Research Article |
Horst Aspöck‡, Ulrike Aspöck§|
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Corresponding author: Ulrike Aspöck ( ulrike.aspoeck@nhm-wien.ac.at ) Academic editor: Dominique Zimmermann
© 2023 Horst Aspöck, Ulrike Aspöck.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Aspöck H, Aspöck U (2023) The snakeflies of the Mediterranean islands: review and biogeographical analysis (Neuropterida, Raphidioptera). Deutsche Entomologische Zeitschrift 70(1): 175-218. https://doi.org/10.3897/dez.70.101559
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Abstract
The Mediterranean region representsone of the hotspots of biodiversity of many organisms and among these also of Raphidioptera. With about 250 known species world-wide Raphidioptera is the smallest order of extant holometabolous insects.
The Mediterranean region harbors 113 species (= >45% of the world fauna), and of these 33 (= >13% of the world fauna) have been found on islands of the Mediterranean Sea. In the course of the past 50 years most of the larger islands have been intensively explored for Raphidioptera, particularly all islands (except the Baleares) which had no connection to the mainland throughout the whole Pleistocene. Altogether, 11 species of Raphidioptera (9 Raphidiidae, 2 Inocelliidae) are endemic to (usually)one, rarely to a few Mediterranean islands:
Phaeostigma (Ph.) euboica (Euboea), Ph. (Aegeoraphidia) prophetica (Rhodes), Ph. (Ae.) karpathana (Karpathos), Ph. (Ae.) biroi (Crete), Ph. (Superboraphidia) minois (Crete), Subilla principiae (Sardinia), S. colossea (Rhodes), Raphidia (R.) peterressli (Chios), R. (R.) ariadne (Crete), Fibla (F.) maclachlani (Corsica, Sardinia, Sicily), F. (Reisserella) pasiphae (Crete).
In the Aegean Sea there is a remarkable difference between the eastern and the western islands marked by the so-called Rechinger line based on results of the analysis of the flora. The Raphidioptera confirm this line impressively.
It is rather unlikely that further Raphidioptera species endemic to Mediterranean islands are still to be detected – with one exception: the Baleares. A few additional species known from various parts of the continents surrounding the Mediterranean Sea will probably be found on islands so that a total number may be around 40, possibly around 45 species.
The possible ways and times of colonization of the islands by Raphidioptera are discussed. The present paper may serve as a basis for studies on natural and particularly on anthropogenic dispersals of snakeflies from island to island, from mainland to islands, and from islands to the mainland in connection with phylogeographic investigations.
Key Words
colonization of islands, dispersal to and from islands, endemic species, Inocelliidae, Mediterranean region, Pleistocene, Raphidiidae, Rechinger line, refugial centers
Introduction
Raphidioptera
The Raphidioptera (snakeflies) is the smallest order of holometabolous insects. So far ca. 250 described valid species are known, and although several species are still to be detected, it is very unlikely that the real number of snakefly species harboring our planet presently exceeds 300. The order comprises two families: Raphidiidae (with ca. 210 known species) and Inocelliidae (with > 40 species). The two families have a similar general appearance, but are easily distinguished by several striking features. All snakeflies are characterized by an elongated prothorax, by translucent wings with a sparse wing venation and a pterostigma, by a great variety of the genital sclerites, particularly in males, and by a conspicuous ovipositor of the females (e.g. Fig.
Both families are characterized by long developmental periods. Most species need two years with at leastone hibernation of the mature larva (or rarely pupa). The larvae of the majority of species (many Raphidiidae and probably all Inocelliidae) (e.g. Figs
Today, the distribution of both families of the order is confined to arboreal parts of the Northern Hemisphere (Fig.
Most snakefly species are characterized by small, in many cases extremely small distribution areas largely restricted to glacial or postglacial refugial areas. The capacity to disperse and thus their expansivity are usually very low; there are only very few species with large distributions, e.g. from Central Europe to the Far East. These species have apparently several or even many scattered refugial areas from which they have spread.
These chorological and biogeographical characteristics are also of striking significance for an understanding of the Raphidioptera fauna of the Mediterranean islands.
Monographs and overviews on Raphidioptera: H.
The islands of the Mediterranean Sea
The Mediterranean Sea is the remnant of the ancient Tethys Ocean and today restricted to an area of about 2.1 million km2 between Europe, Asia, and Africa. It is connected with the Atlantic Ocean through the Strait of Gibraltar. At the end of the Miocene, in the Messinian, about 6 mya, the connection to the Atlantic Ocean was closed due to tectonic events. This led to the so-called Messinian salinity crisis and to the evaporation of the Mediterranean Sea. Between about 6 and 5 mya closing and opening of the Strait of Gibraltar happened several times (
Presently, there are more than 4,300 islands in the Mediterranean Sea, most of them small and uninhabited, in many cases only rocks. However, some islands are large (e.g. Sicily with more than 25,000 km2, Sardinia with more than 24,000 km2, eight islands with sizes of 1,000 km2 to almost 10,000 km2, and many of sizes of several hundred km2). Several islands have high elevations (e.g. Sicily >3300 m, Sardinia >1800 m, Corsica >2700 m, Euboea >1700 m, Crete >2450 m, Rhodes 1250 m, Cyprus >1950 m) with large forests and thus excellent ecological conditions for snakeflies.
The genesis of the islands is very different. Some of them represent fragments broken off from the margins of the continent and drifted into the sea, e.g. Sardinia, Corsica, and the Baleares broke off from Iberia in the Oligocene, between 35 and 28 mya. Others represent the exposed elevations of otherwise overflooded parts of adjacent continental areas. Crete is a good example: The island is the southernmost part of Europe in the Eastern Mediterranean. Originally the whole Aegean was land, due to subsequent tectonic events extensive transgressions occurred, which led to the Aegean Sea in the Miocene, about 15 mya. Crete remained as an island (
Sicily is an island composed of three parts of different origin. The northeasternmost part comes from the Apennine Peninsula (which consists of the Apennines onone hand and the Apulian Plate on the other hand). The northwestern part of Sicily (including central parts) derives from Iberia, namely from the land broken off from the continent in the Oligocene. The south of Sicily is of African origin (Rögl, pers. comm.).
Cyprus is derived from the Levantine mainland, the submarine separation occurred probably in the Miocene. This separated part drifted to the west and emerged from the sea later (Rögl, pers. comm.,
Rhodes was once a part of Anatolia and became separated possibly in the Pliocene due to tectonic events and subsequent transgressions (
Most of the islands near the mainland had connections to the mainland during the glacial periods of the Pleistocene, when the sea level was up to 230 m lower than today, so that an exchange of the faunas was possible. A few islands remained, however, isolated or only connected with neighboring islands, but not with the mainland. Sardinia and Corsica (Corsardinia) were sometimes connected to each other, but never to the mainland since their separation from Iberia. Crete was affected by several transgressions, but at no time was the whole island flooded; some parts always remained as islands. There were no land bridges to the European or the African mainland since the appearance of Crete in the Miocene. Neither Cyprus nor Rhodes had connections to the mainland, at least not in the Pleistocene.
Long isolation of an island from the mainland is an important precondition for the evolution of endemic species restricted to a certain island. As regards the Raphidioptera of Mediterranean islands, Sardinia, Corsica, Crete, Karpathos, and Rhodes do harbor endemic snakeflies, and none of these islands had any connection to continental regions at least throughout the whole Pleistocene. Euboea also harbors at leastone endemic snakefly, but this is due to a refugial center in high mountains (see Discussion). Interestingly, Cyprus has no endemic snakefly. So far, the order Raphidioptera has not yet been recorded from the Baleares. However, it is out of the question for snakeflies to occur on these islands, at least on Mallorca with elevations almost up to 1,500 m. See also chapter 4.3.2 on endemic species.
History of research on Raphidioptera of the islands of the Mediterranean
The first record of a snakefly from Mediterranean islands dates back to Dominicus Scina (1818), who listed “Raphidia ophiopsis” among the insects of Palermo. Raphidia ophiopsis does not occur in Sicily, so we do not know which species he had really found. At that time almost every snakefly was called “Raphidia ophiopsis”.
In addition, several species known from mainland regions were found on Mediterranean islands for the first time: Venustoraphidia nigricollis, Parvoraphidia microstigma, Ornatoraphidia flavilabris, O. christianodagmara, Phaeostigma (Magnoraphidia) major, Ph. (M.) wewalkai, Ph. (Aegeoraphidia) raddai, Subilla confinis, S. artemis, Ulrike syriaca, R. (R.) beieri, R. (R.) mysia, R. (R.) ambigua, Parainocellia ressli.
During the past fifteen years no new species have been found on any of the islands, and only further records of known species were made. Thus, the time has come to review the present state of knowledge and to analyze the genesis of the composition of the snakefly fauna of the Mediterranean islands.
Materials and methods
This comprehensive review and overview is based on several thousand specimens of Raphidioptera from Mediterranean islands identified by us over the past 60 years. Many of these studies – particularly all descriptions of new taxa – were published in many papers scattered in many journals. All publications on Raphidioptera which appeared before 1991 have been considered and cited in our monograph (
The maps showing the distribution of Raphidioptera on Mediterranean islands were provided with ArcGIS/ArcMap ver. 10.3.1.4959. Source of the map: National Geographic-Weltkarte – Content may not reflect National Geographic’s current map policy. Sources: National Geographic, ESRI, DeLorme, HERE, UNEP-WCMC, USGS, NASA, ESA, METI, NRCAN, GEBCO, NOAA, increment P Corp.
The systematic order follows
Abbreviations used in the synonymy lists: anat = anatomy; anncat = annotated catalogue; asl. = above sea level; bibliogr = bibliography; biogeogr = biogeography; biol = biology; cat = catalogue; charact = characterization; chorol = chorology; com = comment; compmorph = comparative morphology; descr = description; distr = distribution; distrmap = distribution map; ecol = ecology; ethol: cop = ethology, copulation; etymol = etymology; FD! = false determination; gs = genital segments; ill = illustration; imag = imago (adult); la = larva; list = listed; molecsyst = molecular systematics; mon = monograph; nom = nomenclature; odescr = original description; overv = overview; paras = parasites, parasitoids; phyl = phylogeny; phylogeogr = phylogeography; phyltree = phylogenetic tree; pu = pupa; rec = record; s.l. = sensu lato; s. str. = sensu stricto; synlist = synonymy list; syst = systematics; tax = taxonomy
An annotated catalogue of the Raphidioptera of the Mediterranean islands with comments on the systematics, taxonomy, biology, ecology, chorology and biogeography
In the following, the 33 species of Raphidioptera recorded from Mediterranean islands are treated with respect to their synonymies, taxonomy, systematics, biology, ecology, distribution, and biogeography (Tables
| Corsica | Sardinia | Sicily | Elba | Giglio | Krk | Hvar | Corfu | Levkas | Kefalonia | Thasos | Samothrace | Skopelos | Skyros | Euboea | Andros | Aegina | Hydra | Naxos | Paros | Crete | Karpathos | Lesbos | Samos | Icaria | Chios | Rhodos | Cyprus | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Raphidiidae | ||||||||||||||||||||||||||||
| Venustoraphidia | x | |||||||||||||||||||||||||||
| V. nigricollis | x | |||||||||||||||||||||||||||
| Xanthostigma | x | x | x | x | x | |||||||||||||||||||||||
| X. corsica | x | x | x | x | x | |||||||||||||||||||||||
| X. aloysiana | x | |||||||||||||||||||||||||||
| Parvoraphidia | x | |||||||||||||||||||||||||||
| P. microstigma | x | |||||||||||||||||||||||||||
| Ornatoraphidia | x | x | ||||||||||||||||||||||||||
| O. flavilabris | x | |||||||||||||||||||||||||||
| O. christianodagmara | x | |||||||||||||||||||||||||||
| Phaeostigma | x | x | x | x | x | x | x | x | x | x | x | x | x | |||||||||||||||
| Phaeostigma s.str. | x | |||||||||||||||||||||||||||
| Ph. (Ph.) euboica | x | |||||||||||||||||||||||||||
| Graecoraphidia | x | |||||||||||||||||||||||||||
| Ph. (G.) divina retsinata | x | |||||||||||||||||||||||||||
| Crassoraphidia | x | |||||||||||||||||||||||||||
| Ph. (C.) cyprica | x | |||||||||||||||||||||||||||
| Magnoraphidia | x | x | x | x | ||||||||||||||||||||||||
| Ph. (M.) major | x | x | ||||||||||||||||||||||||||
| Ph. (M.) flammi | x | x | ||||||||||||||||||||||||||
| Ph. (M.) wewalkai | x | |||||||||||||||||||||||||||
| Pontoraphidia | x | |||||||||||||||||||||||||||
| Ph. (P.) setulosa aegea | x | |||||||||||||||||||||||||||
| Aegeoraphidia | x | x | x | x | x | x | x | |||||||||||||||||||||
| Ph. (Ae.) raddai | x | x | x | x | ||||||||||||||||||||||||
| Ph. (Ae.) prophetica | x | |||||||||||||||||||||||||||
| Ph. (Ae.) karpathana | x | |||||||||||||||||||||||||||
| Ph. (Ae.) biroi | x | |||||||||||||||||||||||||||
| Superboraphidia | x | |||||||||||||||||||||||||||
| Ph. (S.) minois | x |
| Corsica | Sardinia | Sicily | Elba | Giglio | Krk | Hvar | Corfu | Levkas | Kefalonia | Thasos | Samothrace | Skopelos | Skyros | Euboea | Andros | Aegina | Hydra | Naxos | Paros | Crete | Karpathos | Lesbos | Samos | Icaria | Chios | Rhodos | Cyprus | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Subilla | x | x | x | x | ||||||||||||||||||||||||
| S. confinis | x | |||||||||||||||||||||||||||
| S. artemis | x | |||||||||||||||||||||||||||
| S. principiae | x | |||||||||||||||||||||||||||
| S. colossea | x | |||||||||||||||||||||||||||
| Ulrike | x | |||||||||||||||||||||||||||
| U. syriaca | x | |||||||||||||||||||||||||||
| Raphidia | x | x | x | x | x | x | x | x | x | x | x | x | x | x | x | x | ||||||||||||
| Raphidia s.str. | x | x | x | x | x | x | x | x | x | x | x | x | x | x | x | x | ||||||||||||
| R. (R.) mediterranea | x | x | x | x | x | x | x | x | x | |||||||||||||||||||
| R. (R.) beieri | x | x | ||||||||||||||||||||||||||
| R. (R.) peterressli | x | |||||||||||||||||||||||||||
| R. (R.) mysia | x | |||||||||||||||||||||||||||
| R. (R.) ambigua | x | x | ||||||||||||||||||||||||||
| R. (R.) ariadne | x | |||||||||||||||||||||||||||
| Dichrostigma | x | x | x | |||||||||||||||||||||||||
| D. flavipes | x | x | x | |||||||||||||||||||||||||
| Inocelliidae | ||||||||||||||||||||||||||||
| Fibla | x | x | x | x | ||||||||||||||||||||||||
| Fibla s.str. | x | x | x | |||||||||||||||||||||||||
| F. (F.) maclachlani | x | x | x | |||||||||||||||||||||||||
| Reisserella | x | |||||||||||||||||||||||||||
| F. (R.) pasiphae | x | |||||||||||||||||||||||||||
| Parainocellia | x | |||||||||||||||||||||||||||
| P. ressli | x |
Family Raphidiidae Latreille, 1810
Family Raphidiidae Latreille, 1810: H.
