Ann. Mus. civ. St. nat. Vol. 8 2005 [2007] pp.95-106 ISSN 1127-4476

The Raphidioptera of the Apennines Peninsula: a biogeographical analysis*

Horst Aspock 1 & Ulrike Aspock2

1) Department of Medical Parasitology, Clinical Institute of Hygiene and Medical Microbiology, Medical Uni­ versity of Vienna, Kinderspitalgasse 15, A-1095 Vienna, , e-mail: [email protected]; 2) Department of Entomology, Museum of Natural History, Burgring 7, A-1010 Vienna, Austria, e-mail: [email protected]

So far, 24 species of Raphidioptera - 21 of /3 of Inocelliidae - have been found in , 22 (20/2) of these on the mainland only. Five species (4/1) have been detected only in the northernmost parts (Southern ), 17 species (16/1) have been recorded from various parts of the Apennines Peninsula (AP). By far most of these are monocentric Adriatomediterranean faunal ele­ ments (8/1), 3 species of Raphidiidae are polycentric Adriatomediterranean-Balkanopontomediter­ ranean faunal elements. Four species of Raphidiidae are endemic to the AP. Three of these spe­ cies, each representing a systematically isolated monotypic confined to the south of the AP, are most probably ancient inhabitants of the AP and of Iberian origin: Ita/oraphidia so/ariana, Ca/a­ broraphidia renate and Tjederiraphidia santuzza. The fourth endemic species, Phaeostigma (Pon­ toraphidia) grandii, although more recent, is possibly a remainder of the old fauna of the Apulian platform. Of the remaining monocentric Adriatomediterranean elements only Subilla confinis shows an expansivity which has led to a (mainly postglacial) colonisation of larger parts of Central, west­ ern and eastern . The others have hardly extended their distributions beyond the AP (Ra­ phidia ligurica, Parainocellia bic%r) or they have colonised only small parts of the south of Central Europe, southern , and northern Spain (Phaeostigma ita/ogallica, Xanthostigma a/oysiana, X. corsica). Regarding the polycentric species, two (Ornatoraphidia flavilabris, Venustoraphidia ni­ gricollis) also inhabit many extramediterranean parts of Europe, but these colonisations occurred from the Balkan Peninsula (BP) and not from the AP. The remaining species (, Turcoraphidia amara, Raphidia mediterranea, ratzeburgi) are recent invaders (du­ ring the ) from the east and/or from the north. The Raphidioptera species presently in­ habiting the AP exhibit extensive differences in age and origin reflecting the complex geomor­ phological history of the peninsula.

Keywords - Raphidioptera, Raphidiidae, Inocelliidae, Apennines Peninsula, Italy, biogeography, endemisms.

Introduction nean parts of Europe have been inten­ sively studied throughout the past deca­ The Raphidioptera of the Mediterra- des, in particular, also those of the Apen­ nines Peninsula (AP). Since the latest ac­ counts and biogeographical analyses (H. * Gratefully, cordially, and respectfully dedicated to Professor Maria Matilde Principi on the occasion of Aspock et al., 1991, 2001; [Bernardi] lori her 90th birthday (May 4th, 2005). With her papers et al., 1995), besides new records of seve­ on Raphidioptera published during the 1950s and ral species, two surprising discoveries 1960s she has initiated a new era in Raphidioptera have been made on the mainland of Italy, research. Her studies, descriptions, conclusions, and considerations, and, in particular, also her outstan­ one of them leading to the description of a ding drawings have been of utmost influence for our new species belonging to a new genus further neuropterological work. (Rausch et al., 2004; Fig. 1), the other

95 Proceedings of the IX International Symposium on Neuropterology

Fig. 1 - Calabroraphidia renate Rausch & H. Aspock & U. Aspock, 2004, c\. Italia, , Cosenza, Sila Grande, near Vivaio, 39° 23' 38" N 1160 36' 24" E, 1300 m, 18 June 2005, H. & R. Rausch leg. (Photograph by Hubert Rausch). representing a species and a genus so far highest number of species (about 110), only known from Pontomediterranean re­ however, most of them are restricted to gions (Letardi, 2004), and, in addition, a small areas in mountain ranges of the spectacular new species has been de­ peninsulas of Southern Europe, northwest tected in (Pantaleoni et al., Africa, Anatolia, and a few other parts of 2005). Moreover, research on the palaeo­ the Near East respectively. No snake-fly geography and palaeoclimatology of the species exists with a distribution compri­ Mediterranean has made considerable sing all three south European peninsulas, progress so that the time has come to re­ and very few species occur in the AP as consider the genesis of the Raphidioptera well as the Iberian Peninsula (IP) or the fauna of the AP and to summarise our Balkan Peninsula (BP). knowledge on snake-flies of this part of Snake-flies in general are characteri­ Europe in light of biogeography. sed by an extremely low expansivity. The The Raphidioptera (snake-flies) with majority of species from both families on about 215 valid (and possibly 250 exi­ all continents inhabit small areas, in many sting) species is the smallest and more cases just a few mountain ranges. Several ancestral order of the Holometabola. Du­ species exist which have been recorded ring the Mesozoic snake-flies occurred in from a single mountain only (stationary a significantly greater diversity than today, species). Rarely do species occupy large and they also inhabited tropical regions distribution areas. Examples are, on one (H. Aspock, 1998, 2004; U. Aspock & H. hand, a few Euro-Siberian elements with Aspock, 2003, 2004, 2005a). Nowadays distributions covering large parts of Euro­ they are designated as living fossils, while pe and northern Asia (they are particularly their present distribution is restricted to associated with the belt of the coniferous arboreal parts of the Northern hemi­ forests), and on the other hand, several sphere. Nearctic species occurring along the The western Palaearctic harbours the Rocky Mountains from the north of Mexico

