9). Altitudinal range: m. Highest site: 15. Northeast Mediterranean. Pachymerium ferrugineum insularum is very common in the southernmost areas of Crete. It was found on the small satellite islands to the south of Crete (see Fig. 9) and on almost every coastal area around Crete (these records are recent and therefore not presented in Fig. 9). Adults occur in the spring as well as in late autumn and early winter. It prefers coastal phryganic areas and sand dunes. It was also collected on a plateau dominated by Berberis cretica, Genista acanthoclada, Sarcopoterium spinosum, Phlomis sp., and some Quercus coccifera and Acer sempervirens. Clinopodes C. L. Koch, Clinopodes flavidus C. L. Koch, 1847 CRETE (Verhoeff, 1925: 659; Zapparoli, 1993: 95; 2002: 99; Stoev, 1997: 103): Chania: Gallos (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); Georgioupoli (Zapparoli, 2002: 99); Imbros Gorge (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); Lakki (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); Mournies (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); Omalos (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); Vrises, Kournas Lake (Zapparoli, 2002: 99). Rethymno: Oros Idi (Dobroruka, 1977: 164; Zapparoli, 2002: 99); Oros Idi, m (Zapparoli, 1994: 18; 2002: 99); Oros Idi, slope of Anogia, 900 m (Zapparoli, 2002: 99); Oros Idi, slope of Anogia, 1500 m (Zapparoli, 2002: 99); Oros Idi, slope of Anogia, above 1500 m (Zapparoli, 2002: 99); Rethymno (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99). Irakleio: Dafnes (Attems, 1902: 572; Kanellis, 1959: 12; Zapparoli, 2002: 99); near Gonies (Zapparoli, 1994: 18; 2002: 99); Kastelli Viannos, Afrati, 500 m, olive grove (Zapparoli, 2002: 99). Lasithi: Agios Nikolaos, Ammoudara, 50 m near sea shore, phrygana (Zapparoli, 2002: 99); Exo Potami (Zapparoli, 2002: 99); Oros Thrypti,

34 400 S. SIMAIAKIS ET AL. Isr. J. Zool. Kato Chorio, 100 m (Zapparoli, 2002: 99); Oros Thrypti, Afentis Stavromenos, at top, 1476 m (Zapparoli, 2002: 99); Vidiatis, Agios Georgios (Zapparoli, 2002: 99). UNCERTAIN DATA Chaui [or Chani?] Koku (Zapparoli, 2002: 99). Chaui Koku is the locality Kokkini Chani next to Irakleio. CRETE: Chania: 13 (a 1 ); 15 (b 3 ; c 1 ); 16 (b 1 1 ; c 1 3 ). Rethymno: 34 (b 1 ); 35 (b 1 ; c 1 ); 36 (a 1 3 ; b 1 ; c 2 1 ; e 2 ; f 1 1 ); 37 (a 1 2 ; b 1 ); 38 (a 3 2 ); 43 (b 1 ). Irakleio: 50 (a 2 3 ); 51 (a 1 ); 54 (a 1 ). Lasithi: 58 (a 15 5 ); 62 (a 1 ); 64 (a 1 ); 65 (a 1 ; b 1 1 ); 72 (b 1 ). Cretan (Fig. 10). Altitudinal range: m. Highest site: 38. Mainland and insular Greece, Albania, Austria, Bosnia Herzegovina, Bulgaria, Caucasus, Crimea, Croatia, Cyprus, European Russia, former Czechoslovakia, FYR Macedonia, Italy, Montenegro, Palestine, Poland, Romania, Serbia, Sicily, Slovenia, Syria, Turkestan, Turkey (Zapparoli, 2002). Central and South European West Asiatic. Clinopodes flavidus was found from western to eastern Crete, up to 2400 m, mainly on the mountains. It occurs during the wet periods in autumn and in early spring. It has been caught in mountainous places with Pinus brutia and Quercus coccifera, in cultivated lands with olive trees, and in mediterranean phrygana dominated by Phlomis lanata, Sarcopoterium spinosum, Calicotome villosa, and Satureja thymbra, as well as in subalpine and alpine sparse phrygana at the edge of plateaus. Geophilus Leach, Geophilus carpophagus Leach, 1815 CRETE: Rethymno: Oros Idi, Psiloreitis, direction to Anogia, near Idaion Andron, next to Skinakas, 1400 m (Zapparoli, 2002: 101). Lasithi: Lato (Zapparoli, 2002: 101); ascent to Oros Selena, m (Zapparoli, 2002: 101). CRETE: Chania: 7 (a 1 ); 10 (g 1 ; h 1 ); 11 (g 1 ); 22 (a 1 ; b 1 ). Lasithi: 65 (a 1 ). Cretan (Fig. 10). Altitudinal range: m. Highest site: 11.

