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The family (Neuropterida: Raphidioptera): A review of present knowledge

Horst Aspöck1, Xingyue Liu2, Ulrike Aspöck3 1 Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University of Vienna 2 Department of Entomology, China Agricultural University, Beijing 3 Natural History Museum Vienna; Department of Evolutionary Biology, University of Vienna

Abstract: Die Familie Inocelliidae (Neuropterida: Raphidioptera): Eine Übersicht des gegen- wärtigen Forschungsstandes Die Inocelliidae, eine der beiden Familien der reliktären Insektenordnung Raphidioptera (Kamel­ halsfliegen), erfuhren zuletzt vor 20 Jahren eine zusammenfassende Darstellung. Seither hat sich die Zahl der beschriebenen und als valide anerkannten Arten von 20 auf 31 erhöht, was auf die Entdeckung neuer Arten im südlichen Nordamerika, vor allem aber in Ost- und Südost-Asien und besonders in China zurückgeht. Damit ergaben sich zugleich neue Informationen über die Biologie, Ökologie, Chorologie und Biogeographie dieser durchwegs seltenen Insekten. Bisher kennt man die Larven von 12 Spezies, sie sind durchwegs kortikol. Die Larvalperiode umfasst mindestens zwei, zumeist mehrere (bis sieben) Jahre. Das Paarungsverhalten und die Art der Kopulation wurden bei mehreren Arten studiert; es bestehen markante Unterschiede gegenüber den Raphidiiden. Fossile Inocelliiden sind aus dem Eozän, Oligozän und Miozän bekannt, vermutlich hat die Familie – als einer der an winterliche Kälte und damit an gemäßigte Zonen adaptierten Zweige – bereits vor dem K/T-Impakt (der zu einer Beinahe-Auslöschung der Raphidiopteren führte) existiert. Die Verbreitung der Inocelliidae umfasst bestimmte arboreale Teile der Paläarktis und der Ne- arktis. In der Nearktis beschränkt sie sich auf südwestliche und südliche Teile Nordamerikas. Sie ist insgesamt ähnlich jener der , jedoch etwas kleiner. Es ist bemerkenswert, dass Inocelliiden in einem Großteil der Gebirge Zentralasiens (wo zahlreiche Raphidiiden- Spezies vorkommen) fehlen. Hingegen stellen sie sowohl in Amerika als auch in Asien die südlichsten Vorkommen der Raphidiopteren dar. Die Suche nach Inocelliiden in großen Höhen in noch weiter südlich gelegenen Gebirgen erscheint erfolgversprechend. Zukünftige Arbeiten werden besonders auch die Klärung der phylogenetischen Beziehungen der sieben Gattungen auf molekularbiologischer­ Basis zum Ziel haben.

Keywords: Raphidioptera, Inocelliidae, review, systematics, species list, phylogeny, biology, chorology, biogeography.

H. Aspöck, Institute of Specific Prophylaxis and Tropical Medicine, Medical Parasitology, Medical University of Vienna, Kinderspitalgasse 15, 1095 Vienna, Austria; E-Mail: [email protected] X. Y. Liu, Department of Entomology, China Agricultural University, Beijing 100193, China, E-Mail: [email protected] U. Aspöck, Natural History Museum Vienna, Burgring 7, 1010 Vienna, Austria; Department of Evolutionary Biology, University of Vienna, Althanstraße 14, 1090 Vienna, Austria, E-Mail: [email protected]