96 H. Aspock & U. Aspock - The Raphidioptera of the Apennines Peninsula: a biogeographical analysis to the south of western Canada. So far it Aspock et at. (1991); additional records in is unknown why so many Raphidioptera Italy: Pantaleoni (1990a, 1990b); Letardi species are markedly stationary although (1991, 1993, 1994, 2004); Pantaleoni et it seems that identical or, at least suitable al. (1994); Hellrigl & Holzel (1996); Letardi ecological conditions exist in adjacent re­ & Pantaleoni (1996); Pantaleoni & Letardi gions. (1998); GOsten (1998a, b, c); Letardi & The Raphidioptera of Italy also contain Maltzeff (2001); Nicoli Aldini & Baviera several extremely stationary species (l. (2001); Rausch et al. (2004); Pantaleoni so/ariana, C. renate, T. santuzza) as well (2005). as a number of species with very low (e.g. Phaeostigma (Phaeostigma) nolala Ph. grandii) or modest expansivity (e.g. (Fabricius, 1781) Parainocellia bicolor), but only one really Widely distributed in extra mediterranean expansive species (Subilla confinis). parts of Europe. Recorded in Italy only in the The biogeographical terminology used in northernmost parts, i.e. in the Southern Alps. this article follows that in the Biogeography Phaeostigma (Phaeostigma) ita/ogallica Glossary in H. Asp6ck et al. (1991). (H. Aspock & U. Aspock, 1976) Known only from the south of Italy and from Annotated list of the Raphidioptera southern France. of Italy Phaeostigma gal/oitalica (H. Aspock & U. Aspock, 1976) At present, the following species have Recorded from the AP, southern France been found in continental parts of Italy; the and Dalmatia. list is made complete by the addition of Phaeostigma (Pontoraphidia) grandii comments on the snake-flies of the Italian (Principi, 1960) islands. For distribution maps see H. Recorded only from the southern half of the

Fig. 2 - Documented distribution of Phaeostigma (Pontoraphidia) grandii, of the subgenus Pontoraphidia, and of the genus Phaeostigma.

97 Proceedings of the IX International Symposium on Neuropter%gy

AP (Fig. 2). Ornatoraphidia flavilabris (Costa, 1855) Dichrostigma flavipes (Stein, 1863) Widely distributed in the BP, in the AP, in Widely distributed in the BP and (mainly) the south of France, and in southern parts of eastern extramediterranean parts of Europe. Central Europe. Recorded in Italy only in the north and south to Xanthostigma xanthostigma (Schummel, 1832) and (Bernardi lori et al., Widely distributed in extra mediterranean 1995). parts of Europe and in palaearctic parts of Asia Tjederiraphidia santuzza as far as Sachalin. The only record for Italy is (H. Aspock & U. Aspock & Rausch, 1980) from Southern Tyrol (Alto Adige). Known only from the Aspromonte (Fig. 3). Xanthostigma corsica (Hagen, 1867) Turcoraphidia amara In scattered populations widely distributed (H. Aspock & U. Aspock, 1964). in the AP, Elba, Giglio, , Sardinia, Cor­ In scattered populations in high altitudes, sica, the south of France and the IP. widely distributed in the BP, the Crimean pen­ Xanthostigma aloysiana (Costa, 1855) insula and western Anatolia. Recently detected In scattered populations widely distributed for Italy in the Majella Nat. Park () (Le­ in the AP, the south of Central Europe and of tardi, 2004) (Fig. 3). France, and the north of Spain. Subilla confinis (Stephens, 1836) Raphidia (Raphidia) ophiopsis Linnaeus, 1758 Widely distributed in extra mediterranean Widely distributed in extra mediterranean parts of Europe. Several records in the sou­ parts of Europe and palaearctic Asia as far as thern half of the AP. Found probably in scat­ Irkutsk. In Italy only in the northernmost parts tered populations throughout the whole pen­ (Southern Alps) (Hellrigl & Holzel, 1996). insula. Letardi (1998) and Pantaleoni (1999) question whether the southern Italian popula­ Raphidia (Raphidia) mediterranea tions might represent a distinct subspecies; so H. Aspock & U. Aspock & Rausch, 1977 far this could not be verified by us. Widely distributed in the BP, the southeast

Fig. 3 - Distribution of the genera Subilla, Turcoraphidia (with the isolated record of Turcoraphidia amara in Italy), and Tjederiraphidia. - In the Apennines Peninsula and in Central Europe Subilla is represented by S. confinis only.