35 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 401 Fig. 10. Dignathodon microcephalus, Clinopodes flavidus, Geophilus insculptus, G. carpophagus, G. linearis, and Tuoba poseidonis distribution on Crete. Insular Greece, Albania, Algeria, Balearic Is., Belgium, Canary Is., Corsica, Croatia, Denmark, former Czechoslovakia, France, Germany, Ireland, Italy, Montenegro, Morocco, Netherlands, Norway, Palestine, Portugal, Romania, Sardinia, Sicily, Slovenia, Spain, Sweden, Tunisia, Turkey, United Kingdom (Zapparoli, 2002). European Mediterranean. Geophilus carpophagus was found from western to eastern Crete, up to 2000 m. It occurs mainly on plateaus above the timberline with prostrate shrubs, mainly Juniperus oxycedrus oxycedrus, Berberis cretica, Prunus prostrata, and Satureja spinosa. It has also been caught in subalpine and alpine phryganic ecosystems as well as on sand dunes. 36. Geophilus insculptus Attems, 1895 CRETE (Attems, 1903b: 227; Verhoeff, 1925: 659; Zapparoli, 1993: 95; 2002: 103; Matic and Stavropoulos, 1993: 103; Stoev, 1997: 103): Chania: Mournies (Attems, 1902: 572; Kanellis, 1959: 11; Zapparoli, 2002: 103).

36 402 S. SIMAIAKIS ET AL. Isr. J. Zool. West-central Crete (Fig. 10). Altitudinal range: 100 m. Highest site: Mournies. Mainland and insular Greece, Austria, Bulgaria, Corsica, Croatia, Denmark, European Russia, France, Germany, Hungary, Ireland, Italy, Montenegro, Morocco, Netherlands, Norway, Romania, Sardinia, Sicily, Slovenia, Sweden, Tunisia, United Kingdom (Zapparoli, 2002). European North and West Mediterranean. The habitat preferences of Geophilus insculptus in Crete are unknown. 37. Geophilus linearis C. L. Koch, 1835 CRETE: Zapparoli, 1993: 95. CRETE: Chania: 6 (i 1 ). Rethymno: 31 (a 1 ); 32 (b 3 ); 35 (d 2 ). Lasithi: 65 (c 1 ; d 1 ). Cretan (Fig. 10). Altitudinal range: m. Highest site: 65. Mainland and insular Greece, Austria, Bulgaria, Corsica, Croatia, Denmark, England, Finland, Germany, Italy, Netherlands, Poland, Romania, Sardinia, Slovenia, South France, Sweden, Turkey, Wales (Zapparoli, 2002). European North Mediterranean. Geophilus linearis was found from western to eastern Crete, up to 1750 m, mainly on the mountains. It was collected in mountainous places with Pinus brutia and Quercus coccifera as well as in subalpine and alpine sparse phrygana dominated by Satureja thymbra, Berberis cretica, and Astragalus angustifolius. 38. Geophilus naxius Verhoeff, 1901 CRETE (Zapparoli, 1993: 95; Stoev, 1997: 103; Zapparoli, 2002; 104): Rethymno: Oros Idi, m (Zapparoli, 1994: 20; 2002: 104). Lasithi: direction Tzermiado, Agios Timios Stavros, m (Zapparoli, 2002: 104); direction Tzermiado, m (Zapparoli, 2002: 104). CRETE: Chania: 15 (a 1 1 ). Rethymno: 31 (a 1 1 ); 32 (c 1 ). Irakleio: 58 (a 1 ). Lasithi: 67 (a 2 ); 71 (a 1 ); 72 (b 1 ); 76 (a 2 1 ).

37 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 403 Fig. 11. Geophilus naxius, G. pygmaeus, Insigniporus sturanyi, and Strigamia acuminata distribution on Crete. Cretan (Fig. 11). Altitudinal range: m. Highest site: Oros Idi. Mainland and insular Greece, Palestine, South Turkey (Zapparoli, 2002). Northeast Mediterranean. In Crete, Geophilus naxius occurs mainly in altitudes of about 1500 m. Adults are found in late autumn and in early winter. It was collected on the western and central mountain massifs, in open valleys with Berberis cretica, Genista acanthoclada, Sarcopoterium spinosum, and Phlomis sp. together with some Quercus coccifera and Acer sempervirens. It was also found among riparian vegetation with Nerium oleander, Coridothymus capitatus, and Calicotome villosa. 39. Geophilus pygmaeus Latzel, 1880 CRETE: Attems, 1902: 572; Verhoeff, 1925: 659. CRETE: Irakleio: 50 (a 2 ).