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Introductory remarks The Inocelliidae (Figs. 1, 2, 3, 4) is one of the two families of the endopterygotan order Raphidioptera (=). Raphidioptera is one of the smallest insect orders comprising altogether about 230 valid and clearly characterized species, 31 of which belong to the family Inocelliidae. In the Mesozoic, Raphidioptera experienced a period of flowering; they occurred in several families with many species and extended into tropical zones and the Southern Hemisphere. It is assumed that Raphidioptera almost became extinct during the period after the K/T-impact (H. As p ö c k 1998). Since the beginning of the Cenozoic, however, only two families are known to exist, Raphidiidae and Inocelliidae, both of which are adapted to a cold climate. Possibly, the ability to persist throughout cold seasons was a prerequisite for the survival of these lines of snakeflies after theK/T -impact (H. As p ö c k & U. As p ö c k 2009). The latest comprehensive review on the Inocelliidae was published in our monograph 20 years ago (H. As p ö c k & al. 1991). During the past two decades a number of publications devoted to Inocelliidae has appeared with a considerable amount of new information on , biology, chorology and biogeography (see references). Thus, a review of the present state of knowledge seems useful. History of research Li n n a e u s (1758) described the first , a Raphidiid, and it was not until 1781 when an additional valid species of this family was described. In 1832 the first representative of the present-day Inocelliidae was described from Silesia (at that time a province in the east of Germany) as crassicornis by Sc h u mm e l (1832) and shortly thereafter transferred to a new subgenus, Inocellia, by Sc h n e i d e r (1843). Subsequently, Inocellia was raised to a , later to a tribe (Inocelliini) and finally to a family (for details see H. As p ö c k & al. 1991). Fig. 5 shows the progress in the discovery of Inocelliidae over the past 180 years. In the course of the past century, five more valid genera (and two subgenera) were described with altogether 31 valid species (Tab 1): Na v á s , 1915 (including the subgenus Reisserella H.A. & U.A., 1971), Na v á s , 1916, Parainocellia H.A. & U.A., 1968 (including the subgenus Armurinocellia H.A. & U.A., 1973, which was later elevated to the rank of a genus), Indianoinocellia U.A. & H.A., 1970, and Sininocellia Ya n g , 1985. In a recent paper (U. As p ö c k & al. 2011), four more species will be described from Thailand, thus uncovering a new, unknown and probably very important evolution center of the family in Southeast Asia. Taxonomy All known species are morphologically well investigated and characterized. The differentiation of the genera is based, at least in part, on eidonomic characters (mainly head, antennae, wings and wing venation, size), but for the identification of species, characters of the genital structures, preferably of the male genitalia, are essential. Of the 31 described species (plus one subscpecies), 26 are known in both sexes, 4 species are known in the male sex only and 2 species only in the female sex (Tab. 1). The wings are translucent as in all Raphidiidae. Recently, however, a spectacular species, Inocellia elegans (Fig. 6), which is isolated in the genus, was described with large, dark patches on fore and hind wings (Li u & al. 2009b). Keys are available for the identification of all species (H. As p ö c k & al. 1991, U. As p ö c k & H. As p ö c k 1999b, Li u & al. 2009a, 2010a, b). Currently, only few genomic data for a small number of species are available (Ha r i n g & al. 2011). Biology and Ecology As far as we know, larvae of all species of Inocelliidae develop under bark of trees; one very unusual exception is Parainocellia bicolor which also develops on old grapevines, where it may act as a predator of pest (Pa n t a l e o n i 1990, 2007). At present, the larvae of only 12 species (plus one subspecies) are known, but all adults have been found on or around trees so that the conclusion is justifiable that development takes place exclusively under bark. In the northern parts of their distribution, Inocelliidae may be found from sea level to almost the timberline. In the southern parts, they are apparently confined to higher altitudes, usually above 1,000 m, e.g. up to 1,700 m (Mexico) and up to 2,600 m (India).

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Fig. 1: Inocellia crassicornis Sc h u m ., male (Austria, Eichkogel near Vienna). Length of forewing: 9.5 mm. (Orig., F. Anderle phot.)

Fig. 2: Fibla (Reisserella) pasiphae H.A. & U.A., male (Crete, Omalos). Length of forewing: 15 mm. (Orig., H. Aspöck phot.)

Fig. 3: Fibla (Reisserella) pasiphae H.A. & U.A., larva (Crete, Omalos). Length of larva: 23 mm. (Orig., F. Anderle phot.).

Fig. 4: Fibla (Reisserella) pasiphae H.A. & U.A., pupa (Crete, Omalos). Length of pupa: 25 mm. (Orig., F. Anderle phot.)

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The larval period lasts at least two years, but in many species it is apparently much longer, up to four, five, six or (at least in captivity) seven years, and possibly longer. The number of larval stages is around ten, but it is not fixed and may (particularly in case of long development) reach 15 instars. Hard data are, however, not available. Larvae of Inocelliidae are entomophagous, feeding on any small, soft- bodied (e.g. aphids, caterpillars, larvae of beetles and small maggots). According to our information, all snakeflies (Inocelliidae as well as Raphidiidae) need a cold snap in winter; otherwise they will not develop adequately and will not pupate. It is, however, not known how long this period must last and at what temperature. In the species of the Old World, pupation takes place in spring and adults appear (after two or three weeks of a pupal stage) in May or June. In the species of the New World, in particular those of the genus Indianoinocellia, pupation takes place at nearly any time of the year since adults have been found from February to September. Adults of Inocelliidae are (in contrast to Raphidiidae) not predators; when dissecting adult specimens, pollen is sometimes found in the gut; but usually the gut is entirely empty so that one may conclude that they take up no food at all. Lifespan of adults lasts only several days and in males is shorter than in females. The mating behavior of Inocelliidae differs considerably from that in Raphidiidae (U. As p ö c k & al. 1995, U. As p ö c k & H. As p ö c k 1999a; H. As p ö c k 2002, Pa n t a l e o n i 2007). The male crawls beneath the abdomen of the female, then attaches his head by eversible sacs emerging from the basis of the antennae to the 5th sternite of the female and finally bends his abdomen so that the 9th gonocoxites of the male can grasp the abdomen of the female. This mating behavior has been observed in phylogenetically distant genera of Inocelliidae; most probably it is characteristic for the whole family thus representing a synapomorphy of the Inocelliidae.