98 H. Aspock & U. Aspock - The Raphidioptera of the Apennines Peninsula: a biogeographical analysis of extramediterranean Europe and western Widely distributed in Central and eastern Anatolia. In Italy in isolated populations in Europe, the BP (north of the gulf of Corinth), (as far north as Gargano), and the AP as far south as the Aspromonte. and . Inocellia crassicornis (Schummel, 1832) Raphidia (Raphidia) ulrikae H. Aspock, 1964 Widely distributed in extramediterranean In scattered populations in various ex­ parts of Europe and the more northern parts of tramediterranean parts of Europe. In Italy only palaearctic Asia eastwards to Sachalin. In Italy in the northernmost parts. restricted to the northernmost parts, known Raphidia (Raphidia) ligurica Albarda, 1891 only from South Tyrol (Alto Adige). In isolated populations in , Parainocellia bicolor (Costa, 1855) Calabria and the south of . Widely distributed in the AP and the south Italoraphidia solariana (Navas, 1928) of France and southwestern parts of Central Restricted to the southern parts of the AP Europe. (Fig. 4). Puncha ratzeburgi (Brauer, 1876) The islands belonging to Italy, even the Widely distributed in Central and eastern large ones (Sardinia, Sicily) harbour a Europe, in isolated populations in the north of surprisingly low number of Raphidioptera the BP and the AP, and southern France (Fig. species. Besides Xanthostigma corsica, 4). which has been recorded on several is­ Calabroraphidia renate lands (see above), only the following two (Rausch & U. Aspock & H. Aspock, 2004) species have been found: The discovery of this species (Fig. 1) in a Subilla principiae small area in the Sila Grande in 2003 (Rausch Pantaleoni & U. Aspock & Cao & H. Aspock, 2005 et al., 2004) was quite a surprise. It was recap­ This species - one of the recent discove­ tured in 2005 (Rausch, pers. comm.) and 2006 ries - has been found only in a Quercus pu­ (unpubl.) in a few localities in the Sila. bescens habitat in the Gennargentu on Sar­ Venustoraphidia nigricollis (Albarda, 1891) dinia (Pantaleoni et al., 2005).

Fig. 4 - Distribution of the genera of group II of Raphidiidae.

99 Proceedings of the IX International Symposium on Neuropterology

Fibla (Fib/a) maclachlani (Albarda, 1891) that in reality it is closer to group II. Recorded only from Sicily, Sardinia and Group II comprises 8 genera with a Corsica. distribution confined to Europe and the northwest of Africa. Five of these, which Present state of classification of have a distribution restricted to western Raphidioptera Europe and the northwest of Africa, form a monophyletic clade within group II (Fig. 4). As a precondition to understanding the The remaining genera assigned to group II chorological facts and biogeographical - Puncha Navas, 1915, Italoraphidia H. conclusions it would seem necessary to Aspock & U. Aspock, 1968, and Cala­ provide a condensed account of the con­ broraphidia Rausch & U. Aspock & H. firmed or suspected relationships of the Aspock, 2004 - differ in several striking genera within the families of the Raphi­ characters and probably constitute the dioptera. The order comprises two extant adelphotaxon to the rest of group II. All families, Raphidiidae and Inocelliidae. three occur in Italy, two of them (lta/ora­ They differ in many morphological and phidia, Calabroraphidia, both monotypic) biological characters, presenting a number are even endemic to the south of the AP of striking autapomorphies (H. Aspock et (Fig. 4). al., 1991, 2001; H. Aspock, 2002). Up to Group III consists of two markedly dif­ the present, the classification of each of ferent genera, Mauroraphidia H. Aspock & the two families has been founded solely U. Aspock & Rausch, 1983 (monotypic on morphological characters; an analysis and endemic to the Atlas mountains) and of relationships based on molecular bio­ Venustoraphidia H. Aspock & U. Aspock, logy is, however, in progress. 1968, with two species, one of them con­ The family Raphidiidae comprises 26 fined to the Peloponnisos, the other widely genera, which have been assigned to 7 distributed in Central Europe, the BP and presumably monophyletic groups (groups the AP. I-IV and groups VI-VIII; the dubious group Groups IV to VIII are not represented V was revealed as part of group VI) (H. in Europe. Aspock et al., 1991, 1998). The relationships between these 7 Group I comprises 12 Raphidiidae ge­ groups of Raphidiidae have not yet been nera with a total of more than 90 species. verified. The current classification is: ««(11 It is the dominant group in Europe, the + VII) + IV) + III) + I) + VI) + VIII, despite Caucasus region, Anatolia, other parts of several weak points; the morphological the eastern Mediterranean, the Near East findings urgently need molecular biological as well as the northern parts of Asia. In examination (which is currently in pro­ Italy this group is represented by the ge­ gress). nera Phaeostigma Navas, 1909, Dichro­ Of particular interest is the genus stigma Navas, 1909, Turcoraphidia H. Aguila (group VII), which is restricted to Aspock & U. Aspock, 1968, Subilla Navas, North America and probably is the sister­ 1916, Ornatoraphidia H. Asp6ck & U. group of group II. If this is true, the rela­ Aspock, 1968, Xanthostigma Navas, 1909, tionship must date back to the Mesozoic. and Raphidia Linnaeus, 1758. The mono­ The Inocelliidae is a small family com­ typic genus Tjederiraphidia H. Aspock & prising 21 described valid species which U. Aspock & Rausch, 1985, endemic to have been assigned to 6 genera Fibla Calabria, was assigned - with some hesi­ Navas, 1915, with four species, one of tation - to group I (H. Asp6ck et aI., 1991). them endemic to the islands of Corsica, In the meantime, we have developed seri­ Sardinia and Sicily. Parain 0 cellia H. ous doubts regarding its systematic posi­ Aspock & U. Aspock, 1968, widely distri­ tion, and we cannot exclude the possibility buted in several southern parts of Europe