38 404 S. SIMAIAKIS ET AL. Isr. J. Zool. Central Cretan (Fig. 11). Altitudinal range: 200 m. Highest site: 50. Bosnia Herzegovina, Croatia, Germany, Serbia, Slovenia (Verhoeff, 1925; Stoev, 1997). Balkan Central European. Geophilus pygmaeus was only collected in a dense pine forest. Tuoba Chamberlin, Tuoba poseidonis (Verhoeff, 1901) CRETE: Rethymno: 29b (a 11 8 ). Central Cretan (Fig. 10). Altitudinal range: 0 5 m. Highest site: 29b. Mainland and insular Greece, Corsica, Egypt (Red Sea), Italy, Jordan (Death Sea), Sardinia, Sicily, Slovenia, Somalia (Indian ocean), South France (Zapparoli, 2002). First record for Crete. Mediterranean South West Asiatic. It was collected only on sandy seashore with Posidonia sp., Tamarix sp., and the common reed Phragmites sp. Insigniporus Attems, Insigniporus sturanyi Attems, 1903 CRETE: Chania: 15 (a 1 ); 16 (a 1 1 ). West Cretan (Fig. 11). Altitudinal range: m. Highest site: 15. North mainland Greece, FYR Macedonia (Zapparoli, 2002). First record for Crete. Balkan.

39 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 405 It was only collected from two localities on the western mountains of Crete, in alpine shrubs (Berberis cretica, Genista acanthoclada, Sarcopoterium spinosum, Phlomis sp.) with some Quercus coccifera and Acer sempervirens as well as among ferns and Platanus orientalis. LINOTAENIIDAE Cook, 1904 Strigamia Gray, Strigamia acuminata (Leach, 1814) CRETE: Chania: 11 (c 3 1 ; g 1 1 ); 19 (b 1 ). Rethymno: 35 (d 1 ); 36 (a 1 ; d 2 1 ); 38 (a 1 ). Irakleio: 57 (a 1 ). Lasithi: 65 (c 2 1 ; d 3 3 ). Cretan (Fig. 11). Altitudinal range: m. Highest site: 38. Balkans, Caucasus, Europe, Turkey (Kos, 1992; Zapparoli, 1999). First record from Greece and Crete. European Northeast Mediterranean. Strigamia acuminata is widespread in the subalpine and alpine zone from western to eastern Crete, at altitudes between 1200 and 2400 m. The peak of activity in both males and females is from late autumn to early winter. It occurs in phrygana dominated by Berberis cretica, Astragalus angustifolius, and Satureja spinosa. It was also collected on the island of Gavdos in phrygana dominated by Sarcopoterium spinosum, Coridothymus capitatus, and sparse maquis with Pistacia lentiscus, Pinus brutia, Juniperus phoenicea, and J. macrocarpa. DISCUSSION Forty-two taxa are reported, belonging to 17 genera and 9 families. One species (Strigamia acuminata) is recorded for the first time from Greece while 6 additional species (Lithobius lucifugus, Lithobius pamukkalensis, Haploschendyla europea, Nannophilus eximius, Tuoba poseidonis, and Insigniporus sturanyi) are recorded for the first time from Crete. Nannophilus eximius is the only species captured on Grantes islet but not on Crete. Of the taxa examined, 36% have Mediterranean affinities s. l. (Fig. 12). Seven of them are Mediterranean, six are northeast Mediterranean, and two have an east Mediterranean distribution (Table 1). 25% have European affinities s. l. (Fig. 12). Two have a