Phylogeny and Inocelliidae is ipso facto the sister taxon of Raphidiidae, the only other family of the order. There is no indication that an extinct earlier sister taxon of the Inocelliidae ever existed. The oldest fossils which can be clearly attributed to Inocelliidae are from the Eocene and Oligocene. These were found in Colorado (Florissant) and Europe (Baltic Amber), respectively. These species were attributed to the genus Fibla Na v á s on one hand (En g e l 1995, 2002) and to a new genus, Succinofibla U.A. & H.A., on the other hand (U. As p ö c k & H. As p ö c k 2004). Moreover, a new genus, Miofibla Ne l , was described based on a new species from the Miocene of France (Ne l 1993)1. All these fossils were found within the distribution range of present-day Inocelliidae. Although they represent remarkable findings, they have not yet contributed to an understanding of the generic structure of the family. There are no comprehensive modern studies on the phylogenetic structures within the family. Twenty years ago, a hand-made dendrogram was published (H. As p ö c k & al. 1991) with the following relationships: (Inocellia + Parainocellia) + [Indianoinocellia + (Negha + Fibla)]. In a molecular study on the phylogeny of the Raphidiidae (Ha r i n g & al. 2011), representatives of 4 species from 4 genera of Inocelliidae were included, in which the following relationships emerged: [(Inocellia + Parainocellia) + Fibla] + Negha. A comprehensive molecular study, which should include distinctly more species of Inocelliidae, is still pending.

Distribution and biogeography Fig. 7 shows the world distribution of the Inocelliidae. The records of new species described elsewhere (U. As p ö c k & al. 2011) do not significantly alter the pattern. The world distribution of the Inocelliidae largely coincides with that of the Raphidiidae (for comparison: Fig. 8, world distribution of Raphidioptera). The following differences merit attention:

1 A from the Oligocene of British Columbia (Canada), described as Archiinocellia oligoneura Ha n d l i r s c h , is probably not an Inocelliid (En g e l 2009). Another fossil found in Xiaoershijiazi (China) was described by Wa n g (1987) as Sinoinocellia liaoxiensis and assigned to the family Inocelliidae. In our opinion, however, this fossil certainly does not represent an Inocelliid species.

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Fig. 5: The progressive discovery of species of Inocelliidae world-wide (1832–2011).

Fig. 6: Inocellia elegans Li u , H.A., Ya n g & U.A. from Li u & al. 2009b. Scale bar: 1.0mm

Inocelliidae occur somewhat more south than Raphidiidae in the New World as well as in the Old World. In America the southernmost findings of Inocelliidae are at the Mexican/ Guatemalan border (U. As p ö c k & al. 1992). (Raphidiidae have been found in the State of Oaxaca, but not further south.) In Asia, the southernmost findings of Raphidiidae are in Taiwan, those of Inocelliidae in Thailand. Like Raphidiidae, Inocelliidae do not occur in northern or eastern North America, and their whole distribution range is distinctly smaller. The northernmost records are around 50°N (in the Old World around 66°N and of Raphidiidae in the New World around 53°N) and the easternmost records in America north of Mexico are around 111°W (of Raphidiidae around 98°W).

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Surprisingly there is a considerably large area in Central Asia (Southern Kazakhstan, Kyrgyzstan, Uzbekistan, Tajikistan; possibly also in the north of Iran) where many species of Raphidiidae occur, while no Inocelliid species has yet been found despite intensive field studies by several specialists throughout several years. Table 1 gives a brief overview of the distribution of the 31 species of Inocelliidae. Most species show a very low degree of expansivity and are thus more or less confined to their Pleistocene refugial centres. In the Old World, one species (I. crassicornis) has a large distribution ranging from the Far East throughout Northern Asia to Central Europe. In America, three species (N. inflata, N. longicornis, N. meridionalis) show a considerable degree of expansivity, which results in larger distribution areas in western North America. Seven species are Mediterranean faunal elements. In Asia, distribution centers are situated in the north of India and, in particular, in the east and southeast of Asia. During the past years, 9 new species of Inocelliidae were described from mainland China (Li u & al. 2009a, b, 2010a, b) and Taiwan (U. As p ö c k & al. 2009). In sum, southeast Asia represents a particularly important distributional centre of the family (U. As p ö c k & al. 2011)

Fig. 7: World distribution of the family Inocelliidae.