100 H. Aspock & U. Aspock - The Raphidioptera of the Apennines Peninsula: a biogeographical analysis

(including the AP and the BP), the eastern from the Atlantic Ocean in the late Mio­ Mediterranean and Eastern Asia; five spe­ cene (about 6 millions years ago) - led cies have been described, one of them largely to a desiccation of the Mediterra­ occurs throughout the whole mainland of nean Sea (lasting about 500.000 years) Italy. /nocellia Schneider, 1843, with 6 and to the formation of numerous land species and widely distributed in large bridges and connections. Many of these - parts of Europe and Asia; one species has particularly those deeply below the old sea been found also in the northernmost parts level - were, however, of only limited of Italy. /ndianoinocellia U. Aspeck & H. value for migration of and plants. Aspeck, Negha Navas, and Sininocellia They were basins with, at least, a partly Yang do not occur in Europe. lagoon-like character, high concentrations The relationships within the family Ino­ of salt (Sabhka facies) and high tempera­ celliidae have not yet been clarified. How­ tures. Nevertheless, the Messinian crisis ever, it seems clear that Fib/a is the sister was of great significance for the faunal taxon to the remaining species, and that exchange within the ancient Mediterra­ Parainocellia and /nocellia are probably nean Sea and the continents. sister groups. Another important period for massive changes of land and water in the Mediter­ Palaeogeography of the Apennines ranean were the glacial periods of the Pleistocene. The sea level fell to about Peninsula 100 - 130 m (for short periods possibly even more) below the present sea level. During the past three decades impres­ During this period, many land bridges sive progress has been made in our un­ emerged, which again facilitated migra­ derstanding of the genesis of the Mediter­ tions of organisms on a large scale. It is ranean basin (Boccaletti et a/., 1990; Regl, important to mention that the Northern 1998; Popov et aI., 2005). had largely dried out resulting The AP is composed of several parts in a faunal exchange between southeast having different provenances and ages. Europe and the AP. Thus, it is easily un­ The largest part can be traced back to derstandable why the Raphidioptera fauna lands which were primarily united with the of the AP is markedly heterogeneous from Iberian land-mass, but which separated in a biogeographical, as well as from a sy­ the Oligocene and drifted eastwards in a stematic point of view. rotating movement. In particular, this area encompasses large parts of the Apenni­ nes and parts of the present Calabria. Provenance of the Raphidioptera Also Corsica, Sardinia, and northern parts of the Apennines Peninsula of Sicily were once eastern parts of the Iberian land-mass which separated during Adriatomediterranean species the Oligocene - about 30 millions years ago - and drifted to the east. Endemic species - Of the 22 species In the east, the Apulian platform exi­ of Raphidioptera found on the mainland of sted since the Mesozoic; it also rotated to Italy, four are endemic and restricted to the northeast during the Miocene and sub­ the AP: Ita/oraphidia so/ariana, Ca/abro­ merged under the Dinarides. Only a small raphidia renate, Tjederiraphidia santuzza, narrow stripe has remained as a part of and Phaeostigma (Pontoraphidia) grandii. present-day Apulia and Calabria. Of these, I. so/ariana, C. renate and T. The famous Messinian salinity crisis - santuzza are representatives of monotypic caused as a result of the closure of the genera with distinctly isolated systematic Mediterranean Sea and its separation positions. Ita/oraphidia and Calabroraphi-

