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ZOOSYSTEMATICA ROSSICA, 22(1): 129–140 25 JUNE 2013

Grassflies of the family (Diptera) on Mediterranean islands Злаковые мухи семейства Chloropidae (Diptera) на островах Средиземного моря

E.P. NARTSHUK

Э.П. НАРЧУК

E.P. Nartshuk, Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia. E-mail: @zin.ru

One hundred sixteen of Chloropidae (Diptera) are recorded from several Mediter- ranean islands: 66 on Sardinia, 46 on Sicilia, 45 on the Balearic Islands, 39 on the Maltese Islands, 31 on Corsica, 22 on Cyprus and 6 on Crete. The number of species recorded depends more on the level of knowledge of fauna than on the area of the islands. The recorded species belong to no less than ten chorotypes, most of them, to Euro-Mediterranean, Macaronesian- Mediterranean or Mediterranean chorotypes. Two species, aterrima (Duda) and Oscinimorpha tenuirostris (Duda), are known on islands and in North Africa (Tunisia) but not recorded from mainland Europe. Two species are East Mediterranean, meridiana Dely-Draskovits and pori Harkness et Ismay. Scoliophthalmus trapezoides Becker and Anacamptoneurum obliquum Becker, which are recorded only from Cyprus, are distributed also in Africa, Arabia and the Oriental Region. Five species are up-to now found only on islands. This review includes 22 species (16 valid), which have type localities on the islands. На островах Средиземного моря найдено 116 видов злаковых мух (Chloropidae): 66 ви- дов на Сардинии, 46 – на Сицилии, 45 – на Балеарских о-вах, 39 – на Мальтийских о-вах, 31 – на Корсике, 22 – на Кипре и только 6 – на Крите. Число обнаруженных видов опре- деляется степенью изученности фауны, а не площадью острова. По типу распростране- ния найденные виды принадлежат не менее, чем к 10 хоротипам; большинство из них имеют евро-средиземноморский, макаронезийско-средиземноморский или средизем- номорский тип ареала. Два вида – Lasiambia aterrima (Duda) и Oscinimorpha tenuirostris (Duda) – известны с островов и из Северной Африки (Тунис), но не отмечены в мате- риковой части Европы. Два вида – Tricimba meridiana Dely-Draskovits и Trachysiphonella pori Harkness et Ismay – имеют восточносредиземноморский тип ареала. Scoliophthalmus trapezoides Becker и Anacamptoneurum obliquum Becker, которые найдены только на Ки- пре, распространены также в Африке, Аравии и Ориентальной области. Пять видов из- вестны только с островов. Для 22 видов, из которых 16 в настоящее время валидные, на рассматриваемых островах расположены типовые местности. Key words: grassflies, distribution, range types, review, Mediterranean islands, Diptera, Chloropidae Ключевые слова: злаковые мухи, распространение, типы ареалов, обзор, острова Среди- земного моря, Diptera, Chloropidae

© 2013 Zoological Institute, Russian Academy of Scienсes 130 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS

INTRODUCTION ranean basin dated to 5.96 ± 0.02 million years ago in the Upper Miocene (Krijgsman Investigations of island faunas deal with et al., 1999). The first fossil Chloropidae are many general problems of ecology and bio- known from the Eocene Baltic (Hen- geography, especially the composition of in- nig, 1965; Tschirnhaus & Hoffeins, 2009). sular faunas. The biogeographical theory of The fossil Chloropidae from the Miocene MacArthur and Wilson (MacArthur & Wil- are still undescribed, but more or less di- son, 1967) is based mostly on research of is- verse Chloropidae fauna probably had ex- land faunas. Some authors (Gressit, 1955; isted at that time. The Balearic Islands, Cor- Gilbert, 1980; Chernov, 1982) considered sica, Sardinia, Sicily, Malta and Cyprus were local ecological conditions as more impor- connected with the mainland Eurasia dur- tant for of insular faunas. The ing the Middle Pleistocene, at the Calabrian material on Chloropidae of some (mostly and Sicilian stages (Gratsianskiy, 1971). large) Mediterranean islands obtained in All large islands except Cyprus are for- the last decades by the author and other mally assigned to Europe, but Cyprus, to dipterists allows one to evaluate the biodi- Asia. Location of islands in different parts versity of grassflies on these islands. of the Mediterranean Sea, different size of The Chloropidae is a species-rich dipter- islands and their different history may be of an family characterized by high biological interest to analyze their Chloropidae fauna. diversity. Many chloropid species are phy- tophagous with larvae developing in shoots MATERIAL and seeds of grasses and sedges ( and Cyperaceae), many use rotting tissues The paper reviews the published data of plants, fungi, decaying wood, excrements, and the unpublished material identified by frass and eggs of spiders and , nests of the author. The following publications were birds and and even live and dead used in the paper: for the Balearic Islands, vertebrates as substrates for development Ebejer (2006) and Carles-Tolrá & Ventura (Ferrar, 1987; Ismay & Nartshuk, 2000). (2009); for Corsica, Becker et al. (1910) Most species of Chloropidae are rather small and Séguy (1934); for Crete, Nartshuk in size, and they were often found during (1984); for Cyprus, Georghiou (1977) and trapping of air-borne insects (Yoshimoto et Nartshuk (1990, 2010b); for the Maltese Is- al., 1962; Yoshimoto & Gressit, 1963). lands, Ebejer (2010); for Sardinia, Nartshuk There are no special publications that (1995a, 2009b, 2011) and Merz (2005); for review insular faunas of Chloropidae. How- Sicily, Bezzi & De Stefani-Perez (1897) and ever, Chloropidae collected from many is- Nartshuk (1995a). lands of the World Ocean were studied by In addition to these published data, the several researchers, mostly by Sabrosky material was used collected from Sardinia, (1952, 1955, 1957, 1962, 1976, 1977, 1978, Cyprus, Sicilia and Corsica and sent to the 1997), also by Bezzi (1928), Frey (1945, author for identification. The author also 1949, 1958), Frey & Stora (1937), Wheeler identified the material from Mediterranean & Forrest (2002, 2003) and others. Some islands in the following European museums: new species, including endemic ones, were Finnish Museum of Natural History (Hel- described from islands. sinki), Museum of Zoology of the Lund Uni- All Mediterranean islands are situated versity, Museum für Naturkunde (Berlin), close to mainland Europe. They were united Zoologische Staatssammlung München, with the mainland in prehistoric times and Naturhistorisches Museum Wien. some of them connected Europe with North Chloropidae of some small islands of Africa in the past. The Messinian salinity Italy and Greece are practically unknown, crisis and desiccation of the entire Mediter- only few species were recorded from these

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS 131 islands. Oscinimorpha longirostris (Loew, of known species does not correspond with 1858) and bimaculata (Loew, the square of island. The greatest number of 1845) were described from Rhodos, Chlo- species is recorded from Sardinia. This re- rops lucens Becker, 1910, from Paros flects the large size of the island, its diverse (Greece), and a small list was published for habitats and geology and the fact that Sar- Capri (Italy) (Nartshuk, 2005). Therefore dinia has been the subject of special investi- these islands are not discussed. gation (Nardi et al., 2011). The Chloropidae of Crete are the least known. The same is RESULTS AND DISCUSSION also true for the fauna of Greece in compari- son with the faunas of Spain (without the A total of 116 species is recorded on the Canarian Islands) and Italy (see Figure). islands under study. For a few species only Species of the subfamilies Siphonellop- the generic name is known. The list of spe- sinae and Rhodesiellinae, which are not cies with the data on their distributions by numerous in the European fauna, are rep- islands is given in Table 1; some quantita- resented by only one species each. Species tive data are summarized in Table 2 and in of the subfamily predominate the Figure. The islands under consideration over species of the subfamily . are arranged according to the size of the ter- The numbers of species of Siphonellopsinae: ritory in the following order: Sicily, Sardin- Rhodesiellinae: Oscinellinae: Chloropinae ia, Cyprus, Corsica, Crete, the Balearic Is- are in the ratio 1: 1: 71: 43. This ratio agrees lands, and the Maltese Islands. The number with the data on the Chloropidae fauna of

Figure. Number of species of Chloropidae (Diptera) recorded up to now on islands and peninsulas of the Mediterranean [for details, see Tables 1, 2 and text; data on Turkey are taken from Nartshuk (2012a)].