40 406 S. SIMAIAKIS ET AL. Isr. J. Zool. Fig. 12. Chorotypes of Crete centipede species. BAL: Balkanic, EME: east Mediterranean, EN: endemic, EUR: European, MED: Mediterranean, NEM: northeast Mediterranean, PA: Palearctic, SEU: south European, WA: west Asiatic. Table 1 Chorotype and distribution of centipede taxa on Crete. BAL: Balkanic, BAL_CNM: Balkanic central-north Mediterranean, BAL_EME: Balkanic east Mediterranean, BAL_MED: Balkanic Mediterranean, BAL_SEU: Balkanic south European, CEU_MED: central European Mediterranean, CSE_WA: central-south European west Asiatic, CWP: central-west Palearctic, EME: east Mediterranean, EN_EME: endemic with east Mediterranean affinities, EN_SEU: endemic with south European affinities, EUR: European, EUR_CWM: European central-west Mediterranean, EUR_MED: European Mediterranean, EUR_NEM: European northeast Mediterranean, EUR_NME: European north Mediterranean, EUR_NWM: European northwest Mediterranean, EUR_WA: European west Asiatic, MED: Mediterranean, MED_SWA: Mediterranean southwest Asiatic, NEM: northeast Mediterranean, NEM_WA: northeast Mediterranean west Asiatic, SBA: south Balkanic, SEU_MED: south European Mediterranean, SEU_NEM: south European northeast Mediterranean, SWP: southwest Palearctic, WPA: west Palearctic Species Chorotype Distribution on Crete Scutigera coleoptrata SEU_MED Cretan Eupolybothrus litoralis BAL_EME Cretan Lithobius agilis EUR West Central Cretan Lithobius cretaicus EN_SEU Central Cretan Lithobius creticus EN_SEU Cretan Lithobius erythrocephalus EUR_WA Cretan Lithobius lucifugus EUR_WA Cretan Lithobius nigripalpis NEM Cretan Lithobius piceus EUR Central Cretan Lithobius viriatus NEM_WA Cretan Lithobius micropodus SEU_NEM Central Cretan Lithobius aeruginosus EUR_NEM West Cretan Lithobius crassipes WPA West Central Cretan Lithobius pamukkalensis NEM East Cretan Scolopendra cretica EN_EME Cretan Cryptops anomalans SWP West Central Cretan

41 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 407 Table 1 continued Species Chorotype Distribution on Crete Cryptops beroni EN_SEU West Cretan Cryptops hortensis CWP Cretan Cryptops kosswigi EME Cretan Cryptops trisulcatus MED Cretan Bothriogaster s. graeca NEM Cretan Bothriogaster s. thesei EN_EME Cretan Dignathodon microcephalum BAL_MED Cretan Henia athenarum NEM West Cretan Henia devia BAL West East Cretan Henia bicarinata CEU_MED Cretan Henia pulchella BAL_CNM Cretan Haploschendyla europea SBA South Cretan Nannophilus ariadnae SBA West East Cretan Nannophilus eximius MED East Cretan Schendyla nemorensis EUR_CWM Cretan Pachymerium f. ferrugineum SWP Cretan Pachymerium f. insularum NEM South Cretan Clinopodes flavidus CSE_WA Cretan Geophilus carpophagus EUR_MED Cretan Geophilus insculptus EUR_NWM West Central Cretan Geophilus linearis EUR_NME Cretan Geophilus naxius NEM Cretan Geophilus pygmaeus BAL_SEU Central Cretan Tuoba poseidonis MED_SWA Central Cretan Insigniporus sturanyi BAL West Cretan Strigamia acuminata EUR_NEM Cretan south European distribution and nine are European (Table 1). Four species have a Palaearctic distribution s. l. (10%), two species have Asiatic affinities s. l., and five species are Balkanic s. l. (Table 1). Only four species (Lithobius cretaicus, Lithobius creticus, Scolopendra cretica, and Cryptops beroni) and one subspecies (Bothriogaster signata thesei) are considered endemic to Crete (Table 1). Besides, there are significative data concerning the zoogeographical affinities of Corsica, Sardinia, Sicily (Foddai et al., 1995) and Malta (Zapparoli, 2004: personal communication) (Table 2), whilst the centipede fauna of Cyprus is poorly known. Thus, in Crete, the dominant element is the Mediterranean s. l. accompanied by the European s. l. and with a marginal presence of the Balkanic, the Palearctic, and the Asiatic species. The same pattern is present in Sardinia, Sicily, and Cyprus, while in Corsica (Foddai et al., 1995) and Malta (Zapparoli, 2004: personal communication) the dominant element is the European s. l. accompanied by the Mediterranean s. l. Four species and one