101 Proceedings of the IX International Symposium on Neuropterology dia are probably related to Puncha, ano­ insula - The origin and biogeographical ther monotypic genus which is distributed assignment of these species (Ph. ita/oga/­ throughout large parts of Central and ea­ fica, Ph. gal/oita/ica, S. confinis, O. fla­ stern Europe (Fig. 4). vilabris, X. aloysiana, X. corsica, R. /iguri­ The sister taxon of Puncha + Ita/ora­ ca, V. nigricoJlis, P. bicolor) is very diffe­ phidia + Ca/abroraphidia is probably a rent. The two species, Ph. ita/ogaJlica and group of genera occurring in the western Ph. gal/oita/ica, are related to Phaeo­ Mediterranean (Fig. 4). Thus it is reaso­ stigma (Ph.) notata (Fabricius, 1781), a nable to assume that Ita/oraphidia, as well species with a large distribution in ex­ as Ca/abroraphidia, can be traced back to tramediterranean parts of Europe, more­ the ancient invaders coming from Iberia over, to Phaeostigma (Ph.) promethei H. via the drifted platform which nowadays Aspbck & U. Aspbck & Rausch, 1983, re­ forms part of Calabria. This is comparable stricted to the Caucasus, and to Phaeo­ to the origin of Fibla mac/ach/ani (occur­ stigma (Ph.) euboica (H. Aspbck & U. ring in Corsica, Sardinia and Sicily) and to Aspbck, 1976), endemic to mountains of Subilla principiae (most probably endemic Evia. It is likely that the evolution of these to Sardinia), both of which are apparently species took place in the BP during the ancient Tyrrhenian elements derived from Pleistocene and that Ph. italogal/ica and Iberian ancestors. Ph. gal/oita/ica (or their common ancestor) The origin of Tjederiraphidia is not invaded the AP either from the north or clear. Until now the genus was assigned from the east via the dried Adria. to group I (see above), but this seems in­ Two species, O. flavilabris and V. ni­ creasingly more doubtful, and it is quite gricollis, occur also in large parts of the possible that the genus is also an ancient BP (moreover, in parts of Central Europe element with ancestors of Iberian origin. and in the south of France). Per definition The fourth endemic species, Ph. (P.) they represent polycentric Adriato-Bal­ grandii, belongs to a subgenus (Pon­ kanpontomediterranean elements. Maybe toraphidia) which altogether comprises they also reached the AP in the Pleisto­ four species and has a range of distribu­ cene in a similar way as the Phaeostigma tion covering - besides the south of the species. It is of interest that both of the AP - parts of the southeast of Europe, two genera, Ornatoraphidia H. Aspbck & northern Aegean islands, Anatolia, and U. Aspbck, 1968, and Venustoraphidia H. the Caucasus region (Fig. 2). Thus, Pon­ Aspbck & U. Aspbck, 1968, comprise a toraphidia is of distinctly eastern origin second species, and both occur in very and it cannot be doubted that (the ance­ small mountain areas in Greece. stor of) Ph. (P.) grandii came from the P. bicolor is a typical monocentric east. It is certainly much younger than the Adriatomediterranean faunal element with other three endemic species dealt with a distribution throughout the whole above, but it is the sister taxon to the re­ mainland of Italy, from the south of maining three species of the subgenus Calabria to the Southern Alps, but with and thus certainly is not a recent acquisi­ very few records in adjacent parts of tion to the AP. Ph. grandii (or its immedia­ southern France and Switzerland. P. bi­ te ancestor respectively) must have rea­ color is the sister species to the closely ched the peninsula before the glacial pe­ related P. braueri (Albarda, 1891), which riods, i.e. prior to the Pleistocene. Possibly occurs in the southeast of Europe. Both it is the sole remainder of the Raphi­ are the sister group to P. ressli (H. Aspbck dioptera fauna of the old Apulian platform. & U. Aspbck, 1965), the only Inocelliid species of Anatolia. No doubt, P. bic%r is Adriatomediterranean elements not of eastern origin and represents a com­ restricted to the Apennines Pen- paratively recent acquisition to the Raphi-

102 H. Aspock & U. Aspock ~ The Raphidioptera of the Apennines Peninsula: a biogeographical analysis dioptera fauna of the AP. It is a reasona­ racterised by more or less similar mon­ ble assumption that P. bicolor (or its an­ strous-like genitalia. One of these species cestor) reached the peninsula from the is the recently discovered S. principiae, east in the Pleistocene. possibly endemic to Sardinia, the other The genus Xanthostigma with 5 known four species are restricted to the eastern species, distributed from the north of the Mediterranean region (Rhodes, southern IP to the east of Asia, is most probably an Anatolia). Pantaleoni et al. (2005) have ancient element of the Raphidioptera of hypothesised that S. principiae might be the AP. It is likely that it invaded the pen­ an old Tyrrhenian element of Iberian ori­ insula from the north some time before the gin; there is indeed no plausible bio­ glacial periods, perhaps in the late Plio­ geographical argument which might sup­ cene. Both species occurring in Italy, X. port a close relationship between S. prin­ corsica and X. a/oysiana, exhibit some cipiae and the other four species of this degree of expansion by reaching the (probably not monophyletic) group. Mo­ south of France and the north of the IP. lecular biological studies should help cla­ R. ligurica is a typical monocentric rify the question. Adriatomediterranean faunal element with a somewhat isolated systematic position. The genus Raphidia (with 15 to 18 spe­ Balkanopontomediterranean ele­ cies, depending on the taxonomic status ments with recent immigration of some phena) is distributed throughout history large parts of Central and southeast Europe, Anatolia, the Caucasus region By "recent" we mean invasions to the and in the east as far as the Baikal lake. AP that occurred during the Pleistocene Thus, the most plausible assumption is via the Adria bridge. that R. ligurica (or its ancestor) invaded As discussed above, O. flavilabris and the AP from the north or from the east be­ V. nigricollis are of Balkan origin. Three fore the Pleistocene, possibly during the additional species of Balkan origin in­ late Pliocene. vaded the AP recently, i.e. during the gla­ One species remains to be discussed cial period: Dichrostigma flavipes, Raphi­ in this chapter: Subilla confinis. This spe­ dia mediterranea, and Turcoraphidia cies is an Adriatomediterranean faunal amara. element showing a considerable expansi­ D. flavipes occurs in Italy only in the vity. It has been found in various parts of northern parts (the southernmost records the AP and is widely distributed in Central are in Tuscany and Marche), but the spe­ Europe extending to large parts of France, cies is widely distributed in the BP north of England and Romania. In the IP on one Middle Greece (Sterea Elias). Other spe­ hand and in the BP and Anatolia on the cies of the genus occur in eastern Europe, other hand, altogether four other closely Anatolia, and the Near East. related species of the genus occur. It is R. mediterranea, although a species suggested that all these 5 species evolved with a broad ecological spectrum, has in the Pleistocene due to a splitting of a been found only in a relatively small part once continuous distribution area of the of Italy, mainly in the southeast, but re­ hypothetic stem species. The AP was cently also in Lazio. Our previous assump­ colonised by Subilla from the north or from tion that it was introduced to Italy by hu­ the east, possibly in the late Pliocene or man activities in historic or prehistoric early Pleistocene. It is of great interest times (H. Aspock et al., 1991; 2001) that, besides the above mentioned 5 spe­ seems unlikely now as pointed out by cies (confinis-group), the genus Subilla Pantaleoni (2005). Presently, we believe comprises 5 more species which are cha- that it has reached the AP via the dried