Venustoraphidia
Venustoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia nigricollis Albarda, 1891]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae corticolous on a great variety of deciduous and coniferous trees. Development usually two years. Last hibernation stage: full-grown larva. Adults: (EIV)V–VI(BVII).
Distribution
Apennine Peninsula, Balkan Peninsula, northern part of Iberian Peninsula, Central and Eastern Europe.
Venustoraphidia nigricollis
Raphidia nigricollis
Albarda, 1891 (odescr): H.
Taxonomy
H.
Biology and ecology
Larvae exclusively corticolous on Quercus, Malus, Pyrus, Acer and many other deciduous trees, but also on Pinus in light forests and in wild gardens and even in urban parks in altitudes up to ca. 500 m (Central Europe), records in Southern Europe 700 to 1100 m. Development at least two years. Last hibernating stage: full-grown larva. Adults: V–VII.
Records on Mediterranean islands
(Fig.
Continental distribution
Balkan Peninsula as far south as to the gulf of Korinthos, Eastern Europe, Apennine Peninsula (Northern Italy, Calabria), Central Europe as far north as to Northern Germany, Eastern and Southern France.
Biogeography
Polycentric Adriato-Balkanopontomediterranean faunal element with high expansivity.
Xanthostigma
Xanthostigma
Navás, 1909 (described as a section of Raphidia L.) [type species by absolute tautonymy: Raphidia xanthostigma Schummel, 1832]: H.
Raphidilla
Navás, 1915b [type species by original designation: Raphidia xanthostigma Schummel, 1832]: H.
Rhaphidilla
Navás, [1919] 1918b [unjustified emendation of Raphidilla Navás, 1915]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae of X. corsica probably mainly in upper layers of soil, but also corticolous (Quercus, Pinus). Larvae of X. xanthostigma (probably strictly) corticolous on a great variety of deciduous as well as of coniferous trees. Substrate of larvae of other species unknown. Development – as far as known – usually two years. Last hibernation stage: full-grown larva. Adults: IV–VII.
Distribution
Almost whole of Europe from southernmost parts (X. corsica) to northern parts of Scandinavia over the North of Asia as far as to Far East (X. xanthostigma). Caucasus region, Mongolia, northern China.
Xanthostigma corsica
Raphidia corsica
Hagen, 1867 (odescr): H.
Raphidia insularis
Albarda, 1891 (odescr): H.
Puncha italica
Navás, 1927b (odescr): H.
Xanthostigma corsica
(Hagen, 1867): H.
Xanthostigma corsicum
(Hagen):
Taxonomy
H.
Biology and ecology
Larvae mainly terricolous, however, several records also under bark of Pinus and of Quercus. Many records of adults mainly on low vegetation and on bushes (particularly Genista) in a great variety of different biotops: light oak and/or pine forests, various kinds of light mixed forests with a rich bush vegetation from sea level to 1500 m. Development usually two years. Last hibernation stage: full-grown larva. Adults: IV–VII.
Records on Mediterranean islands
(Fig.
Continental distribution
The species occurs on the Apennine Peninsula from the south of Calabria to Tuscany in the north; moreover it has been recorded in the south of France and the west of Spain. It is possible that the western populations are remnants of a very old invasion and might be differentiated phylogenomically from the Eastern populations. The populations on the Italian islands and on Corsica seem to be rather homogenous. One may assume that this rather frequent species is more or less regularly transferred by anthropogenic activities from the mainland or fromone island to another.
Biogeography
Polycentric Tyrrhenic-Adriato-Atlantomediterranean faunal element.
Xanthostigma aloysiana
Raphidia aloysiana
Costa, 1855 (odescr): H.
Raphidilla puella
Navás, 1915c (odescr): H.
Raphidilla soror
Navás, 1915c (odescr): H.
Xanthostigma aloysiana
(Costa): H.
Xanthostigma aloysianum
(Costa):
Taxonomy
H.
Biology and ecology
Poorly known. Larvae probably terricolous. Adults in light pine forests or light mixed forests with rich bush vegetation and in macchias of different structure in altitudes of 600 to 1200 m, usually found in single specimens only. Adults: V–VII.
Records on Mediterranean islands
(Fig.
Continental distribution
Records in all major parts of the Apennine Peninsula, in the south of Switzerland, southern France, and in the northeast of Spain in altitudes between 600 and 1200 m.
Biogeography
Monocentric Adriatomediterranean faunal element with moderate expansivity.
Parvoraphidia
Parvoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia microstigma Stein, 1863]: H.
Taxonomy and systematics
H.
Biology and ecology
The larvae of all species of the genus live in upper layers of soil. Development probably (mainly)one year. Last stage of hibernation: (probably) full-grown larva.
Distribution
Parvoraphidia is confined to the southern and southwestern parts of the Balkan Peninsula.
Biogeography
The three species represent Balkanopontomediterranean faunal elements with low expansivity.
Parvoraphidia microstigma
Raphidia microstigma
Stein, 1863 (odescr): H.
Raphidia (Parvoraphidia) microstigma
Stein: H.
Parvoraphidia microstigma
(Stein, 1863): H.
Taxonomy
H.
Biology and ecology
Larvae probably exclusively terricolous. Numerous records of adults on low vegetation, bushes and trees in various habitats: light oak and pine forests, on clearings within fir tree forests, on slopes with Genista, on hedgerows at fields in altitudes of 650–1200 m. Development possibly onlyone year. Last hibernation stage: probably full-grown larva. Adults: V–VII.
Records on Mediterranean islands
(Fig.
Continental distribution
Greece north of the gulf of Korinthos, Albania, North Macedonia.
Biogeography
Stationary monocentric Balkanopontomediterranean faunal element.
Ornatoraphidia
Ornatoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia etrusca Albarda, 1891]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae living in upper layers of soil around roots of bushes. Development two years, see under O. flavilabris. Last hibernation stage: pupa or full-grown larva. Adults: IV–VI.
Distribution
Balkan Peninsula, Apennine Peninsula, southern parts of Central Europe.
Ornatoraphidia flavilabris
Rhaphidia ophiopsis var. flavilabris
Costa, 1855 (odescr):
Raphidia etrusca
Albarda, 1891 (odescr): H.
Ornatoraphidia etrusca
(Albarda):
Ornatoraphidia flavilabris
(Costa): H.
Raphidia luigionii
Navás, 1927a (odescr): H.
Raphidia regisborisi
Navás, 1929 (odescr): H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Continental distribution
The distribution of O. flavilabris comprises southern and northwestern parts of the Balkan Peninsula with a gap in large central parts, the whole Apennine Peninsula, eastern and southern parts of Austria, northwestern parts of Hungary, and the southeast of France.
Biogeography
O. flavilabris represents a polycentric Balkanopontomediterranean-Adriatomediterranean element with moderate expansivity. The populations of the southern Balkan Peninsula show considerable genetic (but not morphological) differences to populations from the Apennine Peninsula and from Central Europe.
Ornatoraphidia christianodagmara
Raphidia (Ornatoraphidia) christianodagmara
H. Aspöck & U. Aspöck, 1970 (odescr, ecol); H.
Ornatoraphidia christianodagmara
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae are soil-dwelling (although this needs confirmation). Development two years. Pupation in late autumn or even in winter, inone case pupation took place in March. Adults: V–VI. Findings of adults in light Castanea sativa forests (Euboea) and in light fir tree (Abies cephalonica) forests (Parnis mountains).
Records on Mediterranean islands
(Fig.
Continental distribution
The species is only known from the Parnis Mts. in Attica, and in the Athmanon Mts. in Thessalia, Greece, where it occurs in altitudes of 850–1120 m.
Biogeography
Monocentric, stationary Balkanopontomediterranean faunal element.
Phaeostigma
Phaeostigma
Navás, 1909 (described as a section of Raphidia L.) [type species by subsequent designation: Raphidia notata Fabricius, 1781]:
Taxonomy and systematics
H.
Biology
Larvae of many species corticolous, of many other species soil-dwelling. Development usually two or three years. Last hibernation stage: full-grown larva. Adults: IV–VII(VIII).
Distribution
The distribution of Phaeostigma s.l. comprises Europe (except the northernmost parts of the continent, the largest part of the Iberian Peninsula, Sicily, Sardinia, Corsica and several islands in the Aegean Sea) as far as to the Ural, Anatolia, the Caucasus region, Lebanon, Syria, northern Iraq, northern Iran. Several Mediterranean islands harbor at leastone species: Levkas, Kefalonia, Thasos, Skopelos, Euboea, Crete, Karpathos, Ikaria, Samos, Lesbos, Chios, Rhodes, Cyprus.
Phaeostigma
Phaeostigma
Navás, 1909 (described as a section of Raphidia L.) [type species by subsequent designation: Raphidia notata Fabricius, 1781]: H.
Erma
Navás, 1918a (odescr) [type species by subsequent designation and monotypy: Erma abdita Navás, 1918a]: H.
Navasana
Steinmann, 1963 (odescr) [type species by original designation: Navasana perumbrata Steinmann, 1963]: H.
Note
The subgenus Phaeostigma comprises six known species,one of these occurs on the island of Euboea.
Phaeostigma (Phaeostigma) euboica
Raphidia (Phaeostigma) euboica
H. Aspöck & U. Aspöck, 1976 (odescr): H.
Phaeostigma (Phaeostigma) euboica
(H. Aspöck & U. Aspöck, 1976): H.
Taxonomy
H.
a. Venustoraphidia nigricollis, ♂. Lower Austria, Eichkogel near Mödling. Foto: H. Bruckner; b. Venustoraphidia nigricollis, ♀. Austria, Vienna. Foto: H. Bruckner; c. Venustoraphidia nigricollis, larva. Austria, Vienna. Foto: H. Bruckner; d. Xanthostigma corsica, ♀. Italy, Calabria, Aspromonte, Montalto. Foto: P. Sehnal; e. Xanthostigma corsica, larva. Itlay, Calabria, Aspromonte, Montalto. Foto: F. Anderle, now Denner; f. Parvoraphidia microstigma, ♀. Greece, Phokis, Lidorikion mts. Foto: P. Sehnal; g. Ornatoraphidia flavilabris, ♀. Greece, Phokis, Lidorikion mts. Foto: P. Sehnal; h. Ornatoraphidia flavilabris, ♂. Greece, Viotia, Parnassos. Foto: P. Sehnal.
a. Ornatoraphidia flavilabris, larva. Italy, Calabria, Sila Grande. Foto: F. Anderle, now Denner; b. Phaeostigma (Graecoraphidia) d. divina, ♂. Greece, Viotia, Parnassos. Foto: P. Sehnal; c. Phaeostigma (Graecoraphidia) d. divina, ♀. Greece, Viotia, Parnassos. Foto: P. Sehnal; d. Phaeostigma (Crassoraphidia) cyprica, ♂. Cyprus, Troodos Mountain. Foto: H. Aspöck; e. Phaeostigma (Magnoraphidia) major, ♂. Lower Austria, Dürnstein. Foto: H. Bruckner; f. Phaeostigma (Magnoraphidia) major; ♀. Lower Austria, Dürnstein. Foto: H. Bruckner; g. Phaeostigma (Magnoraphidia) major, larva. Lower Austria, Eichkogel near Mödling. Foto: H. Bruckner; h. Phaeostigma (Magnoraphidia) flammi, larva. Greece, Pilion. Foto: H. Bruckner.
a. Phaeostigma (Magnoraphidia) wewalkai, ♂. Greece, Korinthia, Oros Onia. Foto: F. Anderle, now Denner; b. Phaeostigma (Magnoraphidia) wewalkai, ♀. Greece, Korinthia, Oros Onia. Foto: P. Sehnal; c. Phaeostigma (Aegeoraphidia) biroi, ♂. Greece, Crete, Anogia > Axos. Foto: P. Sehnal; d. Phaeostigma (Aegeoraphidia) biroi, ♀. Greece, Crete, Anogia > Axos. Foto: P. Sehnal; e. Phaeostigma (Aegeoraphidia) biroi, larva. Greece, Crete, Anogia > Axos. Foto: H. Bruckner; f. Phaeostigma (Superboraphidia) minois, ♂. Greece, Crete, Levka Ori, W Omalos. Foto: H. Aspöck; g. Subilla confinis, ♂. Lower Austria, Klosterneuburg. Foto: H. Bruckner; h. Subilla principiae, ♂. Italy, Sardinia, Gennargentu, Fonni. Watercolour by Wilhelm Zelenka (1936–2011), Vienna. In the possession of H. & U. Aspöck.
a. Ulrike syriaca, ♂. Cyprus, Limassol Forest. Watercolour by Wilhelm Zelenka (1936–2011), Vienna. In the possession of H. & U. Aspöck; b. Raphidia mediterranea, ♂. Upper Austria, Pelmberg near Helmonnsödt. Foto: H. Bruckner; c. Raphidia mediterranea, ♀. Upper Austria, Pelmberg near Helmonnsödt. Foto: H. Bruckner; d. Raphidia mediterranea, larva. Upper Austria, Pelmberg near Helmonnsödt. Foto: H. Bruckner; e. Raphidia ariadne, ♂. Greece, Crete, near Males. Foto: H. Bruckner; f. Raphidia ariadne, ♀. Greece, Crete, Anogia. Foto: H. Bruckner; g. Raphidia ariadne, larva. Greece, Crete, near Males. Foto: H. Bruckner; h. Dichrostigma flavipes, ♂. Lower Austria, Eichkogel near Mödling. Foto: F. Anderle, now Denner.
a. Dichrostigma flavipes, ♀. Lower Austria, Dürnstein. Foto: F. Anderle, now Denner; b. Dichrostigma flavipes, larva. Greece, Olympos. Foto: H. Bruckner; c. Fibla (F.) maclachlani, ♂. Italy, Sardinia, Supramonte. Foto: H. Aspöck; d. Fibla (F.) maclachlani, ♀. Italy, Sardinia, Gennargentu. Foto: H. Aspöck; e. Fibla (F.) maclachlani, larva. Italy, Sardinia, Gennargentu. Foto: F. Anderle, now Denner; f. Fibla (Reisserella) pasiphae, ♂. Greece, Crete, Levka Ori, W Omalos. Foto: H. Aspöck; g. Fibla (Reisserella) pasiphae, ♀. Crete, Triphti Forest. Foto: H. Paulus; h. Fibla (Reisserella) pasiphae, larva. Greece, Crete, Anogia > Axos. Foto: F. Anderle, now Denner.
Biology and ecology
Larvae most probably exclusively corticolous, preferably on conifers (Abies, Pinus). Development two to three years. Last hibernation stage: full-grown larva. Adults: V–VI in light coniferous forests.
Records on Mediterranean islands
(Fig.
Biogeography
Ph. euboica is probably an endemism of Euboea. It represents a monocentric, extremely stationary Balkanopontomediterranean faunal element.
Graecoraphidia
Graecoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia divina H. Aspöck & U. Aspöck, 1964a]: H.
Taxonomy
H.
Biology
Larvae (of all species and subspecies?) (Fig.
Distribution
The distribution of Graecoraphidia is restricted to a small part of southern Greece.
Phaeostigma (Graecoraphidia) divina
Raphidia divina
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Phaeostigma (Graecoraphidia) divina
(H. Aspöck & U. Aspöck): H.
Note
Within this species three subspecies are differentiated, which occur in perfect geographic vicariance in small parts of southern Greece. One of these occurs on the island of Euboea.