42 408 S. SIMAIAKIS ET AL. Isr. J. Zool. subspecies are endemic to Crete (12%): more than those of Sicily and Cyprus (3.4% and 9.5%, respectively) but fewer than those of Corsica and Sardinia (18% and 14.5%, respectively) (Minelli and Zapparoli, 2004: personal communication) (Table 2). Most of the species and subspecies (54%) are widespread on Crete, from western to easternmost areas as well as from northern to southern localities. A few species (10%) are distributed only on western Crete, a few others (11%) on the central part of Crete, and a third group (10%) shows a wider range of distribution on western and central Crete. Lithobius pamukkalensis and Nannophilus eximius are only distributed on eastern Crete while Haploschendyla europea and Pachymerium f. insularum occur in southern Crete. Finally, two species (Henia devia and Nannophilus ariadnae) have been recorded from western and eastern localities on Crete (Fig. 13). Regarding the zoogeographical affinities of centipedes, three of the five endemics are widely distributed on the island (Lithobius creticus, Scolopendra cretica, and Bothriogaster signata thesei), and the other two (Lithobius cretaicus and Cryptops beroni) are recorded from central and western Crete, respectively (Fig. 14). The majority of species with European and Mediterranean affinities s. l. are widespread on Crete. Most of species referred from central Crete have European affinities (Fig. 14), while the Table 2 Number of taxa and endemic species among the Mediterranean islands Islands Species Endemic (%) Corsica 39 7 (17,9) Sardinia 55 8 (14.5) Sicily 59 2 (3.4) Malta 20 Crete 42 5 (11.9) Cyprus 21 2 (9.5) Fig. 13. Centipede distribution in Crete. CCR: central Crete, CRE: Crete, ECR: east Crete, SCR: south Crete, WCC: west central Crete, WCR: west Crete, WEC: west east Crete.

43 Vol. 50, 2004 THE CENTIPEDE FAUNA OF CRETE AND ITS SATELLITE ISLANDS 409 four species referred from western Crete are characterized by four different affinities. The two centipedes collected on eastern Crete (Lithobius pamukkalensis and Nannophilus eximius) have Mediterranean affinities (s. l.), despite the fact that the former has been quoted as a south Anatolian, Taurian endemic (Zapparoli, 2002). It is also remarkable that Haploschendyla europea and Pachymerium f. insularum, recorded from south Crete, have Mediterranean affinities s. l. (Fig. 14). As for the altitudinal distribution of centipedes, most of scolopendromorphs and the only scutigeromorph species are present from coastal and lowland valleys to subalpine phrygana (Fig. 15). For instance, Scutigera coleoptrata, Scolopendra cretica, and Cryptops kosswigi occur up to 1650 m, while Cryptops anomalans reaches altitudes of about 2000 m. The other scolopendromorph species are known to occur on lowlands, up to 600 m. Most lithobiomorphs (Eupolybothrus litoralis, Lithobius agilis, Lithobius creticus, Lithobius erythrocephalus, Lithobius lucifugus, Lithobius nigripalpis, and Lithobius viriatus) are common from low to high altitudes on mountains, on subalpine and alpine phrygana (Fig. 15). Apart from these, some species occur at lower altitudes, up to 800 m (Lithobius cretaicus, Lithobius micropodus, Lithobius aeruginosus, Lithobius crassipes, and Lithobius pamukkalensis). Only Lithobius piceus seems to prefer the high altitudes, and was found from 1650 m upwards (Fig. 15). Most geophilomorph species (Fig. 15) are present from lowland to subalpine or alpine areas (Pachymerium f. ferrugineum, Clinopodes flavidus, Geophilus carpophagus, Geophilus naxius, Bothriogaster s. graeca, Bothriogaster s. thesei, and Dignathodon microcephalus). However, Schendyla nemorensis, Geophilus linearis, and Strigamia acuminata are common on Crete from mountainous to alpine areas, up to Fig. 14. Chorotype local distribution of Crete centipede species. CCR: central Crete, CRE: Crete, ECR: east Crete, SCR: south Crete, WCC: west-central Crete, WCR: west Crete, WEC: west east Crete, EN: endemic, EUR: European, MED: Mediterranean, WPA: west Palearctic.

44 410 S. SIMAIAKIS ET AL. Isr. J. Zool. Fig. 15. Centipede altitudinal distribution in Crete m. The last was also found on Gavdos Island. Henia athenarum and Insigniporus sturanyi have been found in mountainous areas at middle altitudes from 700 to 1100 m. Moreover, Haploschendyla europea, Nannophilus ariadnae, Geophilus insculptus, and Geophilus pygmaeus are not very common on Crete and occur at low altitudes, up to 400 m. Furthermore, three species, Henia devia, Henia bicarinata, and Henia pulchella, are rare on Crete, where they are found at low and middle altitudes, up to 1100, 900, and 800 m respectively. Finally, Pachymerium f. insularum is very common on the adjacent satellite islets in lowland areas, but it has been also found on sandy coasts next to the sea, similar to Tuoba poseidonis. As regards temporal distribution in Chilopoda, Lithobiomorpha is known to be active in autumn and winter, Geophilomorpha from autumn to spring, and Scolopendromorpha