103 Proceedings of the IX International Symposium on Neuropterology out Adriatic Sea during (later periods of) far as Emilia Romagna. the Pleistocene. Finally, it cannot be excluded that D. T. amara is another impressive exam­ fJavipes also immigrated (or partially so) ple of a recent addition to the Raphidiop­ into Italy from the north during postglacial tera of the AP from the BP via the Adriatic periods. land bridge. Turcoraphidia is a genus with a distinct Pontomediterranean distribution Significance of the Apennines (Fig. 3). Five species are currently known. T. amara has recently been found in the Peninsula for the Raphidioptera Majella National Park (Abruzzo) (Letardi, fauna of other parts of Europe 2004). Possibly also Puncha ratzeburgi rea­ Compared to the BP, the AP has con­ ched the AP in the Pleistocene from the tributed only sparingly to the Raphidiop­ BP via the Adriatic land bridge, although it tera fauna of other (particularly northern) seems more likely that the species was a parts of Europe. Most Adriatomediterra­ later immigrant (see below). nean species are either endemic or are characterised by an extremely low expan­ sivity. Only a single species - Subilla con­ Extramediterranean (European finis - has extended its distribution in the or Eurosiberian) and/or Balkano­ postglacial period and reached the nor­ pontomediterranean faunal ele­ thern borders of Central Europe (records ments with postglacial immigra­ in Denmark); moreover, its present distri­ tion bution covers large parts of France, Eng­ land and eastern Europe. The distribution of few species in Italy Few species (Ph. ita/ogallica, X. aloy­ is confined to the northernmost parts. Five siana, X. corsica) have colonised parts of of them - Ph. notata, X. xanthostigma, R. southern France and the north of the IP. It ophiopsis, R. ulrikae, and I. crassicomis - is unlikely that all these events should have only been found in Italy in the Sou­ have occurred after the last glacial period. thern Alps and may have persisted there X. corsica e.g. may have reached the IP throughout the later periods of the Pleisto­ already in earlier periods in the Pleisto­ cene, at least throughout the last glacial cene. period. Three of these (X. xanthostigma, In the Raphidioptera fauna of Central R. ophiopsis, I. crassicomis) are Eurosibe­ Europe, the Adriatomediterranean faunal rian faunal elements with high expansivi­ elements play - with the exception of ties and large distribution areas ranging Subilla confinis - a rather insignificant throughout northern Asia and northern, role. There are a few records of X. aloy­ eastern and Central Europe. They may siana, R. /igurica and P. bicolor in the have evolved in Europe or in northern southernmost parts. The occurrence of O. Asia in the Pliocene. flavilabris as well as V. nigricollis can most Puncha, however, is a very old Euro­ probably be traced back to postglacial pean monotypic genus and systematically immigrations from the BP rather than from isolated. It is nonetheless probably related the AP. O. flavilabris occurs in several to Italoraphidia and Ca/abroraphidia and is parts of eastern Austria, but has not yet assigned to group II (see above). P. been found in the southwest of Central ratzeburgi is an Extramediterranean Euro­ Europe. X. a/oysiana, R. /igurica, P. bi­ pean faunal element which persisted color, and O. flavilabris reach the northern south of the Alps during the last glacial pe­ borders of their ranges in the south of riod and possibly enlarged its distributional Central Europe, where they serve as bio­ area in the AP somewhat to the south as indicators for climatically favoured and