Phaeostigma (Graecoraphidia) divinaretsinata
Raphidia (Graecoraphidia) divina retsinata
H. Aspöck & U. Aspöck, 1973 (odescr): H.
Phaeostigma (Graecoraphidia) divina retsinata
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae most probably soil-dwelling. Development under experimental conditions three years. Last hibernating stage: full-grown larva. Adults: V–VI. Most specimens were collected from fir trees in altitudes of 850–1100 m.
Records on Mediterranean islands
(Fig.
Continental distribution
Greece (Parnis, Pateras mountains).
Biogeography
Ph. (G.) divina retsinata is an extremely stationary Balkanopontomediterranean faunal element with a few refugial subcenters within a small area in Greece.
Crassoraphidia
Crassoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia cyprica Hagen, 1867]: H.
Taxonomy
H.
Biology and ecology
Larvae are mainly soil-dwelling, but sometimes found under bark. Development two to three years. Last hibernation stage: full-grown larva. Adults: IV-VI(VII) in various kinds of light forests in altitudes of 600–1500 m.
Distribution
SW- and S-Anatolia, Cyprus, Lebanon.
Phaeostigma (Crassoraphidia) cyprica
Raphidia cyprica
Hagen, 1867 (odescr): H.
Raphidia phoenicia
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Phaeostigma (Crassoraphidia) cyprica
(Hagen): H.
Phaeostigma cyprica
(Hagen):
Taxonomy
Crassoraphidia cyprica (Fig.
Biology and ecology
Larvae were repeatedly found under the bark of pine trees. Development two to three years. Last hibernation stage: full-grown larva. Adults: IV-VI. Imagines were collected particularly on pine trees, in various light forests and forest-like habitats in altitudes from 1000–1850 m.
Records on Mediterranean islands
(Fig.
Continental distribution
Lebanon.
Biogeography
Syrian (-Cyprian) faunal element. Most probably the refugial center was primarily somewhere in the Near East, from where the species invaded (passively) Cyprus. When and how this event may have happened, is unknown; it could have been during the last glacial period and/or possibly much later in the Holocene, perhaps even (repeatedly) by humans. Specimens from Cyprus cannot be differentiated morphologically from specimens from Near East. One may assume that genomic studies will lead to a better understanding.
Magnoraphidia
Magnoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia major Burmeister, 1839]: H.
Note
Magnoraphidia is – at least on the basis of characters of the male genitalia – a well differentiated subgenus with six known species. Three of these species have been found on Mediterranean islands.
Biology
Larvae of at least five species including those recorded from islands are corticolous. Development two, three or more years. Last hibernating stage: full-grown larva. Adults: (IV)V–VI(VII).
Distribution
Balkan Peninsula, Eastern Europe, Central Europe.
Phaeostigma (Magnoraphidia) major
Raphidia major
Burmeister, 1839 (odescr): H.
Phaeostigma (Magnoraphidia) major
(
Phaeostigma major
(Burmeister): H.
Phaeostigma (Magnoraphidia) majus
(Burmeister):
Phaeostigma majus
(Burmeister):
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Continental distribution
Balkan Peninsula, Central Europe (including eastern France and northern Germany), Eastern Europe.
Biogeography
Ph. (M.) major is a monocentric Balkanopontomediterranean faunal element with considerable expansivity. Ph. major is the only monocentric Balkanopontomediterranean snakefly which has reached the northern coast of Germany.
Phaeostigma (Magnoraphidia) flammi
Raphidia (Magnoraphidia) flammi
H. Aspöck & U. Aspöck, 1973 (odescr): H.
Phaeostigma (Magnoraphidia) flammi
(H. Aspöck & U. Aspöck, 1973): H.
Taxonomy
H.
Biology and ecology
Larvae (probably exclusively) corticolous on a great variety of deciduous trees and conifers in altitudes of 100 m (Skopelos) to 1100 m (Euboea). Development two to three (or more) years. Last hibernating stage: full-grown larva. Adults: V-VI.
Records on Mediterranean islands
(Fig.
Continental distribution
Confined to a small part of Greece (Sterea Ellas: Othrys, Pilion mountains).
Biogeography
Extremely stationary, monocentric Balkanopontomediterranean faunal element with several isolated populations confined to single mountain ranges.
Phaeostigma (Magnoraphidia) wewalkai
Raphidia (Magnoraphidia) wewalkai
H. Aspöck & U. Aspöck, 1971a (odescr): H.
Phaeostigma (Magnoraphidia) wewalkai
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae (probably exclusively) corticolous on deciduous trees and on conifers in various warmer habitats with rich vegetation from sea level to ca. 1000 m asl. Development two, three or more years. Last hibernation stage: full-grown larva. Adults: (IV)V–VI.
Records on Mediterranean islands
(Fig.
Continental distribution
Restricted to a small part of eastern Sterea Ellas.
Biogeography
Extremely stationary, monocentric Balkanopontomediterranean faunal element.
Pontoraphidia
Pontoraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia pontica Albarda, 1891]: H.
Phidiara
U. Aspöck & H. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia grandii Principi, 1960]: H.
Taxonomy
H.
Biology and ecology
Larvae of all species soil-dwelling in detritus between roots of bushes. Development two to three years. Adults: V-VIII, in various, particularly warmer habitats with rich vegetation of bushes in altitudes from 600–2200 m.
Distribution
The distribution comprises the southern parts of the Apennine Peninsula, the northern Balkan Peninsula, Eastern Europe and Anatolia.
Phaeostigma (Pontoraphidia) setulosa
Raphidia setulosa
H. Aspöck & U. Aspöck, 1967b (odescr): H.
Phaeostigma (Pontoraphidia) setulosa
(H. Aspöck & U. Aspöck): H.
Phaeostigma setulosa
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Phaeostigma (Pontoraphidia) setulosaaegea
Phaeostigma (Pontoraphidia) setulosa aegea
H. Aspöck, U. Aspöck & Rausch, 1991 (odescr, mon); H.
Taxonomy
H.
Biology and ecology
Larvae most probably exclusively soil-dwelling. Development probably two to three years. Last hibernating stage: probably full-grown larva. Adults: (V)–VI. Ph. (P.) s. aegea was collected on the island of Thasos in glades within light pine forests at an altitude of about 1000 m in large numbers on fruits of Asphodelus (on the Athos Mountain in 1600 m on pine trees).
Records on Mediterranean islands
(Fig.
Continental distribution
Athos mountain (Chalkidiki, Greece).
Biogeography
Ph. (P.) setulosa aegea is an extremely stationary monocentric Balkanopontomediterranean faunal element. The species – Ph. (P.) setulosa (s. l.) – is a polycentric Balkanapontomediterranean-Anatolopontomediterranean faunal element of moderate expansivity. It will certainly be found in many parts of the Balkan Peninsula and in Anatolia.
Aegeoraphidia
Aegeoraphidia
H. Aspöck & U. Aspöck & Rausch, 1991 (odescr, mon) [type species by original designation: Raphidia (Phidiara) raddai U. Aspöck & H. Aspöck, 1969]. H.
Taxonomy
H.
Biology and ecology
Larvae of all species principally soil-dwelling, but larvae of some species can also be found under bark of various trees. Development two or three years. Adults: IV–VI, in various habitats in altitudes of 0–2200 m.
Distribution
The distribution of Aegeoraphidia comprises Crete, several islands in the eastern Aegean Sea, western, southern, and southeastern Anatolia, and northern Iraq.
Phaeostigma (Aegeoraphidia) raddai
Raphidia (Phidiara) raddai
U. Aspöck & H. Aspöck, 1969 (odescr): H.
Phaeostigma (Aegeoraphidia) raddai
(U. Aspöck & H. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae corticolous as well as soil-dwelling. Findings of larvae under bark of Pinus, but also of Pyrus and Mastix as well as in the detritus between and around roots of Genista and Arbutus. Development two, three or more years. Last hibernating stage: full-grown larva. Adults: IV–VI occurring in various habitats from sea level to 800 m.
Records on Mediterranean islands
(Fig.
Continental distribution
Southwestern Anatolia.
Biogeography
Monocentric, stationary Anatolopontomediterranean faunal element comprising a relatively small area east of the Rechinger line.
Phaeostigma (Aegeoraphidia) prophetica
Raphidia prophetica
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Phaeostigma ressli prophetica
(H. Aspöck & U. Aspöck): H.
Phaeostigma prophetica
(H. Aspöck & U. Aspöck): H.
Phaeostigma (Aegeoraphidia) prophetica
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae mainly soil-dwelling, but also corticolous on various trees (Pinus, Quercus, Amygdalus). Development (at least) two years. Last stage of hibernation: full-grown larva. Adults: (IV)V–VI in various habitats, particularly light forests of Pinus and Quercus. Records from 350–750 m.
Records on Mediterranean islands
(Fig.
Biogeography
Monocentric, extremely stationary Anatolopontomediterranean faunal element. Endemic to Rhodes.
Phaeostigma (Aegeoraphidia) karpathana
Phaeostigma (Aegeoraphidia) karpathana
U. Aspöck & H. Apöck, 1989 (odescr): H.
Taxonomy
U. Aspöck and H. Aspöck (1989). In its eidonomic characters the species agrees with Ph. (Ae.) ressli and Ph. (Ae.) prophetica, but it can easily be differentiated by characters of the male and female genitalia.
Biology and ecology
Larvae soil-dwelling, but exceptionally also corticolous (findings on Pyrus with lichens on the stem). Development two years. Last hibernating stage: full-grown larva. Adults: (IV) V (-VI) in various habitats like pine forests (Pinus halepensis), open slopes with Genista and Crataegus in altitudes from 50–750 m. Adults in extraordinarily high numbers on Genista fasselata and also on young pine trees.
Records on Mediterranean islands
(Fig.
Biogeography
Monocentric, extremely stationary species, endemic to Karpathos. The species can neither be classified as Anatolopontomediterranean nor as a Cretan faunal element. It is assumed that Karpathos has represented a refugial center of its own during the glacial periods.
Phaeostigma (Aegeoraphidia) biroi
Lesna biroi
Navás, 1915a (odescr): H.
Raphidia labyrintha
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Phaeostigma biroi
(Navás): H.
Phaeostigma (Aegeoraphidia) biroi
(Navás): H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Biogeography
Monocentric, stationary Cretan faunal element. Endemic to Crete.
Superboraphidia
Superboraphidia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia auberti H. Aspöck & U. Aspöck, 1966]: H.
Taxonomy
H.
Biology
Larvae of probably all five species are soil-dwelling. Development insufficiently known, probably two years or longer. Last hibernating stage: full-grown larva. Adults: V–VII in various habitats, usually with rich low vegetation in higher elevations (800–1200 m), sometimes even above timberline.
Distribution
The distribution of Superboraphidia comprises southern parts of the mainland of Greece, the Peloponnisos and the west of Anatolia (H.
Phaeostigma (Superboraphidia) minois
Phaeostigma (Superboraphidia) minois
U. Aspöck & H. Aspöck, 1990 (odescr, ecol, distr; ill: ♀ gs, ♂ gs, wings);
Taxonomy
U. Aspöck and H.
Biology and ecology
Larvae probably soil-dwelling. Larvae and development unknown. Adults: V, on Pinus brutia and Cypressus sempervivens, in light forests with Quercus, Crataegus etc. in 830–1100 m.
a. Records of Ornatoraphidia christianodagmara, Phaeostigma (Magnoraphidia) major, Ph. (M.) flammi, Ph. (Pontoraphidia) setulosa aegea, and of Ph. (Superboraphidia) minois, on Mediterranean islands; b. Records of Phaeostigma (Grecoraphidia) divina retsinata, and of species of Aegeoraphidia, Ph. (Aegeoraphidia) raddai, Ph. (Ae.) prophetica, Ph. (Ae.) karpathana, and of Ph. (Ae.) biroi, on Mediterranean islands.
a. Records of species of the genus Raphidia, R. (R.) mediterranea, R. (R.) beieri, R. (R.) peterressli, R. (R.) mysia, R. (R.) ambigua, and R. (R.) ariadne, on Mediterranean islands; b. Records of species of the family Inocelliidae, Fibla (F.) maclachlani, Fibla (Reisserella) pasiphae, and Parainocellia ressli, on Mediterranean islands.
Records on Mediterranean islands
(Fig.
Biogeography
Endemic to Crete. Monocentric, (extremely) stationary Cretan faunal element.
Subilla
Subilla
Navás, 1916 (odescr) [type species by original designation: Raphidia sericea Albarda, 1891]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae of all species corticolous, some species particularly on Quercus, some (also or preferably) on conifers (Pinus, Abies). Development at least two or three years. Last hibernating stage: full-grown larva. Adults: (IV)V–VI(VII).
Distribution
All European peninsulas, Central, Western and Eastern Europe, Anatolia. Altogether four species have been found on Mediterranean islands: Sardinia, Sicily, Levkas, Rhodes. Each of these islands harbors onlyone Subilla species.
Subilla confinis
Raphidia confinis
Stephens, 1836 (odescr): H.
Raphidia cognata
Rambur, 1842 (odescr): H.
Raphidia schneideri
Ratzeburg, 1844 (odescr): H.
Raphidia colubroides
Costa, 1855 (odescr): H.
Raphidia sericea
Albarda, 1891 (odescrr): H.
Subilla confinis
(Stephens): H.
Taxonomy
H.
Biology and ecology
Larvae exclusively corticolous on Quercus, Acer, Pinus, Malus, very rarely (in the south of Italy) on Pinus in light forests and gardens, mostly from sea level to ca. 500 m, in the south of Italy up to 1400 m. Development usually two or three years. Last hibernating stage: full-grown larva. Adults: (IV) V–VI (VII).
Records on Mediterranean islands
(Fig.
Continental distribution
Apennine Peninsula, Central Europe, France, England, Denmark, Eastern Europe.
Biogeography
Expansive probably monocentric Adriatomediterranean faunal element.
Subilla artemis
Raphidia (Subilla) artemis
H. Aspöck & U. Aspöck, 1971 (odescr): H.
Subilla artemis
(H. Aspöck & U. Aspöck, 1971): H.
Taxonomy
H.
Biology and ecology
Larvae corticolous on Quercus, Amygdalus, Pyrus, Crataegus and other deciduous trees, very rarely on Pinus in light forests in 100–1100 m. Development two to three years. Last hibernating stage: full-grown larva. Adults: V–VI.
Records on Mediterranean islands
(Fig.
Continental distribution
Greece north of the Gulf of Korinthos, North Macedonia.
Biogeography
S. artemis is a monocentric Balkanopontomediterranean faunal element with moderate expansivity.
Subilla principiae
Subilla principiae
Pantaleoni, U. Aspöck, Cao & H. Aspöck, 2005 (odescr);
Taxonomy
Biology and ecology
So far larvae were found exclusively under the bark of Quercus pubescens. Only known from the type locality – a light oak forest – at 1050 m. Adults: V–VI.
Records on Mediterranean islands
(Fig.
Biogeography
Stationary Tyrrhenian faunal element.
Subilla colossea
Raphidia (Subilla) colossea
H. Aspöck, U. Aspöck & Rausch, 1979 (odescr): H.
Subilla colossea
(H. Aspöck, U. Aspöck & Rausch, 1979): H.
Taxonomy
H.
Biology and ecology
Larvae exclusively corticolous on Pinus and (rarely) on Quercus ilex in light pine forests from 200 to 750 m asl. Development (at least) two or three years. Last hibernating stage: full-grown larva. Adults: (IV)–V.
Records on Mediterranean islands
(Fig.
Biogeography
Stationary Anatolopontomediterranean faunal element.