Fig. 8: World distribution of the order Raphidioptera.

570 Ha l l e (Sa a l e ) 2012 Mi t t . Dt s c h . Ge s . a l l g . a n g e w . En t . 18 X X X X X North America America X SE - Asia X Japan X X X X X X X X X X X X X X X China Taiwan S - Korea N - Korea Asia X X X India Bhutan X X Near East X X Abbreviations: M, male; F, female; L, larva Abbreviations: M, male; F, Asia North. X North Africa Africa X X X X X X Europe Mediterr. Europe X X X Extra - Europe mediterr. M, F, L M, F, Known: M, F, L M, F, M, F, L M, F, F M, F M M, F, L M, F, M, F M, F, L M, F, M, F, L M, F, M, F M, F M, F, L M, F, M, F M, F L M, F, M, F, L M, F, M, F, L M, F, M L M, F, L M, F, M, F F M, F M, F M, F M, F M, F, L M, F, L M, F, M M, F M, F M , a u s c h , 1992 , 1991 a u s c h & U . A , 2010 g , 1958) & U.A., 2010 & U.A., 2010 a n U.A. & H.A., 1968 g g a u s c h a n a n & U . A ., 2009 & U . A , 2010 , 1832) , 1891) g g , 1891) & U . A ., 2010 a e l a g , 1919) d a n a n a r p e n t e r d & H . A ., 2009 , 1855) , 1891) ., Y , H . A ., ( H . A & U ., 1971) , 1917 mm a n i u a , 1985 , H . A & U ., 2009 , 1915 ( C i u d g a v á s l b a r i u ., Y , H . A ., ., Y , H . A ., L l b a r U.A., H.A. & R o t o o s t a , H . A & U ., 2009 c h u , 1936 L a n (U.A. & H.A., 1968) i u i u m ( A i u , 1999 a v á s ( C Y ( A l b a r ( S , 1891) g L ., Y , H . A ., ., Y , H . A ., (H.A. & U.A., 1973) k a L a v á s U.A., 1988 U . A ., L e n i u i u a n ., Y , H . A ., O ( A g N H.A. & U.A. R L H.A. & U.A., 1985 L Y i u a

L ( H Sininocellia gigantos Negha inflata Negha meridionalis Negha longicornis Fibla (F.) maclachlani Fibla (F.) Fibla (F.) peyerimhoffi ( N Fibla (F.) Fibla (Reisserella) pasiphae Fibla (Reisserella) Fibla (F.) hesperica N Fibla (F.) Inocellia crassicornis Inocellia bhutana Inocellia shinohara L Inocellia biprocessus Indianoinocellia mayana Inocellia taiwana Inocellia fulvostigmata H . A ., U & R Inocellia fulvostigmata nigrostigmata Inocellia sinensis Inocellia cheni Inocellia fujiana Inocellia hamata Indianoinocellia pilicornis Inocellia digitiformis Inocellia obtusangularis Inocellia elegans Inocellia japonica Parainocellia braueri Parainocellia bicolor Parainocellia ressli (H.A. & U.A., 1965) Parainocellia ressli Parainocellia burmana 1982 Amurinocellia australis Amurinocellia calida Amurinocellia sinica Tab. 1: Known stages and distribution of the 31 described valid species family Inocelliidae. Tab.

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Focal points of future research Thanks to the development of several new techniques, which allow entirely new insights, in particular microcomputer tomography, larvae as well as adults should be studied. Aside from a remarkable publication on a Raphidiid larva (Be u t e l & Ge 2008), nothing else has been published about Raphidioptera. Presently, various studies using these methods on Neuroptera are being carried out, e.g. on the tentoria, but no examinations of Raphidioptera – neither Raphidiidae, nor Inocelliidae – have yet been performed. Another important step will be a molecular study on the phylogeny on the family at generic level based on as many species as possible. It will be of particular interest to confirm or to confute that the American genus Negha is actually the sister group to the rest. Moreover, it will be important to clarify the systematic position of Indianoinocellia U.A & H.A. Furthermore, it will be an important challenge to analyze the phylogeny of the species of the large and widely distributed genus Inocellia. Last but not least, much taxonomic, biological and chorological work needs to be done. Many species are known as single specimens only, and the larvae of most species are still unknown. Large parts of the presumed distributional area of the family have never been explored, and most likely several new species will be detected. It will be a challenge to determine the southernmost occurrence of Inocelliidae.

Acknowledgements Grateful thanks to Dr. John Plant, Vienna, Austria, for linguistic improvement. This research was partly supported by the National Natural Science Foundation of China (No. 31000973 and No. 31110103002 to Xingyue Liu).

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