104 H. Aspock & U. Aspock - The Raphidioptera of the Apennines Peninsula: a biogeographical analysis particularly rich, but vulnerable biocoe­ demiae Scientiarum Hungaricae, 48 (Suppl. 2): 35-50. noses which merit protection (U. Aspock & Aspock H., 2004 - Kamelhalsfliegen -Iebende Fossilien: Eine der Endkreide-Katastrophe entkommene Tier­ H. Aspock, 2005b). gruppe. Verhandlungen Westdeutscher Entomologen­ tag, Dusseldorf 2002, 1-6. Aspock H., Aspock U. & Rausch H., 1991 - Die Outlook Raphidiopteren der Erde. Eine monographische Darstellung der Systematik, Taxonomie, Biologie, The Raphidioptera fauna of Italy can Okologie und Chorologie der rezenten Raphi­ be regarded as largely comprehended, diopteren der Erde, mit einer zusammenfassenden Obersicht der fossi/en Raphidiopteren (Insecta: although a few additional species so far Neuropteroidea). Goecke & Evers, Krefeld, 730 recorded from regions outside the AP will pp.; 550 pp. certainly be found. It is, however, unlikely Aspock H., U. Aspock U. & Rausch H., 1998 - Was - although it cannot be excluded - that ist Usbekoraphidia turkestanica (H. Aspock & U. Aspock & Martynova 1968)? Zur Kenntnis der new species might be detected in isolated Taxonomie, Okologie und Chorologie mittelasia­ mountainous regions in higher altitudes. tischer Raphidiiden (Insecta: Raphidioptera: Raphi­ Forthcoming biogeographical studies diidae). Stapfia 55: 421-457. will particularly be based upon the results Aspock H., Holzel H. & Aspock U., 2001 - Kommentierter Katalog der Neuropterida (Insecta: of molecular biological investigations. This Raphidioptera, Megaloptera, Neuroptera) der concerns the clarification of the relation­ Westpalaarktis. Denisia 02, Biologiezentrum des ships of the genera (and subgenera) on 00. Landesmuseums, Linz, 612 pp. one hand and the provenance of certain Aspock U. & Aspock H., 2003 - Raphidioptera (). In: V.H. Resh & R.T. Carde (eds.) - populations on the other hand. Encyclopedia of . Academic Press, Amster­ Phylogeography will become an impor­ dam etc., 973-975. tant tool also to clarify open questions on Aspock U. & Aspock H., 2004 - Two significant new structure and biogeographical history of snakeflies from Baltic amber, with discussion on autapomorphies of the order and its included taxa the snake-flies of Italy. (Raphidioptera). Systematic Entomology, 29: 11-19. Aspbck U. & Asp6ck H., 2005a - 28. Ordnung Raphidioptera, Kamelhalsfliegen. In: H. H. Dathe (Hrsg.) - 5. Tei/: Insecta. Acknowledgements In: Lehrbuch der Speziellen Zoologie. BegrOndet von Alfred Kaestner. ZweiteAuffage. KorrigierterNeudruck. H.-E. Gru­ We wish to express our grateful thanks to ner (Hrsg.): Band I: Wirbellose Tiere. Spektrum Aka­ Dr. Fred Ragl and Univ. Doz. Dr. Mathias demischer Ver1ag Heidelberg, Ber1in, 542-552, 888-889. Harzhauser (Geological and Palaeontological Aspock U. & Aspock H., 2005b - Verbreitungs­ Department of the Natural History Museum grenzen von Neuropterida in Mitteleuropa. Linzer bio/ogische Beitrage (Verhan-dlungen des XVIII. Vienna) for valuable discussions on the pa­ Internationalen Symposiums uber Entomo-faunistik laeogeography of the Mediterranean; to Hubert Mitteleuropas [SIEECJ), 37/1: 29-38. Rausch (Scheibbs) for the photograph of [Bernardi] lori A., Kathirithamby J., Letardi A., Ca/abroraphidia renate; to Mag. Franziska An­ Pantaleoni R.A. & Principi M.M., 1995 - Neuro­ derle (Vienna) for producing the maps and for pteroidea (Megaloptera, Raphidioptera, Planipen­ technical assistance with the manuscript; and nia), Mecoptera, Siphonaptera, Strepsiptera. In: to Dr. John Plant (Vienna) for linguistic im­ Minelli A., Ruffo D. & La Posta S. (eds.) - Checklist provement. Finally, the reviewers - Profs. delle specie della fauna italiana, Calderini, Bolo­ gna, 62: 1-20. Achille Casale, Roberto A. Pantaleoni, and Boccaletti M., Ciaranfi N., Cosentino D., Deiana G., Sandro Ruffo - are gratefully acknowledged for Gelati R., Lentini F., Massari F., Moratti G., Pesca­ their valuable comments. tore T., Ricci Lucchi F. & Tortorici L., 1990 - Palin­ spastic restoration and palaeogeographic recon­ struction of the peri-Tyrrhenian area during the Bi bl iography Neogene. Palaeography, Palaeoclimatology, Pa­ laeo-ecology, 77: 41-50. Aspock H., 1998 - Distribution and biogeography of Glisten R, 1998a - Segnalazioni faunistiche italiane. the order Raphidioptera: updated facts and a new 331 - Raphidia mediterranea H. Aspock, U. Aspock & hypothesis. Acta Zoologica Fennica, 209: 33-44. Rausch, 1977 (Raphidioptera Raphidiidae). Bol/ettino Aspock, H., 2002 - The biology of Raphidioptera: A del/a Societa Entomo/ogica /tafiana, 130: 80-81. review of present knowledge. Acta Zoologica Aca- Glisten R., 1998b - Segnalazioni faunistiche italiane.