Ulrike
Ulrike
H. Aspöck, 1968 (odescr) (described as subgenus of Raphidia L.) [type species by original designation: Agulla attica H. Aspöck & U. Aspöck, 1967a]: H.
Taxonomy and systematics
H.
Biology
See Ulrike syriaca.
Distribution
Greece (Attica), Cyprus, Near East.
Ulrike syriaca
Raphidilla syriaca
Steinmann, 1964 (odescr): H.
Ulrike syriaca
(Steinmann, 1964): H.
Taxonomy
H.
Biology and ecology
Larvae probably soil-dwelling. Development two years. Last hibernating stage: full-grown larva. Adults: IV–V in light forests, particularly on pines at altitudes from 100–1500 m.
Records on Mediterranean islands
(Fig.
Continental distribution
Lebanon, Syria, Jordan, Israel.
Biogeography
(Possibly polycentric) Syrian (-Cyprian) faunal element with low expansivity. Most probably the refugial center was primarily somewhere in the Near East, from where the species invaded (passively) Cyprus (see also Phaeostigma (Crassoraphidia) cyprica).
Raphidia
Raphidia
Linnaeus, 1758 (odescr) [type species by monotypy: Raphidia ophiopsis Linnaeus, 1758]: H.
Taxonomy and systematics
H.
All species can easily be differentiated by characters of the male, partly also of female genitalia.
Biology
Larvae of some species strictly corticolous, of others partly or strictly soil-dwelling. Development in some (most?) species mainlyone year, in others two (or – rarely – three) years. Last hibernating stage: full-grown larva. Adults: (III)IV–VII(VIII).
Distribution
Europe (except W- and SW-Europe), Aegean islands, Anatolia, Caucasus, N-Iran, northern and northeastern Asia.
Raphidia
Raphidia
Linnaeus, 1758 (odescr) [type species by monotypy: Raphidia ophiopsis Linnaeus, 1758]: H.
Pretzmannia
H. Aspöck & U. Aspöck, 1968 (described as a subgenus of Raphidia L.) [type species by original designation: Raphidia euxina Navás, 1915d]: H.
Raphidia (Raphidia) mediterranea
Raphidia (Raphidia) ophiopsis mediterranea
H. Aspöck, U.Aspöck & Rausch, 1977 (odescr): H.
Raphidia (Raphidia) mediterranea
H. Aspöck, U. Aspöck & Rausch: H.
Raphidia mediterranea
H. Aspöck, U. Aspöck & Rausch: H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Continental distribution
Greece, Bulgaria, Romania, Hungary, Austria, Italy, NW-Anatolia.
Biogeography
Pontomediterranean faunal element, probably monocentric, with the refugium in the south of the Balkan Peninsula. We assume that the occurrence in Eastern and Central Europe, in Italy, in NW-Anatolia and also on some islands is the result of anthropogenic dispersal, possibly already in antiquity and also presently (H.
Raphidia (Raphidia) beieri
Raphidia beieri
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Raphidia schizurotergalis
Bartoš, 1965 (odescr): H.
Raphidia (Raphidia) beieri
H. Aspöck & U. Aspöck: H.
Taxonomy
H.
Biology and ecology
Euryoecious! Larvae corticolous and soil-dwelling; many records under bark of Abies, Pinus, Quercus, Pyrus and Prunus, and in the litter around roots of bushes. Development usually two years, rarelyone or three years. Last hibernating stage: full-grown larva. Adults: IV–VI. In various habitats – light forests with conifers as well as with only deciduous trees in altitudes from 100–1700 m. On Thasos in light pine forests with Rosa, Prunus, Crataegus, Juniperus in 900–1100 m asl., on Samothraki in pastures with single old oak trees in 1200 m asl.
Records on Mediterranean islands
(Fig.
Continental distribution
Northeastern parts of Greece, North Macedonia, Kosovo, Bulgaria, Romania, Moldavia, Ukraine, NW-Anatolia.
Biogeography
Polycentric Pontomediterranean faunal element. Phylogeographic studies are urgently needed.
Raphidia (Raphidia) peterressli
Raphidia (Raphidia) peterressli
H. Aspöck & U. Aspöck, 1973 (odescr): H.
Raphidia peterressli
H. Aspöck & U. Aspöck: H.
Taxonomy
H.
Biology and ecology
Larvae not yet described, probably mainly soil-dwelling, but findings of a few larvae under bark of Pistacia lentiscus. Development probably similar to that of the closely related species. Euryoecious! Records in almost all types of vegetations with trees or bushes (pine-forests, olive-yards, all types of maquis) in altitudes from 0–1000 m.
Records on Mediterranean islands
(Fig.
Continental distribution
No records.
Biogeography
Probably endemic to Chios and thus an extremely stationary Anatolopontomediterranean faunal element.
Raphidia (Raphidia) mysia
Raphidia (Raphidia) mysia
H. Aspöck, U. Aspöck & Rausch, 1991 (mon, odescr): H.
Taxonomy
H.
Biology and ecology
Larvae at least also corticolous (findings on oak), but probably mainly soil-dwelling. Development two years. Last hibernating stage: full-grown larva. Adults: V(–VI). Euryoecious! Often high population densities in various habitats (e.g. light forests, pastures with small areas of shrubs) and on different plants: Pistacia terebinthus, Pistacia lentiscus, Phillyrea trifolia, Pinus halepensis.
Records on Mediterranean islands
(Fig.
Continental distribution
NW-Anatolia.
Biogeography
Stationary Anatolopontomediterranean faunal element.
Raphidia (Raphidia) ambigua
Raphidia ambigua
H. Aspöck & U. Aspöck, 1964b (odescr): H.
Raphidia (Raphidia) ambigua
H. Aspöck & U. Aspöck: H.
Taxonomy
H.
Biology and ecology
Larvae corticolous on conifers and deciduous trees as well as soil-dwelling. Developmentone year. Last hibernating stage: full-grown larva. Adults: (IV)V–VI(VII). Euryoecious! In various habitats: light (mixed) pine forests, oak forests, bush-forests, fruit-gardens, in maquis of various structure, usually in high population densities, in altitudes of 200 to 1600 m.
Records on Mediterranean islands
(Fig.
Continental distribution
Most parts of Anatolia, from the Mediterranean coast until the east.
Biogeography
Monocentric Anatolopontomediterranean faunal element with high expansivity.
Raphidia (Raphidia) ariadne
Raphidia ariadne
H. Aspöck & U. Aspöck, 1964a (odescr): H.
Raphidia (Raphidia) ariadne
H. Aspöck & U. Aspöck: H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Biogeography
Monocentric Cretan faunal element.
Dichrostigma
Dichrostigma
Navás, 1909 (described as a section of Raphidia L.) [type species by subsequent designation: Raphidia flavipes Stein, 1863]: H.
Lesna
Navás, 1915b (odescr) [type species by original designation: Raphidia adanana Albarda, 1891]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae soil-dwelling. Development two years. Last hibernating stage: full-grown larva. Adults: IV–VII. Euryoecious. In light pine forests as well as in habitats with deciduous trees or bushes only, in altitudes from sea level to 1800 m.
Distribution
Central-, E-, SE-Europe, Anatolia, Near East. One species occurs on Mediterranean islands.
Dichrostigma flavipes
Raphidia ophiopsis
var. e: Schummel, 1832 (descr, distr): H.
Raphidia affinis
Schneider, 1843 (odescr, ecol, distr):
Raphidia flavipes
Stein, 1863 (odescr): H.
Raphidia (Dichrostigma) flavipes
Stein: H.
Subilla sulfuricosta
Steinmann, 1963 (odescr): H.
Raphidia sinica
Steinmann, 1964 (odescr): H.
Raphidia maculicaput
Steinmann, 1964 (odescr): H.
Raphidia dichroma
Steinmann, 1964 (odescr): H.
Raphidia durmitorica
Steinmann, 1964 (odescr): H.
Raphidia monotona
Steinmann, 1964 (odescr): H.
Subilla balesdenti
Poivre, 1991 (odescr; ill: imag, gs, head, wings): H.
Raphidia (Lesna) flavipes
Stein: H.
Dichrostigma flavipes
Stein: H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Continental distribution
Balkan Peninsula (except southern parts of Greece), eastern Europe (till the Ural), Central Europe, northern Italy.
Biogeography
(Probably monocentric) Balkanopontomediterranean faunal element with high expansivity.
Family Inocelliidae Navás, 1913
Family Inocelliidae Navás, 1913: H.
Fibla
Fibla
Navás, 1915b (odescr) [type species by original designation: Fibla hesperica Navás, 1915]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae are exclusively corticolous on various trees. Development at least two, usually three (or more) years. Last hibernating stage: full-grown larva. Adults: IV–VII.
Distribution
Iberian Peninsula, Tyrrhenic islands, Sicily, Apennine Peninsula (sporadically), Crete, N-Africa.
Fibla
Subgenus Fibla Navás, 1915b (odescr) [type species by original designation: Fibla hesperica Navás, 1915]: H.
Burcha
Navás, 1915b (odescr) [type species by original designation: Inocellia maclachlani Albarda, 1891]: H.
Estoca
Navás, 1919 (odescr) [type species by monotypy: Estoca peyerimhoffi Navás, 1919]: H.
Taxonomy
H.
Biology and ecology
See Fibla s.l. and F. maclachlani.
Distribution
Iberian Peninsula, Tyrrhenian islands, Sicily, Apennine Peninsula, N-Africa.
Fibla (Fibla) maclachlani
Inocellia crassicornis
auct. (nec Schummel!): H.
Inocellia maclachlani
Albarda, 1891 (odescr): H.
Burcha maclachlani
(Albarda): H.
Burcha sicula
Navás, 1915a (odescr): H.
Fibla maclachlani
(Albarda): H.
Fibla (Fibla) maclachlani
(Albarda, 1891): H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Continental distribution
Apennine Peninsula, probably introduced from Sardinia with cork; old and recent records in Calabria need confirmation (
Biogeography
Polycentric (?) stationary Tyrrhenian-(Adriatomediterranean?) faunal element.
Reisserella
Reisserella
H. Aspöck & U. Aspöck, 1971 (odescr) (described as a subgenus of Inocellia Schneider) [type species by original designation: Inocellia (Reisserella) pasiphae H. Aspöck & U. Aspöck, 1971]: H.
Taxonomy
H.
Biology and ecology
See Fibla (Reisserella) pasiphae.
Distribution
Endemic to Crete.
Fibla (Reisserella) pasiphae
Fibla
sp.: H. Aspöck & U. Aspöck, 1966 (tax, distr). H.
Inocellia (Reisserella) pasiphae
H. Aspöck & U. Aspöck, 1971b (odescr): H.
Filba elkeweimarae
Lauterbach, 1972 (odescr): H.
Fibla (Reisserella) pasiphae
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae (Fig.
Records on Mediterranean islands
(Fig.
Biogeography
Endemic to Crete. Monocentric Cretan faunal element.
Parainocellia
Parainocellia
H. Aspöck & U. Aspöck, 1968 (odescr) (described as a subgenus of Inocellia Schneider) [type species by original designation: Inocellia ressli H. Aspöck & U. Aspöck, 1965b]: H.
Taxonomy and systematics
H.
Biology and ecology
Larvae corticolous on many coniferous as well as deciduous trees from sea level to (at least) 1200 m. Development at least two or three years. Last hibernating stage: full-grown larva. Adults: V–VII(VIII).
Distribution
Apennine Peninsula, Balkan Peninsula, Central and Eastern Europe, Samos, Anatolia.
Parainocellia ressli
Inocellia ressli
H. Aspöck & U. Aspöck, 1965b (odescr): H.
Inocellia (Parainocellia) ressli
H. Aspöck & U. Aspöck: H.
Parainocellia ressli
(H. Aspöck & U. Aspöck): H.
Parainocellia (Parainocellia) ressli
(H. Aspöck & U. Aspöck): H.
Taxonomy
H.
Biology and ecology
Larvae corticolous on Pinus, Quercus (and probably many other trees). Typical habitats: light pine forests and mixed pine-oak forests, also on single old trees, e.g. in pastures from sea level to (at least) 1800 m. Development: at least three (possibly also two) years. Last hibernation stage: full-grown larva. Adults: V–VII.
Records on Mediterranean islands
(Fig.
Continental distribution
South and east Anatolia.
Biogeography
Monocentric Anatoloponotmediterranean faunal element with low expansivity.
Discussion
The Raphidioptera of the Mediterranean islands in the mirror of the Raphidioptera of the world
Presently we know 252 described valid species of the order Raphidioptera: 206 species of Raphidiidae and 46 species of Inocelliidae. Fig.
Only 5 species (= 2%) have so far been found in Africa (restricted to the northwestern parts of the continent). America (a relatively small part of North America only: see Fig.
Fig.
Geographical definitions: Middle East comprises Turkey, Cyprus, Syria, Lebanon, Jordan, Israel, Armenia, Georgia, Azerbaijan, Iraq, Iran, and Afghanistan. Central Asia comprises Kazakhstan, Kyrgyzstan, Uzbekistan, Tajikistan, Pakistan, northern India, and Mongolia. East Asia comprises the East Asian part of Russia, China, Taiwan, Japan, Myanmar, and Thailand. In America Raphidioptera occur in Canada, USA, and Mexico.
How have the Mediterranean islands been colonized by Raphidioptera?
Tables
The Raphidioptera of the Mediterranean region a. Raphidiidae of the world (206 spp.). Two species (Raphidia ophiopsis and Xanthostigma xanthostigma) occur in Europe, Northern Asia, and Far East. In this graph they are included only in Europe and therefore counted only once; b. Raphidiidae of the Mediterranean region (104 spp.); c. Inocelliidae of the world (46 spp.). One species (Inocellia crassicornis) occurs in Europe, Northern Asia, and Far East. In this graph it is included only in Europe and therefore counted only once; d. Inocelliidae of the Mediterranean region (8 spp.).
The palaeogeography of the Mediterranean islands is very diverse – a fact which is impressively reflected also by their diverse Raphidioptera fauna. The most important factors are origin, emergence, age, and development of the different islands onone hand and whether they had connections to continental areas during the Pleistocene or whether they have been isolated for long periods, on the other hand. The desiccation by evaporation of the Mediterranean Sea between 5 and 6 mya led to many land bridges but these were of no major significance for the dispersal of the snakeflies as the dried-up areas were largely covered by evaporates or had the characters of lagoons.
Here we try to analyze the origin of the Raphidioptera fauna of those Mediterranean islands where snakeflies have been found:
Tyrrhenian islands
The Tyrrhenian islands Sardinia and Corsica were separated from the Iberian Peninsula in the Oligocene more than 32 mya and since that time never connected with continental Europe. In the Pleistocene, i.e. during 2.5 mya, they were repeatedly connected to each other for several 10,000 years so that an extensive exchange of organisms could occur. In Corsica two snakefly species have been found (Xanthostigma corsica and Fibla (F.) maclachlani), in Sardinia four species (X. corsica, X. aloysiana, Subilla principiae, and F. (F.) maclachlani). X. corsica occurs also on islands of the Tuscany archipelago (Elba, Giglio) and in Sicily. Moreover, the species is widely distributed in the Apennine Peninsula, in the southeast of France and also in isolated areas in Spain (H.