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 132 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS other islands, for example, the islands of The distribution ranges of chloropid spe- Macaronesia (Nartshuk, 1995b) and some cies occurring on the Mediterranean islands others (Nartshuk, 2012b). There may be are diverse. Seven chorological elements two explanations for this phenomenon. were earlier distinguished within the Chlo- First, species of the subfamily Oscinellinae ropidae fauna of Cyprus (Nartshuk, 1990). are generally smaller in size in comparison A similar diversity of range types was ob- with species of the subfamily Chloropinae, served in chloropids occurring on all other and are therefore more likely to be abun- Mediterranean islands. Six species have a dant in aerial plankton where they have multiregional distribution, 9 species have a better opportunities for dispersal to other Holarctic distribution, 9 species are transpa- territories. Second, most of the species of laearctic, 16 ones are Euro-Asian, 42 species Oscinellinae are saprophagous or phyto- have Euro-Mediterranean, Macaronesian- saprophagous, and are therefore less influ- Mediterranean or Mediterranean distribu- enced by the presence of specific host plants tion ranges, and 5 species occur in southern than are species of Chloropinae, which are Europe, Mediterranean area and to the east usually phytophagous and associated with as far as Kazakhstan, Central Asia and Af- more specific host plants. These peculiari- ghanistan. Two species, Tricimba meridiana ties may give to the Oscinellinae a greater and Trachysiphonella pori, are known only chance for colonization of new areas. from the Eastern Mediterranean. Scolioph- The Chloropidae fauna on all the exam- thalmus trapezoides has a mainly Afrotropi- ined islands is relatively rich. The number cal distribution and is known also from Is- of species found on the islands comprises rael. Anacamptoneurum obliquum is known 57.7% of the species number recorded in from Africa, Arabia, India, Turkey and Israel. Italy, 201 species (Nartshuk, 1995a, 2009a, Most of species have a Euro-Mediterranean 2009b; Merz, 2005) and is superior to the or Mediterranean distribution. Scolioph- fauna of Spain except Canary Islands, 97 thalmus trapezoides and Anacamptoneurum (De Bruyn & Baez, 2002; Nartshuk, 2004), obliquum occur only on Cyprus. Each of the and Greece, 51 (Nartshuk, 2010a). The ex- five species, Lasiambia parallela from Crete, amined islands are situated near Europe, laminata from Corsica, Aphanotri- and some of them have been connected with gonum lanceolatum and sardous the mainland in the past. The fauna of Chlo- from Sardinia, and Speccafrons cypria from ropidae of distant oceanic islands is usually Cyprus, are known from only one island. more meager (Nartshuk, 2012b). Some spe- Elachiptera sarda described from Sardinia cies have been described from the islands probably occurs also on the Balearic Islands mentioned in this review, and 16 of these 22 (M.J. Ebejer, pers. comm.). Some of these species are considered valid taxa. However, species eventually may be found on other although their type localities are situated islands or in mainland Spain, Italy, south- on the islands, many of these species are ern France, Greece and Turkey. The faunas now known to have a more extensive distri- of Chloropidae of the Balearic Islands, Cor- bution. Some species occur on all (Thauma- sica, Sardinia, Sicily and the Maltese Islands tomyia notata) or nearly all the islands (Ela- are by their nature mainly Mediterranean; chiptera bimaculata and frit, on six some Afrotropical and Arabo-Oriental ele- islands; E. megaspis, Speccafrons halophila, ments appear only in the fauna of Cyprus. Trachysiphonella ruficeps, Tricimba hume- In general, the Chloropidae on the Mediter- ralis, Assuania thalhammeri, fulvi- ranean islands represent the same pattern frons and pachycera, on of chorological elements as chloropids from five islands). Six of them belong to Oscinel- the Mediterranean mainland areas, except linae and four to Chloropinae. These species for the five species known from one island have a very wide distribution. only (see above).

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS 133 Chorotypes + + Bal Cor Sar Sic Mal Cre Cyp n) 1820 + Transpalaearctic é (Strobl)(Strobl)Nartshuk(Fall 1909Duda 1893 +Collin 1962 +Becker +(Haliday)Nartshuk + 1932(Macquart) 1946 1910 1838 + 2008 1835 + + + + + + + Euro-Asian + + + + Transpalaearctic + South-Euro-Mediterranean Transpalaearctic Euro-Mediterranean Euro-Mediterranean Euro-Mediterranean Holarctic Sardinia StroblBeckerBecker 1906NartshukNartshuk 1903(Becker)Collin 1803Collin 2003Nartshuk + 1964Dely-Draskovits 1903 + 1945 1981 1964 2008 + + + + + + + Mediterranean + + Near East, Afrotropical + + + + Africa, India, Arabia Mediterranean + Mediterranean-Turanian + Macaronesian, Mediterranean Euro-Mediterranean Mediterranean, Afghanistan Euro-Mediterranean Sardinia sp. sp. Subfamilies and species Authors Year minima Calamoncosis stipae Calamoncosis gallarum Conioscinella mimula Dicraeus nigropilosus Dicraeus raptus Dicraeus sardous Dicraeus tibialis Calamoncosis duinensis Conioscinella frontella Siphonellopsinae Siphonellopsis lacteibasis Rhodesiellinae Scoliophthalmus trapezoides Oscinellinae Anacamptoneurum obliquum parahastatum Aphanotrigonum Aphanotrigonum anderssoni Aphanotrigonum bicolor Aphanotrigonum favillaceum Aphanotrigonum femorellum Aphanotrigonum inerme Aphanotrigonum lanceolatum List of Chloropidae species on islands the Mediterranean. 1 2 3 4 5 6 7 8 9 13 14 15 17 18 19 20 21 22 12 16 10 11 Table 1. Table