105 Proceedings of the IX International Symposium on Neuropter%gy

332 - Raphidia figurica Albarda, 1891 (Raphi­ siciliano S. IV, 25 (3-4): 345-376. dioptera Raphidiidae). Bollettino della Sociefa En­ Pantaleoni RA, 1990a -I Neurotteri (Neuropteroidea) tomologica Italiana, 130: 81. della Valle del Bidente-Ronco (Appennino Roma­ Glisten R, 1998c - Segnalazioni faunistiche italiane. gnolo). Bollettino dell'lstituto di Entom%gia "Guido 333 - Tjederiraphidia santuzza (H. Aspack, U. Grandi" dell'Universila di Bologna, 44: 89-142. Asp6ck & Rausch, 1980) (Raphidioptera Raphi­ Pantaleoni RA., 1990b - Un nuovo ausiliario nel diidae). Bol/ettino della Societa Entomologica Ita­ vigneto: Parainocellia bic%r (Costa). Informatore liana, 130: 81. Fitopatologico, 4: 39-43. Hellrigl K. & Hblzel H., 1996 - Neuropteroidea - Pantaleoni RA, 1999 - Neuropterida Described by Netzflligler. In: Hellrigl K. - Die TielWelt SOdtiro/s. A Costa with Type Designation. Deutsche Ento­ Band 1 der Veroffentlichungen des Naturmuseums m%gische Zeitschrift, Berlin, 46: 249-261. SOdtiro/, Bozen, 515-521. Pantaleoni RA., 2005 - Interpretation of Achille Letardi A, 1991 - Neuropteroidea: Raphidioptera, Costa's data on Neuropterida. Bulletin of /nsecto­ Inocelliidae; Planipennia, Chrysopidae. Bollettino logy, 58 (1): 71-92. dell'Associazione Romana di Entom%gia, 45: 132. Pantaleoni RA., Aspbck U., Cao G.V. & Aspbck H., Letardi A., 1993 - Nota sui reperti di Neurotteri dei 2005 - Subilla principiae n. sp., a new spectacular Monti Lepini (Lazio) (Neuropteroidea). Quaderni from Sardinia (Raphidioptera Raphidii­ del Museo di Storia Naturale di Patrica, 4: 93-99. dae). Redia, 87/2004: 3-6. Letardi A, 1994 - Dati sulla distribuzione italiana di Pantaleoni RA, Campadelli G. & Crudele G., 1994- Megaloptera Sialidae, Raphidioptera Inocelliidae e Nuovi dati sui Neurotteri dell'alto Appennino roma­ Planipennia Mantispidae, con particolare riferi­ gnolo. Bofleffino dell'lstituto di Entomologia "Guido mento all'ltalia centrale. Bollettino della Societa Grandi" dell'Universifa di Bologna, 48: 171-183. Enfomologica Italian a , 125 (3): 199-210. Pantaleoni RA & Letardi A, 1998 - I Neuropterida Letardi A., 1998 - Present knowledge of Italian della collezione dell'lstituto di Entomologia "Guido Neuropterida: history, distribution and current pro­ Grandi" di Bologna. Bollettino dell'lstituto di Ento­ blems. Acta Zoologica Fennica, 209: 145-149. mologia "Guido Grandi" dell'Universita di Bologna, Letardi A, 2004 - Turcoraphidia amara (Aspbck & 52: 15-45. Aspbck, 1964), genere e specie nuovi per l'Italia. Popov S.v., Ragl F., Rozanov AY., Steininger F.F., Bollettino dell'Associazione Romana di Entomolo­ Shcherba I.G. & Kovac M. (eds.), 2005 - Lithological­ gia, 59 (1-4): 99-102. Paleaogeographic maps of Paratethys. 10 maps Late Letardi A. & Maltzeff P., 2001 - Neurotteridi e Eocene to Pliocene. CFS Courier Forschungsinstitut Mecotteri della Tenuta Presidenziale di Castel­ Senckenberg, p. 1-46 + 10 maps + 1 table. porziano e delle aree limitrofe. Bollettino dell'Asso­ Rausch H., Aspack H. & Aspack U., 2004 - Calabro­ ciazione Romana di Entomologia, 56 (1-4): 49-62. raphidia renate n. gen., n. sp. - eine neue Spezies Letardi A & Pantaleoni RA., 1996 - I Neurotteroidei und ein neues Genus der Familie Raphidiidae aus W-Paleartici della collezione del Museo di Zoologia Sliditalien (Neuropterida, Raphidioptera). Entomolo­ dell'Universita di Roma (Neuropteroidea). Frag­ gische Nachrichten und Berichte, 48(3/4): 159-165. menta entomologica, 28 (2): 277-305. Ragl F., 1998 - Palaeogeographic Considerations Nicoli Aldini R & Baviera C., 2001 - Nuove for Mediterranean and Paratethys Seaways (Oli­ acquisizioni sulla NeuroUerofauna di Sicilia e isole gocene to Miocene). Annalen des Naturhistori­ circumsiciliane (Insecta Neuropterida). Natura/ista schen Museums Wien, 99A: 279-310.

106 Bibliography of the Neuropterida

Bibliography of the Neuropterida Reference number (r#): 11958

Reference Citation: Aspöck, H.; Aspöck, U. 2005 [2007.??.??]. The Raphidioptera of the Apennines Peninsula: a biogeographical analysis. in Pantaleoni, R. A.; Letardi, A.; Corazza, C. (eds.). Proceedings of the Ninth International Symposium on Neuropterology (20-23 June 2005, Ferrara, Italy). Annali del Museo Civico di Storia Naturale di Ferrara 8:95-106.

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File: File produced for the Bibliography of the Neuropterida (BotN) component of the Lacewing Digital Library (LDL) Project, 2012.