A third species of the family Raphidiidae occurring in Sardinia was detected at the beginning of this century: Subilla principiae. The species is rather isolated within the genus, and the sister taxon cannot be identified. S. principiae is endemic to Sardinia (perhaps it can be found also in Corsica) and it is certainly a very old Tyrrhenian faunal element. When and how it has come to Sardinia, is, however, unknown. The Iberian Peninsula harbors a Subilla species, S. aliena (Navás, 1915), but this species is closely related to Subilla confinis (Stephens, 1836), which is an expansive Adriatomediterranean faunal element. Moreover, all other Subilla species are Pontomediterranean faunal elements distributed in the Balkan Peninsula or in Anatolia. It is therefore unlikely,one can even exclude that S. principiae is of Iberian origin; the more enigmatic is the presence of this Tyrrhenian Subilla species in Sardinia. The fourth species of Raphidioptera occurring in Sardinia (and Corsica), Fibla (F.) maclachlani, is clearly of Iberian origin and thus a very old faunal element in Corsardinia. It occurs also in Sicily, but primarily not in the Italian mainland and can thus be regarded as an endemism of Corsardinia and Sicily. However, in recent years it has been introduced to the mainland by human activities when cork was brought from Sardinia to Tuscany (
Adriatic and Ionian islands
The islands in the Adriatic Sea as well as those in the Ionian Sea had connections to the continent, i.e. the Balkan Peninsula, during the Pleistocene. This led to an immigration of species from the continent into the former islands.
The Raphidioptera fauna of the islands of Krk, Hvar, Korfu, Levkas, and Kefalonia comprises only snakefly species which occur also in the adjacent continent: Dichrostigma flavipes (Krk, Hvar, Korfu), Parvoraphidia microstigma (Levkas), Ornatoraphidia flavilabris (Levkas), Subilla artemis (Levkas), Phaeostigma (Magnoraphidia) major (Levkas, Kefalonia).
Northern and western Aegean islands
Also the islands of the Northern Aegean Sea harbor only snakefly species of the adjacent continental regions, to which they were connected by land bridges in the Pleistocene: Phaeostigma (Pontoraphidia) setulosa aegea (Thasos), Raphidia (R.) beieri (Thasos, Samothraki), Venustoraphidia nigricollis (Thasos).
Among the Sporades only Skopelos and Skyros have been investigated for Raphidioptera. Skopelos harbors Phaeostigma (Magnoraphidia) flammi, a species which had been described from Euboea, but later had been found in the Pilion mountains on the continent. Euboea, Skopelos and the other Sporades and the continental area where the Pilion mountains are situated were connected toone land mass in the Pleistocene.
On the island of Skyros so far onlyone snakefly species has been found: Raphidia mediterranea, which also occurs in the adjacent continental parts of Greece and in Euboea.
The large island of Euboea with high mountains (up to almost 1800 m) in the north, in the center, and in the south harbors (at least) six snakefly species: Ornatoraphidia christianodagmara, Phaeostigma (Ph.) euboica, Ph. (Graecoraphidia) divina retsinata, Ph. (Magnoraphidia) flammi, Ph. (M.) wewalkai, and Raphidia (R.) mediterranea. During the Pleistocene Euboea was repeatedly widely connected with continental Greece for long periods so that there must have been an intensive faunal exchange. It is thus surprising thatone of the six species of the snakeflies of Euboea, Ph. euboeica, has so far not been found outside the island and has therefore until now been regarded as an endemism of Euboea. Another species of the subgenus Phaeostigma, Ph. (Ph.) pilicollis (Stein), is distributed all over Greece. The sister taxon to Ph. euboica is, however, not Ph. pilicollis, but the species of the Ph. notata group of the subgenus which occur in Northern parts of the Balkan Peninsula, in the Apennine Peninsula, and large parts of East, Central, and West Europe. Ph. euboica must therefore be an older isolate (possibly early Pleistocene).
Ph. (G.) divina retsinata was primarily thought to be endemic to Euboea, but was later found in two further isolated populations in two mountain ranges in Attika (Parnis, Pateras). The other two subspecies of Ph. (G.) divina, Ph. d. divina and Ph. d. simillima, are confined to small areas in continental Greece north of the Gulf of Korinthos. Two other species of the subgenus Graecoraphidia, Ph. (G.) hoelzeli H. Aspöck & U. Aspöck, 1964, and Ph. (G.) albarda Rausch & H. Aspöck, 1991, occur in small isolated areas in the north of the Peloponnisos. All species of Graecoraphidia are old Balkanopontomediterranean faunal elements of a very low expansivity, which were confined to Greece probably throughout the whole Pleistocene.
Euboea harbors two species of Magnoraphidia: Phaeostigma (M.) flammi and Ph. (M.) wewalkai. Both species occur in restricted areas of the adjacent continental region, from where they may have colonized Euboea in the Pleistocene. Ornatoraphidia christianodagmara is so far known from two localities, Parnis mountain in Attica (850–1120 m) and Ochi mountain range in the south of Euboea (1100 m). Probably the species has reached Euboea from the adjacent continental regions inone of the glacial periods.
The genus Ornatoraphidia comprises onlyone further species: O. flavilabris, which occurs in many parts of Greece, however, neither in Euboea nor in Attica.
Raphidia (R.) mediterranea is a Balkanopontomediterranean faunal element, which has been probably introduced from its primary refugial centers (in Greece) to other parts of the Balkan Peninsula and to Aegean islands, but also to Italy and even to Central Europe, probably mainly by anthropogenic dispersal and perhaps already in antiquity (H.
The only snakefly species so far recorded from Andros, from Aegina, and from Hydra is Raphidia (R.) mediterranea (see above). As regards the Kyklades, snakeflies have been found on the islands of Naxos and Paros. Naxos has been intensively investigated. Onlyone snakefly species has been found on each of these islands: Raphidia (R.) mediterranea (see above). Although the Kyklades comprise more than 30 larger (and many small) islands, it may be assumed that all these islands harbor very few snakeflies, possibly only R. mediterranea. Since the formation of the Aegean Sea due to tectonic events about 10 mya the islands had repeated connection toone another or were overflooded and emerged again, and none of the Kyklades remained isolated for a sufficiently long time to allow the formation of a stable snakefly fauna. Moreover, all these islands lack high mountains, which are an important precondition for the formation of refugial subcenters.
Crete
A totally different situation pertains to the island of Crete. Also, Crete originated when the Aegean Sea was formed. It was and is the southernmost part of Europe. And since its formation Crete remained isolated. It was often partly overflooded, but parts of the island always remained so that the original fauna of the island (at least the majority) could always survive throughout the past for ca. 10 million years. This is why Crete harbors so many endemic animals and plants. Crete harbors four snakefly species – three species of the family Raphidiidae andone Inocelliid species – all of them are endemic to Crete and have not been found elsewhere. Are they the descendants of the old fauna after the origin of Crete? Or have they, or some of them, or their ancestors reached Crete from elsewhere? It is and it will probably not be possible to answer these questions, however,one can at least try to find out the nearest relatives of these species which is always a challenging task of phylogeography.
Phaeostigma (Aegeoraphidia) biroi is related to Ph. (Ae.) karpathana (endemic to Karpathos), Ph. (Ae.) prophetica (endemic to Rhodes), Ph. (Ae.) ressli (SW-Anatolia), and Ph. (Ae.) vartianorum (SW-Anatolia). The whole subgenus is of distinct Anatolian origin, but it remains unknown how and when (the ancestors of) the species have come to Crete. Before the development of the Aegean Sea Greece and Anatolia formedone large land mass. Phaeostigma (Superboraphidia) minois (known only in a few specimens from a few localities in the Levka mountains in the west of Crete) seems systematically isolated and has been assigned to the subgenus Superboraphidia with some hesitation. Superboraphidia comprises four more species, three of them occurring in small areas in high elevations in Greece (Ph. auberti (H. Aspöck & U. Aspöck, 1966), Ph. rauschi (H. Aspöck & U. Aspöck, 1970), Ph. mammaphila (H. Aspöck & U. Aspöck, 1974)) and in the west of Anatolia (Ph. turcica (H. Aspöck & U. Aspöck & Rausch, 1981)); also, each of these species seems somewhat isolated within the subgenus. Ph. (S.) minois may be a descendent of the old fauna of Crete. The fourth snakefly species, the Inocelliid Fibla (Reisserella) pasiphae, represents a total conundrum. The subgenus Fibla is distributed in the Iberian Peninsula, Corsardinia, Sicily, and North Africa, but nowhere else. How is it possible that the sister taxon of Fibla s. str., i.e. Reisserella, occurs in Crete, entirely isolated? At firstone may think of a possible land-bridge to the north of Africa in a glacial period, but according to palaeogeological evidence such a land-bridge never existed: Crete has been isolated as an island (of changing shape) since its origin about 10 mya. It is also extremely unlikely that at some point in the past larvae of a Fibla species (or an ancestor of a Fibla species) reached Crete by wood or by swimming wood from the western part of the Mediterranean Sea. So far the question must remain open.
Karpathos
The palaeogeography of Karpathos and also – mutatis mutandis – that of Rhodes is similar to that of Crete. When the Aegean Sea developed about 10 mya Karpathos emerged as an island. Later, in the Messinian, Karpathos and Rhodes became united, but from the late Pliocene onwards Karpathos became again isolated, while Rhodes was connected with the Anatolian mainland. In the Pleistocene both islands remained isolated. This palaeogeographical history explains the snakefly fauna of Karpathos and Rhodes. Karpathos (the island has been investigated intensively by us for Raphidioptera) harbors two species of Raphidioptera, both Raphidiidae: Phaeostigma (Aegeoraphidia) karpathana and Raphidia (R.) mediterranea. Ph. (Ae.) karpathana is either the descendant of a species which inhabited the island already at the time of its origin or (more probably) of a species which invaded into Karpathos when the island was united with Rhodes in the Pliocene. The populations of R. (R.) mediterranea of Karpathos are slightly different from the populations of other parts (including islands) of Greece as well as of Italy. So far, we have hesitated to describe the Karpathos phaenon as a subspecies of R. mediterranea and we do not want to do it without a molecular systematic study. However, presently there is no adequately preserved material available. R. mediterranea has been most probably introduced from its original refugial area in Greece into many other parts of its present distribution area (H.
Rhodos
Rhodes (Rhodos) is inhabited by three species of Raphidioptera, all Raphidiidae: Phaeostigma (Ae.) prophetica, Subilla colossea, and Raphidia (R.) ambigua. It is out of the question that all three species are of Anatolian origin. R. ambigua is widely distributed in Anatolia, from where it must have colonized Rhodes. Ph. prophetica and S. colossea are endemic to Rhodes, but the sister taxa of both species occur in Anatolia.
Eastern Aegean islands
Four islands whose Raphidioptera faunas have been investigated thoroughly have to be especially considered: Lesbos, Chios, Samos, Ikaria. All four islands were regularly connected with the Anatolian mainland during the Pleistocene and also earlier. Thus, it is not surprising that these islands do not harbor endemic species – with one exception. All four islands are inhabited by Phaeostigma (Aegeoraphidia) raddai, a species which occurs in the west of Anatolia. All four islands harbor a species of Raphidia s. str.: Lesbos is inhabited by Raphidia (R.) mysia, which also occurs in the west of Anatolia, on Samos R. (R.) ambigua occurs, which is widely distributed in Anatolia (and in Rhodes). Ikaria is inhabited by R. (R.) mediterranea, a species known from many parts of Greece including several islands (see above: Euboea, Andros, Naxos, Paros, Karpathos); it has possibly been introduced to Ikaria by human activities. On the island of Chios, the subgenus Raphidia s. str. is, however, represented by Raphidia (R.) peterressli, which has so far never been found outside Chios. Is it really an endemism of Chios or is it still to be detected in Anatolia? It is of interest that all four islands harbor a different species of Raphidia s.str., and all four species occur on “their” island from coastal areas to the highest elevations. On the island of Samos Parainocellia ressli was found. The species is widely distributed in the southern parts of Anatolia almost as far as to the Iranian border.
The great Austrian botanist Karl Heinz Rechinger (1906–1998) carried out several extensive phytogeographical studies in the Aegean area (
Cyprus
Cyprus is inhabited by two snakefly species, both Raphidiidae: Phaeostigma (Crassoraphidia) cyprica and Ulrike syriaca. Both species occur in Asian countries bordering the Mediterranean Sea (Syria, Lebanon, Israel). It is surprising that none of the many Raphidioptera species occurring in the south of Anatolia have been found on the island of Cyprus which is a strong argument for the assumption that Cyprus was never connected to Anatolia. Cyprus is of Levantine origin by a submarine break off, it drifted to the west and emerged from the sea in the [late] Miocene. This means that Cyprus was without terrestrial organisms, when it appeared. It is discussed that there was a connection with the Asian continent in the late Messinian (
Open questions in research on the Raphidioptera of the Mediterranean islands
Degree of explorations of Mediterranean islands for Raphidioptera
The Mediterranean has more than 4.300 islands, from 28 of these records of Raphidioptera are available. Is this representative? This question can be clearly affirmed. The majority of the islands of the Mediterranean Sea are small, frequently rocks of low altitude and uninhabited. Generally speaking, snakeflies need forests or forest-like habitats, preferably in higher altitudes. Lowlands harbor very few species and usually only those which occur on the adjacent mainland. The most favorable conditions for snakeflies are fulfilled in large islands with elevations of 500 to 1500 m.
Of the 28 islands with records of Raphidioptera the following islands have been specifically investigated intensively for their Neuropterida and particularly for their Raphidioptera: Corsica, Sardinia, Sicily, Levkas, Thasos, Samothraki, Skopelos, Skyros, Euboea, Naxos, Crete, Karpathos, Lesbos, Samos, Ikaria, Chios, Rhodes, Cyprus.
These 18 islands comprise – with the exception of Mallorca – the 10 largest islands of the Mediterranean. 10 islands have not been particularly investigated for Raphidioptera, but single records are available: Elba, Giglio, Krk, Hvar, Korfu, Kefalonia, Andros, Hydra, Aegina, Paros. Most of these are small and near the mainland and all of them were connected with the mainland or to each other in the Pleistocene. A few additional species may be found on some of these islands, but certainly only those which occur on the adjacent mainland and certainly no endemisms. The Aegean islands of Limnos and Syros were explored for snakeflies, however none were found. The small islands off the southern coast of Anatolia, in particular Kastelorizo (highest elevation <300 m), may harborone or two snakefly species distributed in southern parts of Anatolia, which is characterized by a rich Raphidioptera fauna.
However, there are at least two groups of islands which must be investigated: the Balearic Islands and Malta. Probably no Raphidioptera will be found on any of the islands of Malta. The highest elevations of Malta are less than 250 m asl. Raphidia mediterranea could occur, if it has been introduced. There is an entirely different situation with the Balearic Islands (Mallorca, Menorca, Ibiza and Formentera). Particularly Mallorca is an island with high mountains (almost 1500 m asl.) and with large forests and forest-like habitats. The Balearic Islands were separated from the Iberian Peninsula in the Oligocene (like the later Tyrrhenian islands). One can reasonably assume that the land broken off from the mainland and having given rise to the Balearic Islands harbors the remnants of the old snakefly fauna. One might expect at least a species of Ohmella H.A. & U.A. (Raphidiidae) (probably undescribed) and a species of Fibla (probably also undescribed). So far there are no records of Raphidioptera from the Balearic Islands.
Endemic species on Mediterranean islands
So far, endemic snakefly species have mainly been found on old islands separated from the mainland by tectonic events long ago (prior to the Pleistocene or Miocene) and have remained isolated.