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 134 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS Chorotypes Mediterranean, Africa, Arabia and Near East + + + Bal Cor Sar Sic Mal Cre Cyp n) 1820 + + Euro-Asian n) 1820 + + + + Transpalaearctic é é (Thalhammer)MeigenLoew 1898Schiner +(Linnaeus) + 1830Beschovski + + 1858 1864 1758 + + + + 1978 + + + + + + + + + + + Euro-Mediterranean to Afghanistan + Euro-Asian + Multiregional Holarctic Euro-Mediterranean (Meigen)(Duda)(Strobl)(Duda) 1830(Duda)(Becker) 1932 1893(Fall +(Becker) 1933 1933 + + 1910 + + 1910 + + + + + Transpalaearctic Euro-Asian Euro-Mediterranean Balearic Islands and Tunis + Mediterranean Euro-Mediterranean + Crete (Loew)(Fall (Loew)DudaNartshuk 1845 + 1858 + 1932 2008 + + + + + + + + + + + + Macaronesian, Mediterranean + Macaronesian, Mediterranean Euro-Mediterranean Sardinia and Balearic Islands sp. sp. 1 sp. 2 Subfamilies and species Authors Year lucens Lipara rufitarsis Lipara similis Oscinella nartshukiana Lasiochaeta pubescens zuercheri Lasiambia albidipennis Lasiambia aterrima Lasiambia brevibucca Lasiambia fycoperda Lasiambia palposa Lasiambia parallela Lasiambia Incertella albipalpis Elachiptera megaspis Elachiptera rufifrons Elachiptera sarda Gaurax Elachiptera bimaculata . Continued. 40 41 42 43 44 39 31 32 33 34 35 36 37 38 29 30 24 25 26 27 28 23 Table 1 Table

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS 135 Chorotypes + + Bal Cor Sar Sic Mal Cre Cyp NartshukHarkness et Ismay(Macquart) 1976 1964 1835 + + + + + + + East-Mediterranean Euro-Mediterranean-Turanian Euro-Mediterranean (Meigen)(Meigen)Nartshuk(Meigen) 1838(Duda) 1830(Duda) 1956 +(Loew) 1830 +(Strobl)(Strobl) 1932 + +(Duda) 1932(Meigen) 1858 + + 1900Seguy + + + 1893(Macquart) +(Meigen) 1933 + + 1830(Loew) + + + + +Nartshuk 1835 1946 + +(Duda) + Holarctic + 1838 Transpalaearctic + + + 1858 Euro-Mediterranean + 1990 + + Transpalaearctic + 1932 + Euro-Asian + + Euro-Mediterranean + Macaronesian, Mediterranean + + + + Euro-Asian + Macaronesian, Mediterranean + + + + Balearic Islands and Tunis + Euro-Asian + + Mediterranean Multiregional Euro-Asian + Subcosmopolitan + Cyprus Euro-Mediterranean (Becker) 1908 + + Macaronesian, Mediterranean costalis nitidigenis sp. aff. sp. aff. Subfamilies and species Authors Year Trachysiphonella pori Trachysiphonella ruficeps Trachysiphonella Trachysiphonella carinifacies Trachysiphonella Oscinella pusilla Oscinella ventricosi Oscinella vindicata Oscinimorpha albisetosa Oscinimorpha arcuata Oscinimorpha longirostris Oscinimorpha minutissima Oscinimorpha novakii Oscinimorpha tenuirostris cognatum Polyodaspis picardi Polyodaspis ruficornis Polyodaspis sulcicollis ornatifrons Speccafrons cypria Speccafrons halophila Speccafrons Oscinella nitidissima Oscinella Oscinella nitidigenis . Continued. 66 67 65 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 47 46 45 Table 1 Table