- Tyrrhenian islands + Sicily (Fibla (F.) maclachlani)
- Sardinia (Subilla principiae)
- Crete (Phaeostigma (Aegeoraphidia) biroi, Ph. (Superboraphidia) minois, Raphidia(R.) ariadne, Fibla (Reisserella) pasiphae)
- Rhodes (Phaeostigma (Aegeoraphidia) prophetica, Subilla colossea)
- In addition to these eight species, two species were found on single islands only:
- Phaeostigma (Ph.) euboica on Euboea and
- Raphidia (R.) peterressli on Chios.
Both islands were connected to the adjacent mainland during the Pleistocene so that it seems unlikely that they really harbor endemic species. This pertains particularly to R. peterressli as this species occurs also at low elevations as well as on the mountains all over Chios. In Phaeostigma on Euboea there is, however, a quite different situation: The species is confined to small areas at high elevations. Possibly these areas are more or less (with shifts of vertical distribution) identical with the refugial center of this extreme stationary species during the last glacial period and possibly there were no other refugial areas. Thus, Ph. euboica could really be endemic to Euboea. Nevertheless, it seems strange that the most closely related species of Ph. euboica occur far distantly on the Balkan Peninsula.
It is rather unlikely that other islands – except the Baleares – harbor endemic species so far undiscovered, but it cannot be excluded, of course.
At least on the island of Mallorca (highest elevation almost 1,500 m) Raphidioptera occur, most probably endemic species (possibly of the genera Ohmella and Fibla respectively).
Suggested molecular taxonomic and phylogeographical studies
So far all identifications of snakeflies from islands have been performed on the basis of morphological characters. All 33 species can be regarded as exactly described and there are no doubtful taxa.
Nevertheless, in a few cases molecular studies would be useful for various reasons:
- Phaeostigma (Graecoraphidia) divina retsinata: This taxon has been recorded from high altitudes in Euboea as well as in the Parnis and Pateras mountains. Other subspecies – Ph. (G.) d. divina and Ph. (G.) d. simillima – have been described from other mountains north of the Gulf of Korinthos. It would be of interest to investigate various populations of this species from various parts of the whole distribution area.
- Phaeostigma (Aegeoraphidia) spp.: Ph. (Ae.) biroi, Ph. (Ae.) karpathana, Ph. (Ae.) prophetica. These three species belong to a group of species of the subgenus to which also several species of the Anatolian mainland have been assigned. It would be of interest to analyze the relationship of these taxa on the basis of molecular phylogeny.
- Phaeostigma (Superboraphidia) minois. This species endemic to Crete has been found in a small area of western Crete on the Omalos plateau in an altitude of 1000 m. Its assignment to the subgenus Superboraphidia was made with some hesitation. Therefore, molecular systematic studies of this and other Superboraphidia species and also of species of other subgenera could clarify the systematic position of Ph. (S.) minois.
- Raphidia (R.) peterressli: This species has been found so far only on the island of Chios. Chios was repeatedly connected to the Anatolian mainland during the Pleistocene, so it is unlikely that it harbors an endemic species. Comparative molecular studies with other species of the subgenus occurring on the adjacent islands and on the Anatolian mainland would be desirable.
-
Raphidia (R.) mediterranea. This species occurs on several islands of the Aegean Sea and also in large parts of continental Greece as well as in Italy, in Eastern Europe and even in an old farmhouse in Central Europe. Comparative molecular studies of specimens from continental Greece, from Italy and from Austria have led to the conclusion that the species has been dispersed by human activities (H.
Aspöck et al. 2017 ). So far it was not possible to study specimens from islands. This would be very interesting, particularly also with respect to the slight morphological differences found in the population of Karpathos.
It is unlikely that cryptic species will be detected on Mediterranean islands by detecting significant genomic differences compared to populations of continental regions. All these species have colonized the islands quite recently. Moreover, genetic differences without morphological correlations can hardly justify the differentiation of separate species (H.
Endangered species of Raphidioptera on Mediterranean islands
Finally, we would like to raise the question whether any of the snakefly species occurring on Mediterranean islands may be endangered. Almost all species found in a certain island are distributed all over the whole island usually in a large vertical range. Despite the fact that many habitats are destroyed these species can survive in so many other areas that they are not endangered. There are, however, a few species which are confined to such small areas that they could possibly be eradicated if their habitats are destroyed: Phaeostigma (Ph.) euboica (Euboea), Ph. (Graecoraphidia) divina retsinata (Euboea), Ornatoraphidia christianodagmara (Euboea), Ph. (Superboraphidia) minois (Crete), and particularly Subilla principiae (Sardinia). This latter species has so far only been found in a small light forest of Quercus pubescens.
Acknowledgements
The photographs of living specimens were taken by Mag. Franziska Anderle, now Denner (Lower Austria, Hörersdorf), Mag. Harald Bruckner and Peter Sehnal (Natural History Museum Vienna). The photograph 6g was kindly provided by Prof. Dr. Hannes Paulus (University of Vienna). Moreover, H. Bruckner provided all distribution maps. Grateful thanks for their patience, skill and conscientiousness. Dr. Alexandra R. Szewczyk, B.A., Dr. Edwin Kniha, MSc, and Saori Huber, BSc (Vienna), and Eva Hitzinger (Lower Austria, Stockerau) were involved in the responsible documentation of the bibliographical data and of the references as well as in the digital care of the manuscript. Grateful thanks for their meticulous work.
We are much indebted to Dr. Fred Rögl (Geological-Paleontological Department, Natural History Museum Vienna) for many stimulating hours of fruitful discussion and for providing literature on the geological history of the Mediterranean Sea. Moreover, we gratefully acknowledge further information on the Mediterranean Sea in the course of the Pleistocene received from Univ. Prof. Dr. Mathias Harzhauser (Geological-Paleontological Department, Natural History Museum Vienna).
Several travels to Mediterranean islands were carried out together with colleagues and friends, to whom we want to express our sincere thanks for their company and friendship and for their assistance and help in many situations and – last but not least – for unforgettable weeks of joint field work in wonderful biotopes harboring snakeflies: Dr. Athanasios Koutsaftikis (Greece, Patras), Dr. Agostino Letardi (Italy, Roma), Dr. Laura Loru (Italy, Sardinia, Sassari), Prof. Dr. Roberto Pantaleoni (Italy, Sardinia, Sassari), Hubert and Renate Rausch (Scheibbs, Lower Austria), and Peter Ressl (Vienna).
Our cordial gratitude, though in deep sadness, goes to our son Christoph (1965–2022), who took eagerly part in searching Raphidioptera in Rhodes and in Sicily in 1979.
Sincere thanks also to Prof. Dr. Dušan Devetak (University of Maribor, Slovenia), Prof. Dr. Peter Duelli (Swiss Federal Institute for Forest, Snow and Landscape Research, Birmensdorf, Switzerland), and Prof. Dr. Michael Ohl (Museum für Naturkunde Berlin, Germany) for thoroughly reviewing and improving the manuscript, and to Dr. Dominique Zimmermann (Natural History Museum Vienna), Editor in Chief of DEZ and Subject Editor, for various good ideas and for carefully supervising the submission and review process of the manuscript.
We gratefully acknowledge the University of Vienna for providing the open access funding.
References
- Ábrahám L (1992) A Boronka-melléki Tájvédelmi Körzet Nagyszárnyú, Tevenyakú és Recésszárnyú faunájának természetvédelmi értékelése (Megaloptera, Raphidioptera, Neuroptera). Dunántúli Dolgozatok Természettudományi Sorozat 7: 107–125. [= The preservation value of the alderfly, snakefly and lacewing faunas of the Boronka-Melléki Nature Conservation Area (Megaloptera, Raphidioptera, Neuroptera).]
- Ábrahám L (2001) Somogy megye recésszárnyú-alkatúinak katalógusa (Megaloptera, Raphidioptera, Neuroptera). Natura Somogyiensis 1(1): 253–260. [= Checklist of the neuropteroid fauna of Somogy county (Megaloptera, Raphidioptera, Neuroptera).] https://doi.org/10.24394/NatSom.2001.1.253
- Ábrahám L (2006) A Mecsek recésszárnyú-alkatú faunája és újabb faunisztikai adatai (Neuropterida: Megaloptera, Raphidioptera, Neuroptera). [= The lacewing fauna of the Mecsek Mountains and its new faunistical data (Neuropterida: Megaloptera, Raphidioptera, Neuroptera). ] Folia comloensis 15: 201–218.
- Ábrahám L, Papp Z (1994) A Magyarországi Neuropteroidea Fauna Kutatásának Története (Neuropteroidea: Megaloptera, Raphidioptera, Neuroptera). Somogyi Múzeumok Közleményei 10: 159–182. [= The story of researching the Hugarian Neuropteroidea fauna (Neuropteroidea: Megaloptera, Raphidioptera, Neuroptera).]
- Ábrahám L, Markó V, Vas J (2003) Investigations on a Neuropteroid Community by Using Different Methods. Acta Phytopathologica et Entomologica Hungarica 38(1–2): 199–207. https://doi.org/10.1556/APhyt.38.2003.1-2.21
- Achtelig M (1997) Kamelhalsfliegen (Ins. Raphidioptera) aus der Umgebung von Würzburg. Galathea, Nürnberg 3(Suppl.): 48–51.
- Aistleitner U, Gruppe A (2009) Nachweise von Netzflüglern aus dem Fürstentum Liechtenstein (Insecta: Neuropterida: Raphidioptera, Megaloptera, Neuroptera). Entomofauna 30(13): 209–220.
- Albarda H (1891) Révision des Rhaphidides. Tijdschrift voor Entomologie 34: 65–184.
- Anderle F, Aspöck U (2007) Neuropterida (Insecta Endopterygota) of the Nature Reserve Eichkogel (Lower Austria): arguments for protecting an insular biocoenosis in the South of Vienna. In: Pantaleoni RA, Letardi A, Corazza C (Eds) Proceedings of the Ninth International Symposium on Neuropterology. Ferrara, Italy, 20.–23. 6. 2005. Annali del Museo Civico di Storia Naturale di Ferrara 8: 139–144.
- Aspöck H (1968) Ein weiteres neues Subgenus der Gattung Raphidia Linnaeus, 1758. Entomologisches Nachrichtenblatt, Wien 15: 65.
- Aspöck H (1990) The Raphidioptera of Africa: a review of present knowledge (Insecta: Neuropteroidea). In: Mansell MW, Aspöck H (Eds) Advances in Neuropterology. Proceedings of the Third International Symposium on Neuropterology. Bergen Dal, Kruger National Park (R.S.A.). Pretoria, 1988: 47–61.
- Aspöck H (1998) Distribution and biogeography of the order Raphidioptera: updated facts and a new hypothesis. In: Panelius S (Ed.) Neuropterology 1997. Proceedings of the Sixth International Symposium on Neuropterology, Helsinki, Finland, 13–16 July 1997. Acta Zoologica Fennica 209: 33–44.
- Aspöck H (2002) The biology of Raphidioptera: A review of present knowledge. In: Sziráki G (Ed.) Neuropterology 2000. Proceedings of the 7th International Symposium on Neuropterology, 6–9 August 2000, Budapest, Hungary. Acta Zoologica Academiae Scientiarum Hungaricae 48(Suppl. 2), 35–50.
- Aspöck H (2012) Ulrike Aspöck zum 70. Geburtstag: Ein wissenschaftshistorischer Blumenstrauß. Entomologica Austriaca: Zeitschrift der Osterreichischen Entomologischen Gesellschaft 19: 101–260.
- Aspöck H, Aspöck U (1964a) Neue Arten des Genus Raphidia L. aus Südosteuropa und Kleinasien. (Vorläufige Beschreibung). Entomologisches Nachrichtenblatt, Wien 11(6): 37–40.
- Aspöck H, Aspöck U (1964b) Eine neue europäische Spezies des Genus Raphidia Linné, R. ambigua nov. spec. (Neuroptera, Raphididae). Nachrichtenblatt der Bayerischen Entomologen 13(12): 113–116.
- Aspöck H, Aspöck U (1965a) Zur Kenntnis der Raphidiiden von Südosteuropa und Kleinasien. (Mit kritischen Bemerkungen zur Klassifikation der Familie). Annalen des Naturhistorischen Museums in Wien 68: 309–364.
- Aspöck H, Aspöck U (1965b) Vorläufige Mitteilung über Untersuchungen an europäischen Inocelliidae (Neuroptera, Raphidiodea). Entomologisches Nachrichtenblatt, Wien 12: 65–67.
- Aspöck H, Aspöck U (1966) Zur Kenntnis der Raphidiodea-Familie Inocelliidae (Insecta, Neuroptera). Annalen des Naturhistorischen Museums in Wien 69: 105–131.
- Aspöck H, Aspöck U (1967a) Agulla attica nov. spec. – eine neue Raphidiiden-Art aus Griechenland (Insecta, Neuroptera). Entomologisches Nachrichtenblatt, Wien 14: 7–11.
- Aspöck H, Aspöck U (1967b) Raphidia setulosa nov. spec. aus dem Balkan-Gebirge. Mit Bemerkungen über die Raphidiiden (Insecta, Neuroptera) Bulgariens.–. Entomologisches Nachrichtenblatt, Wien 14: 17–20.
- Aspöck H, Aspöck U (1968) Vorläufige Mitteilung zur generischen Klassifizierung der Raphidiodea (Insecta, Neuroptera). Entomologisches Nachrichtenblatt, Wien 15(7–8): 53–64.
- Aspöck H, Aspöck U (1970) Raphidia (Ornatoraphidia) christianodagmara n. sp. – eine neue europäische Raphidiiden-Spezies (Insecta, Raphidioptera). Entomologische Berichten (Amsterdam) 30: 99–102.
- Aspöck H, Aspöck U (1971a) Drei neue europäische Raphidiiden-Spezies (Neuropteroidea – Raphidioptera). Nachrichtenblatt der Bayerischen Entomologen 20: 86–88.
- Aspöck H, Aspöck U (1971b) Inocellia (Reisserella n. subgen.) pasiphae n. sp. aus Kreta (Ins., Raphidioptera, Inocelliidae). Entomologische Zeitschrift 81: 270–272.
- Aspöck H, Aspöck U (1973) Neue Raphidiiden-Spezies aus dem ägäischen Raum (Neur., Raphidioptera). Entomologische Zeitschrift 83: 193–199.
- Aspöck H, Aspöck U (1974) Zwei neue Raphidiiden-Spezies aus Griechenland (Neuropt., Raphidioptera). Entomologische Zeitschrift 84(15): 165–169.
- Aspöck H, Aspöck U (1976) Die Auflösung des Raphidia notata-Komplexes (Neur., Raphidioptera, Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 27(3/4): 57–70.
- Aspöck H, Aspöck U (1991) Raphidioptera (Snake-flies, camelneck-flies). In: Naumann ID et al. (Eds) The Insects of Australia. A textbook for students and research workers (2nd edn. Vol. I). Melbourne University Press, 521–524.
- Aspöck H, Aspöck U (2007) The Raphidioptera of the Apennine Peninsula: a biogeographical analysis. In: Pantaleoni RA, Letardi A, Corazza C (Eds) Proceedings of the 9th International Symposium on Neuropterology, Ferrara, Italy, 20–23 June 2005. Annali del Museo Civico di Storia Naturale di Ferrara 8: 95–106.
- Aspöck H, Aspöck U (2009) Raphidioptera – Kamelhalsfliegen. Ein Überblick zum Einstieg. Entomologica Austriaca: Zeitschrift der Osterreichischen Entomologischen Gesellschaft 16: 53–72.
- Aspöck H, Aspöck U (2013) Woher kommen die Namen? Die validen rezenten Taxa der Kamelhalsfliegen der Erde: Systematisches Verzeichnis und Etymologie (Insecta: Endopterygota: Neuropterida: Raphidioptera). Entomologica Austriaca: Zeitschrift der Osterreichischen Entomologischen Gesellschaft 20: 9–155.
- Aspöck H, Aspöck U (2014) Die Autoren der Taxa der rezenten Raphidiopteren (Insecta: Endopterygota). Entomologica Austriaca: Zeitschrift der Osterreichischen Entomologischen Gesellschaft 21: 9–152.
- Aspöck H, Aspöck U (2021) Die Schwarzhalsige Kamelhalsfliege – Venustoraphidia nigricollis. Insekt des Jahres 2022 Deutschland Österreich Schweiz. Herausgeber: Kuratorium Insekt des Jahres, Senckenberg Deutsches Entomologisches Institut. Achtseitiges Faltblatt.
- Aspöck H, Aspöck U (2022a) Die Schwarzhalsige Kamelhalsfliege, Venustoraphidia nigricollis (Albarda, 1891): Insekt des Jahres 2022 (Neuropterida: Raphidioptera: Raphidiidae). Entomologica Austriaca: Zeitschrift der Osterreichischen Entomologischen Gesellschaft 29: 209–220.
- Aspöck H, Aspöck U (2022b) Venustoraphidia nigricollis (Albarda, 1891) (Neuropterida, Raphidioptera, Raphidiidae): Das Insekt des Jahres 2022. Mit einem Überblick über die Ordnung Raphidioptera. Entomologische Nachrichten und Berichte 66: 1–12.
- Aspöck H, Hölzel H (1996) The Neuropteroidea of North Africa, Mediterranean Asia and of Europe: a comparative review (Insecta). In: Canard M, Aspöck H, Mansell MW (Eds) Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology. (Insecta: Neuroptera, Megaloptera, Raphidioptera). Cairo, Egypt, 2–7 May 1994, 31–86.
- Aspöck H, Aspöck U, Gruppe A (2019) Metathetely and its implications for the distribution of Raphidioptera (Insecta, Holometabola: Neuropterida). In: Weihrauch F, Frank O, Gruppe A, Jepson JE, Kirschey L, Ohl M (Eds) Proceedings of the XIII International Symposium of Neuropterology, 17–22 June 2018, Osmylus Scientific Publishers, Germany, 79–93.
- Aspöck H, Aspöck U, Rausch H (1977) Polyzentrische Ausbreitung eines „sibirisch-mediterranen“ Faunenelements am Beispiel der polytypischen Kamelhalsfliege Raphidia ophiopsis L. (Neuroptera, Raphidioptera, Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 28(4): 89–105.
- Aspöck H, Aspöck U, Rausch H (1979) Beschreibung von Raphidia (Subilla) colossea n. sp., einer neuen Raphidiiden-Spezies von Rhodes, und Bemerkungen über Raphidia (Subilla) physodes Navas (Neuropteroidea: Raphidioptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 31(1/2): 28–32.
- Aspöck H, Aspöck U, Rausch H (1981) Raphidia (Superboraphidia) turcica n. sp. – eine neue Raphidiiden-Spezies aus Anatolien (Neuropteroidea: Raphidioptera). Entomologische Zeitschrift mit Insektenbörse, Frankfurt a.M. 91(15): 169–174.
- Aspöck H, Aspöck U, Rausch H (1982) Drei neue Raphidiiden-Spezies aus Anatolien (Neuropteroidea: Raphidioptera). Nachrichtenblatt der Bayerischen Entomologen 31(4/5): 78–90.
- Aspöck H, Aspöck U, Rausch H (1989) The Raphidioptera of the Eastern Mediterranean: A Zoogeographical Analysis. Biologia Gallo-Hellenica 15: 67–111.
- Aspöck H, Aspöck U, Rausch H (1991) Die Raphidiopteren der Erde. Eine monographische Darstellung der Systematik, Taxonomie, Biologie, Ökologie und Chorologie der rezenten Raphidiopteren der Erde, mit einer zusammenfassenden Übersicht der fossilen Raphidiopteren (Insecta: Neuropteroidea). Mit 36 Bestimmungsschlüsseln, 15 Tabellen, ca. 3100 Abbildungen und ca. 200 Verbreitungskarten. Goecke & Evers, Krefeld: 2 Bde: 730 pp. [550 pp.]
- Aspöck U, Aspöck H, Rausch H (1995) Die Kopulation der Raphidiopteren: Eine zusammenfassende Übersicht des gegenwärtigen Wissensstandes (Insecta: Neuropteroidea). Mitteilungen der Deutschen Gesellschaft fur Allgemeine und Angewandte Entomologie 9: 393–402.
- Aspöck H, Aspöck U, Rausch H (1999) Biologische und chorologische Charakterisierung der Raphidiiden der östlichen Paläarktis und Verbreitungskarten der in Kasachstan, Kirgisistan, Usbekistan, Turkmenistan und Tadschikistan nachgewiesenen Arten der Familie (Neuropterida: Raphidioptera: Raphidiidae). In: Aspöck H (Ed.) Neuropterida: Raphidioptera, Megaloptera, Neuroptera. Kamelhälse, Schlammfliegen, Ameisenlöwen. Stapfia 60/ Kataloge des Oberösterreichischen Landesmuseums. Neue Folge 138: 59–84.
- Aspöck H, Hölzel H, Aspöck U (2001) Kommentierter Katalog der Neuropterida (Insecta: Raphidioptera, Megaloptera, Neuroptera) der Westpaläarktis. Denisia 02: 1–606.
- Aspöck H, Liu XY, Aspöck U (2012) The family Inocelliidae (Neuropterida: Raphidioptera). A review of present knowledge. Mitteilungen der Deutschen Gesellschaft fur Allgemeine und Angewandte Entomologie 18: 565–573.
- Aspöck H, Abbt V, Aspöck U, Gruppe A (2018) The Phenomenon of Metathetely, formerly known as Prothetely, in Raphidioptera (Insecta: Holometabola: Neuropterida). Entomologia Generalis 37(3–4): 197–230. https://doi.org/10.1127/entomologia/2018/0646
- Aspöck H, Aspöck U, Walochnik J, Kniha E (2021) Where did the Central European populations of Ornatoraphidia flavilabris (Costa) come from? (Neuropterida: Raphidioptera: Raphidiidae). Deutsche Entomologische Zeitschrift DEZ. https://doi.org/10.3897/dez.68.70814
- Aspöck H, Aspöck U, Gruppe A, Sittenthaler M, Haring E (2017) Anthropogenic dispersal of a snakefly (Insecta, Neuropterida) – a singular phenomenon or a model case in Raphidioptera? Deutsche Entomologische Zeitschrift 64(2): 123–131. https://doi.org/10.3897/dez.64.19859
- Aspöck U (2002) Male Genital Sclerites of Neuropterida: an Attempt at Homologisation (Insecta: Holometabola). Zoologischer Anzeiger 241(2): 161–171. https://doi.org/10.1078/S0044-5231(04)70071-6
- Aspöck U, Aspöck H (1969) Das Subgenus Phidiara Aspöck et Aspöck 1968 (Raphidioptera, Raphidiidae, Raphidia). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 21(4): 109–120.
- Aspöck U, Aspöck H (1989) Phaeostigma karpathana n. sp. – eine neue Kamelhalsfliege von der südostägäischen Insel Karpathos (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 41(1/2): 25–31.
- Aspöck U, Aspöck H (1990) Phaeostigma (Superboraphidia) minois n. sp. – eine neue Kamelhalsfliege aus Kreta (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 41(3/4): 71–76.
- Aspöck U, Aspöck H (1994) Paradoxe Verbreitungsbilder bei Neuropteroidea (Insecta: Raphidioptera, Neuroptera). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 46(1/2): 30–44.
- Aspöck U, Aspöck H (1999) Kamelhälse, Schlammfliegen, Ameisenlöwen ... Wer sind sie? (Insecta: Neuropterida: Raphidioptera, Megaloptera, Neuroptera). In: Aspöck H (Ed.) Neuropterida: Raphidioptera, Megaloptera, Neuroptera. Kamelhälse, Schlammfliegen, Ameisenlöwen... Stapfia 60/ Kataloge des OÖ. Landesmuseums Neue Folge 138: 1–34.
- Aspöck U, Aspöck H (2005a) Verbreitungsgrenzen von Neuropterida in Mitteleuropa. Linzer biologische Beiträge (Verhandlungen des XVIII. Internationalen Symposiums über Entomofaunistik in Mitteleuropa [SIEEC]) 37: 29–38.
- Aspöck U, Aspöck H (2005b) Neuropterida (Neuropteroidea, Neuroptera sensu lato), Ordnungen 28–30. In: Dathe HH (Ed.) 5. Teil: Insecta. Lehrbuch der Speziellen Zoologie. Begründet von A. Kaestner. Zweite Auflage. Korrigierter Nachdruck. Gruner HE (Ed.) Band I: Wirbellose Tiere. Spektrum Akademischer Verlag Heidelberg, Berlin, 540–541. [887–888.]
- Aspöck U, Aspöck H (2007) Verbliebene Vielfalt vergangener Blüte. Zur Evolution, Phylogenie und Biodiversität der Neuropterida (Insecta: Endopterygota). Denisia 20/ Kataloge der oberösterreichischen Landesmuseen. Neue Serie 66: 451–516.
- Aspöck U, Aspöck H (2009) Raphidioptera (Snakeflies). In: Resh VH, Cardé RT (Eds) Encyclopedia of Insects (2nd Edn. ). Academic Press, Elsevier Amsterdam etc., 864–866. [ISBN 978-0-12-374144-8.] https://doi.org/10.1016/B978-0-12-374144-8.00226-5
- Aspöck U, Haring E, Aspöck H (2012) The phylogeny of the Neuropterida: long lasting and current controversies and challenges (Insecta: Endopterygota). Arthropod Systematics & Phylogeny 70(2): 119–129.
- Bache F, Popescu SM, Rabineau M, Gorini C, Suc JP, Clauzon G, Olivet JL, Rubino JL, Melinte-Dobrinescu MC, Estrada F, Londeix L, Armijo R, Meyer B, Jolivet L, Jouannic G, Leroux E, Aslanian D, Reis ATD, Mocochain L, Dumurdžanov N, Zagorchev I, Lesić V, Tomić D, Namık Çağatay M, Brun J-P, Sokoutis D, Csato I, Ucarkus G, Çakır Z (2012) A two-step process for the reflooding of the Mediterranean after the Messinian Salinity Crisis. Basin Research 24(2): 125–153. https://doi.org/10.1111/j.1365-2117.2011.00521.x
- Badano D (2008) Contributo alla Conoscenza dei Neurotteri della Liguria (Insecta, Neuropterida). Annali del Museo Civico di Storia Naturale “G. Doria 99: 535–548.
- Badano D, Letardi A (2010) A review of the Neuropterida of Liguria (north-west Italy). Proceedings of the Tenth International Symposium on Neuropterology, Maribor, 83–87.
- Badano D, Makris C (2020) A new unexpected species of Chrysopa Leach from Cyprus with biogeographic remarks (Neuroptera: Chrysopidae). Annales de la Société entomologique de France (NS) (Vol. 56, No. 2), 125–134. https://doi.org/10.1080/00379271.2020.1756911
- Bartoš E (1965) Raphidia schizurotergalis n.sp., eine Raphidien-Art aus der Moldavischen SSR (Raphidioptera). Acta Entomologica Bohemoslovaca 62: 129–138.
- Burmeister HCC (1839) Handbuch der Entomologie. 2. Bd. Besondere Entomologie. 2. Abt. Kaukerfe. Gymnognatha. (2. Hälfte, vulgo Neuroptera). Th. Chr. F. Enslin, Berlin, 757–1050.
- Canbulat S (2014) Checklist of Turkish Raphidioptera on the basis of distribution pattern and biogeographical analysis. Turkish Journal of Zoology 38: 1–10.
- Canbulat S, Kiyak S (2006) Contribution to the distribution of Raphidioptera (Raphidiidae, Inocelliidae) from the Southwest Anatolia. Acta Musei Reginaehradecensis S. A. 31: 141–147.
- Canbulat S, Özsaraç Ö (2004) Neuropterida (Insecta; Raphidioptera, Neuroptera) fauna of çiçekdaği (Kirşehir Province). Journal of the Institute of Science and Technology. Gazi University 17: 1–9.
- Costa A (1855) Fauna del Regno di Napoli. Nevrotteri. Rafidiidei.: 9 pp. In: Costa A (Ed.) Fauna del Regno di Napoli ossia enumerazione di tutti gli animali che abitano le diverse regioni di questo Regno e le acque che le bagnano e descrizione de’nuovi o poco esattamente conosciuti con figure ricavate daoriginali viventi e dipinte al naturale. Nevrotteri. Stamperia di Antonio Cons, Napoli, 83 pp. [7 pl.]
- Czechowska W (2002) Raphidioptera and Neuroptera (Neuropterida) of the canopy in montane, upland and lowland fir forests of Abies alba Mill. in Poland. Fragmenta Faunistica 45(1): 31–56. https://doi.org/10.3161/00159301FF2002.45.1.031
- Devetak D (1991) Neuropteroidea. Megaloptera, Raphidioptera, Planipennia (Insecta). Fauna Durmitora 4: 135–159.
- Devetak D (1992a) Megaloptera, Raphidioptera and Planipennia (Neuropteroidea, Insecta) of Croatia. Znanstvena Revija 4: 89–114.
- Devetak D (1992b) Mrežekrilci Kraškega roba. Proteus (Shippensburg, Pa. ) 54: 239–242.
- Devetak D (1995) New records for the neuropteroid fauna of Slovenia (Raphidioptera, Neuroptera). Acta Entomologica Slovenica (Ljubljana) 3: 49–57.
- Devetak D, Rausch H (2016) Checklist of Lacewings (Neuropterida: Raphidioptera, Megaloptera, Neurotpera) of Albania. Acta Zoologica Bulgarica 68(4): 457–467.
- Devetak D, Klokočovnik V, Rausch H, Janžekovič F (2015) Fauna of the Neuropterida (Raphidioptera, Neuroptera) of the Protected Area Jasen, Macedonia: A summer flash. Turkish Journal of Zoology 39: 15–27. https://doi.org/10.3906/zoo-1305-43
- Dobosz R (1989) Siatkoskrzydłe (Neuropteroidea) w zbiorach Muzeum Górnośląskiego w Bytomiu. Rocznik Muzeum Górnośląskiego w Bytomiu, Przyroda. [= Annals of the Upper Silesian Museum in Bytom, Natural History. ] 12: 75–90.
- Dobosz R (1991) Snake-Flies [sic!] (Raphidioptera, Neuropteroidea) of Poland – a faunistic review. Annals of the Upper Silesian Museum, Entomology. [= Rocznik Muzeum Górnośląskiego, Entomologia. ] 2: 191–208.
- Dobosz R (1994) Materiały do fauny wielbłądek (Neuropteroidea: Raphidioptera) Polski. Acta Entomologica Silesiana 2: 23. [= The material to the fauna of snake-flies (Neuropteroidea: Raphidioptera) of Poland.]
- Dobosz R (1999) Owady siatkoskrzydłe (Neuropteroidea: Raphidioptera, Neuroptera) Puszczy Białowieskiej – stan poznania. Parki Narodowe i Rezerwaty Przyrody 18(supl.): 125–132.