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 136 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS Chorotypes ++ Bal Cor Sar Sic Mal Cre Cyp n) 1820 + Transpalaearctic n) 1820 + Holarctic é é (Strobl)(Latreille)Loew(Rossi)(Fall 1893 1805(Loew)Meigen +Smirnov 1866 1790 +MeigenMeigen + 1866 +Meigen 1830Meigen + 1967 +Strobl 1830 +Strobl + 1830(Bjerkander) 1830 +Loew 1830Meigen + +Meigen + 1902 + 1778 + +Becker 1893 Euro-Mediterranean + Euro-Mediterranean + + + 1866 1830 1830 + + Mediterranean + + 1912 Mediterranean + + + + Euro-Asian + Euro-Asian + Euro-Mediterranean-Kazakhstanian + + Euro-Asian + + Transpalaearctic European + Euro-Mediterranean-Kasakhstanian Euro-Asian Euro-Asian Euro-Asian European Euro-Asian Euro-Mediterranean Euro-Mediterranean (von Roser)(Meigen)(Loew)(Fall 1840Dely-Draskovits 1830 1983 1858 + + + + + + + + Euro-Asian + Holarctic + Euro-Asian East-Mediterranean sp. Subfamilies and species Authors Year curvipennis Capnoptera pilosa Capnoptera scutata cereris Cetema myopinum Chlorops geminatus Chlorops emiliae Chlorops gracilis Chlorops hypostigma Chlorops interruptus Chlorops laetus Chlorops novakii Chlorops pannonicus Chlorops puncticornis Chlorops scalaris Chlorops serenus Chlorops socius Chloropinae Assuania thalhammeri Tricimba cincta Tricimba humeralis Tricimba lineella Tricimba meridiana Tricimba Tricimba Trachysiphonella scutellata Trachysiphonella . Continued. 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 74 69 70 71 72 73 68 Table 1 Table

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS 137 Chorotypes , Cyprus. Cyp , Crete; Cre ++ , the Maltese Islands; Bal Cor Sar Sic Mal Cre Cyp Mal , Sicily; Sic neville) 1843 + + + Transpalaearctic , Sardinia; é Sar n) 1820 + Holarctic rin-M é é (Zetterstedt)(Meigen)(Duda)(Fall 1848(Strobl)Egger 1830A. Costa + + 1933Meigen(Haliday) + + 1900(Gu 1862 1885 1830 + + 1833 + + + + + + + + + + + + Euro-Asian + Transpalaearctic + Euro-Kazakhstanian Euro-Mediterranean Euro-Mediterranean Euro-Mediterranean Euro-Mediterranean Euro-Mediterranean BeckerDudaMacquartFedoseevaMacquart 1912 1835 1933 1960 1835 + + + + + + + + Euro-Mediterranean European Corsica Euro-Asian Holactic Meigen(Zetterstedt)Strobl(Becker) 1830 1848(Meigen)(Meigen)Macquart + 1902(Becker) 1910 1830 + + 1830 + 1835 + + + 1907 + + + + + + + + + + + European + Holarctic + Mediterranean + Multiregional + Euro-Mediterranean + Holarctic + Transpalaearctic Transpalaearctic , Corsica; Cor sp. , the Balearic Islands; Subfamilies and species Authors Year Bal : flavitarsis Cryptonevra nigritarsis messoria Diplotoxoides dalmatina calva Eurina ducalis Chlorops troglodytes Eurina lurida Eutropha fulvifrons Lasiosina herpini Lasiosina immaculata Lasiosina laminata femorata Meromyza nigriseta Meromyza nigriventris Meromyza Meromyza variegata Pseudopachychaeta approximatonervis Pseudopachychaeta pachycera elongatula Thaumatomyia rufa Thaumatomyia sulcifrons . Continued. 94 95 96 97 98 99 93 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 Table 1 Table Abbreviations

© 2013 Zoological Institute, Russian Academy of Scienсes, Zoosystematica Rossica 22(1): 129–140 138 E.P. NARTSHUK. CHLOROPIDAE ON MEDITERRANEAN ISLANDS

Table 2. Number of species of Chloropidae in different subfamilies on islands and in some countries of the Mediterranean.

Subfamilies Bal Cor Sar Sic Mal Cre Cyp Spain Italy Greece

Siphonellopsinae 1011000320 Rhodesiellinae 0000001110 Oscinellinae 33 14 40 23 30 3 15 39 91 32 Chloropinae 11 17 25 22 9 3 6 54 107 19 Total 45 31 66 46 39 6 22 97 201 51

Note. Spain including Balearic Islands but excepting Canary Islands, Italy including Sardinia and Sicilia, and Greece including Crete. Abbreviations as in Table 1.

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