- Dobosz R (2003) Owady siatkoskrzydłe (Neuropterida: Megaloptera, Raphidioptera, Neuroptera) Babiej Góry. [= Neuropteroid insects (Neuropterida: Megaloptera, Raphidioptera, Neuroptera) of Mt. Babia Góra.] In: Wołoszyn BW, Wołoszyn D, Celary W (Eds) Monografia fauny Babiej Góry. Komitet Ochrony Przyrody Polskiej Akademii Nauk, Krakow, 223–234.
- Dobosz R (2007) Contribution to the knowledge of snakeflies (Raphidioptera) of Turkey. Annals of the Upper Silesian Museum (Entomology) 14–15: 43–58.
- Fabricius JC (1781) Species insectorum exhibentes eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. Tom 1. Impensis Carol Ernst Bohnii, Hamburgi et Kilonii, 552 pp. https://doi.org/10.5962/bhl.title.36509
- Gruppe A (1997) Beitrag zur Kenntnis der Raphidiopteren-Fauna Oberbayerns (Neuropteroidea). Nachrichtenblatt der Bayerischen Entomologen 46: 26–28.
- Gruppe A (2006a) Lebensraum für Kamelhalsfliegen. Totholz im Kronenbereich – begehrter Lebensraum. LWF aktuell 53/2006: 10–11.
- Gruppe A (2006b) Vergleich der Neuropteren-Zönosen auf Tanne und Fichte in Bayern (Neuropterida: Raphidioptera, Neuroptera). Beiträge zur bayerischen Entomofaunistik 8: 1–11.
- Gruppe A (2008) Diversity and host tree preference of Neuropterida (Insecta) in mixed forest stands in Germany. In: Floren A, Schmidl J (Eds) Canopy Arthropod Research in Europe. Basic and applied studies from the high frontier. Bioform entomology, Nürnberg, 145–156.
- Gruppe A (2015) Netzflügler im Englischen Garten in München. Beiträge zur bayerischen Entomofaunistik 15: 46–50.
- Gruppe A, Müller J (2007) Distribution of Neuropterida in beech dominated forests in southern Germany. In: Pantaleoni RA, Letardi A, Corazza C (Eds) Proceedings of the Ninth International Symposium on Neuropterology, Ferrara, Italy, 20–23 June 2005. Annali del Museo Civico di Storia Naturale di Ferrara 8: 145–152.
- Gruppe A, Schubert H (2001) The spatial distribution and plant specificity of Neuropterida in different forest sites in Southern Germany (Raphidioptera and Neuroptera). Beiträge zur Entomologie 51(2): 517–527. https://doi.org/10.21248/contrib.entomol.51.2.517-527
- Gruppe A, Aspöck H, Aspöck U (2017) Das Strohdach des Mittermayrhofs in Pelmberg (Oberösterreich) – Kinderstube und Puppenwiege einer mediterranen Kamelhalsfliege (Insecta: Neuropterida: Raphidioptera: Raphidiidae). Linzer Biologische Beitrage 49(1): 577–583.
- Gruppe A, Aspöck H, Aspöck U (2023) How many larval instars do Raphidioptera have? (Insecta, Neuropterida). Spixiana 45(2): 241–248.
- Gruppe A, Gossner M, Simon U (2004) Neuropteren in Wäldern Schwabens und Oberbayerns. Beiträge zur bayerischen Entomofaunistik 6: 243–254.
- Gusenleitner F (2004) Dokumente zum wissenschaftlichen Opus von Horst Aspöck. In: Aspöck U (Ed.) Entomologie und Parasitologie. Festschrift zum 65. Geburtstag von Horst Aspöck. Denisia 13: 23–28.
- Güsten R (1998) Segnalazioni faunistiche italiane. 331 – Raphidia mediterranea H. Aspöck, U. Aspöck & Rausch, 1977 (Raphidioptera Raphidiidae). Bollettino della Società Entomologica Italiana 130: 80–81.
- Hagen HA (1867) Notes on the genus Raphidia. The Transactions of the Entomological Society of London 3/5: 493–499.
- Haring E, Aspöck U (2004) Phylogeny of the Neuropterida: A first molecular approach. Systematic Entomology 29(3): 415–430. https://doi.org/10.1111/j.0307-6970.2004.00263.x
- Haring E, Aspöck H, Bartel D, Aspöck U (2011) Molecular phylogeny of the Raphidiidae (Raphidioptera). Systematic Entomology 36(1): 16–30. https://doi.org/10.1111/j.1365-3113.2010.00542.x
- Hellrigl K, Hölzel H (1996) Neuropteroidea – Netzflügler. In: Hellrigl K (Ed.) Die Tierwelt Südtirols. Kommentiertes systematisch-faunistisches Verzeichnis der auf dem Gebiet der Provinz Bozen – Südtirol (Italien) bekannten Tierarten. Veröffentlichungen des Naturmuseums Südtirol, Bozen 1: 515–521.
- Hiermann U, Kopf T, Gruppe A (2018) Nachweise von Kamelhalsfliegen (Insecta: Neuropterida: Raphidioptera) aus Vorarlberg und Nordtirol (Austria occ.), sowie Streudaten aus Europa. Inatura – Forschung online 55: 1–5.
- Hofrichter R (2020) Das Mittelmeer: Geschichte und Zukunft eines ökologisch sensiblen Raums. Springer-Verlag, 1267 pp. https://doi.org/10.1007/978-3-662-58929-8
- Horstmann K (1993) Neue Taxa der Campopleginae aus den Gattungen Campoplex Gravenhorst, Diadegma Förster und Nemeritis Holmgren (Hymenoptera, Ichneumonidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 44: 116–127.
- Horstmann K (1994) Nachtrag zur Revision der westpaläarktischen Nemeritis-Arten (Hymenoptera, Ichneumonidae, Campopleginae). Mitteilungen der Münchner Entomologischen Gesellschaft 84: 79–90.
- Jolivet L, Augier R, Robin C, Suc J-P, Rouchy JM (2006) Lithospheric-scale geodynamic context of the Messinian salinity crisis. Sedimentary Geology 188–189: 9–33. https://doi.org/10.1016/j.sedgeo.2006.02.004
- Kielhorn KH (1991) Fluginsekten im Kronenraum von Kiefern und Eichen. Diplomarbeit. Institut für Angewandte Zoologie und Tierphysiologie, Bodenzoologie und Ökologie, Freie Universität Berlin, 90 pp.
- Klokočovnik V, Devetak D, Gomboc S (2010) Neuropterida (Megaloptera, Raphidioptera, Neuroptera) of Kozjanski Regional Park, Slovenia. In: Devetak D, Lipovšek S, Arnett AE (Eds) Proceedings of the Tenth International Symposium on Neuropterology. Piran, Slovenia, 22–25 June 2008: 171–174. University of Maribor, Maribor.
- Lange L (2010) 936. Ratzeburgs Kamelhalsfliege Puncha ratzeburgi (Brauer, 1876) und andere Kamelhalsfliegen aus Norddeutschland (Raphidioptera). Entomologische Nachrichten und Berichte 54(1): 56–57.
- Latreille PA (1810) Considerations generates sur l‘ordre naturel des animaux composant les classes des Crustaces, des Arachnides et des Insectes; avec un tableau methodique de leurs genre disposes en families. F. Schoell, Paris, 444 pp. https://doi.org/10.5962/bhl.title.13342
- Lauterbach KE (1972) Beschreibung zweier neuer europäischer Inocelliiden (Insecta, Raphidioptera), zugleich ein Beitrag zur vergleichenden Morphologie und Phylogenie der Kamelhalsfliegen. Bonner Zoologische Beitrage 23: 219–252.
- Letardi A (1993) [1994] Nota sui reperti di Neurotteri dei Monti Lepini (Lazio). (Neuropteroidea). Quaderni del Museo di Storia Naturale di Patrica 4: 93–99.
- Letardi A (1994) Dati sulla distribuzione italiana di Megaloptera Sialidae, Raphidioptera Inocelliidae e Planipennia Mantispidae, con particolare riferimento all’Italia Centrale. (Neuropteroidea). Bollettino della Società Entomologica Italiana 125: 199–210.
- Letardi A (2015) Stato delle conoscenze sui Neuropterida dei Monti Lepini (Lazio). In: Corsetti L, Angelini C, Copiz R, Mattoccia M, Nardi G (Eds) Biodiversità dei Monti Lepini. Edizioni Belvedere, Latina, 199–207.
- Letardi A, Maltzeff P (2001) Neurotteridi e Mecotteri della Tenuta Presidenziale di Castelporziano e delle Aree Limitrofe (Neuroptera, Raphidioptera, Mecoptera). Bollettino dell’Associazione Romana di Entomologia 56: 49–62.
- Letardi A, Maltzeff P (2008) I Neuropterida della Tenuta Presidenziale di Castelporziano. Addenda (Neuroptera, Raphidioptera). Bollettino dell’Associazione Romana di Entomologia 63: 83–89.
- Letardi A, Migliaccio E (2002) Neuropterida of the Abruzzo National Park, Italy. In: Sziráki G (Ed.) Proceedings of the Seventh International Symposium on Neuropterology. Budapest, Hungary, 6–9 August 2000. Acta Zoologica Academiae Scientiarum Hungaricae 48 (Supplement 2), 149–154.
- Letardi A, Pantaleoni RA (1996) I Neurotteroidei W-Paleartici della collezione del Museo di Zoologia dell–Università di Roma (Neuropteroidea). Fragmenta Entomologica 28: 277–305.
- Letardi A, Thierry D, Tillier P, Canard M (2008) Mise à jour de la faune des Neuropterida de Corse (Raphidioptera & Neuroptera). Revue de l’Association Roussillonnaise d’Entomologie XVII(3): 95–105.
- Letardi A, Nicoli Aldini R, Pantaleoni RA (2010) The Neuropterida of Triveneto (Northern Italy): an updated faunal checklist with some zoogeographical remarks. In: Devetak D, Lipovšek S, Arnett AE (Eds) Proceedings of the Tenth International Symposium on Neuropterology. Piran, Slovenia, 22–25 June 2008: 181–189. University of Maribor, Maribor.
- Linnaeus C (1758) Sytema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species, cum Characteribus, Differentiis, Synonymis, Locis. Ed. X., Tom. I. , Holmiae, Laurentii Salvii, 824 pp. https://doi.org/10.5962/bhl.title.542
- Liu XY, Aspöck H, Yang D, Aspöck U (2009) Discovery of Amurinocellia H. Aspöck & U. Aspöck (Raphidioptera: Inocelliidae) in China, with description of two new species. Zootaxa 2264(1): 41–50. https://doi.org/10.11646/zootaxa.2264.1.3
- Lu X, Zhang W, Wang B, Engel MS, Liu XY (2020) A new and diverse paleofauna of the extinct snakefly family Baissopteridae from the mid-Cretaceous of Myanmar (Raphidioptera). Organisms, Diversity & Evolution 20(4): 565–595. https://doi.org/10.1007/s13127-020-00455-y
- Monserrat VJ (1996) Lista de los neuropteros de Aragon (Megaloptera, Raphidioptera, Planipennia). Catalogus de la entomofauna aragonesa 11: 11–17.
- Monserrat VJ, Papenberg D (2010) Revisión del género Phaeostigma Navás, 1909 de la Península Ibérica (Insecta, Raphidioptera). Graellsia 66(1): 47–77. https://doi.org/10.3989/graellsia.2010.v66.010
- Monserrat VJ, Papenberg D (2012) Revisión del género Venustoraphidia Aspöck & Aspöck, 1968 de la Península Ibérica (Insecta, Raphidioptera). Graellsia 68(2): 291–304. https://doi.org/10.3989/graellsia.2012.v68.061
- Monserrat VJ, Papenberg D (2015) Los Rafidiópteros de la Península Ibérica (Insecta, Neuropterida: Raphidioptera). Graellsia 71(1): e024. https://doi.org/10.3989/graellsia.2015.v71.116
- Morgan MJ (1993) Recording in Wales: Part 7. Neuro News. The Newsletter of the British Isles Neuroptera Recording Scheme 13: 5–7.
- Morinière J, Hendrich L, Hausmann A, Hebert P, Haszprunar G, Gruppe A (2014) Barcoding Fauna Bavarica: 78% of the Neuropterida Fauna Barcoded! PLoS ONE 9(10): e109719. https://doi.org/10.1371/journal.pone.0109719
- Myers N, Mittermeier RA, Mittermeier CG, Da Fonseca GA, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403(6772): 853–858. https://doi.org/10.1038/35002501
- Navás L (1909) Les Raphidides (Insectes, Neuropteres) du Musée de Paris. Annales de la Société Scientifique de Bruxelles 33: 143–146.
- Navás L (1913) Neurópteros del R. Museo Zoológico de Nápoles. Annuario del Museo Zoologico della R. Università di Napoli 4: 1–11.
- Navás L (1915a) Raphididae Musei Nationalis Hungarici. Annales Historico-Naturales Musei Nationalis Hungarici 13: 331–336.
- Navás L (1915b) Neurópteros nuevos o poco conocidos. Quarta serie & Quinta serie. Memorias de la Real Academia de Ciencias y Artes de Barcelona 11: 373–398. [455–480.]
- Navás L (1915c) Neue Neuropteren. Zweite Serie. Entomologische Mitteilungen 4: 194–202.
- Navás L (1915d) Notas sobre Rafididos (Ins., Neur.). Revista de la Real Academia de Ciencas Exactas & Fisicas y Naturales de Madrid 13: 784–797. [860–874.]
- Navás L (1916) Notas sobre el orden de los Rafidiópteros (Ins.). Memorias de la Real Academia de Ciencias y Artes de Barcelona 3(12): 507–513.
- Navás L (1918a) Neurópteros nuevos ó poco conocidos. X. Memorias de la Real Academia de Ciencias y Artes de Barcelona 14: 339–366.
- Navás L (1918b) Monografia de l’ordre dels Rafidiòpters (Ins.). Arxius de l’Institut de Ciencias, Barcelona 90 pp.
- Navás L (1919) Rafidiópter (Ins.) nou. Publicaciones del Instituto de Ciencias de Barcelona 1918: 91–93.
- Navás L (1927a) Mis excursiones cientificas en 1926. Revista de la Academia de Ciencias Exactas. Fisico-Quimicas y Naturales de Zaragoza 10: 81–124.
- Navás L (1927b) Insecta nova. Series XII. Memorie della Pontificia Accademia Romana dei Nuovi Lincei 10(2): 1–10.
- Navás L (1929) Insectes Nevropteres de Bulgarie. Izvestiya na Tsarskite Prirodonauchni Instituti v Spfiya 2: 140–142.
- Nicoli Aldini R (2005) I Neurotteri (Insecta Neuropterida) della Val Camonica e del Parco Naturale dell’Adamello (Alpi e Prealpi Lombarde): Materiali per uńanalisi zoogeografica del popolamento. Biogeographica 26: 417–430. https://doi.org/10.21426/B626110579
- Nicoli Aldini R, Baviera C (2001) Nuove acquisizioni sulla neurotterofauna di sicilia e isole circumsiciliane (Insecta Neuropterida). Naturalista Siciliano S. IV XXV(3–4): 345–376.
- Nicoli Aldini R, Letardi A, Pantaleoni RA (2012) State of the art on Neuropterida of Sicily and Malta. Biodiversity Journal 3(4): 445–458.
- Novak GB (1891) Terzo cenno sulla Fauna dell’Isola Lesina in Dalmazia. Neuroptera. Glasnik Hrvatskoga Naravoslovnoga Društva 6: 50–58.
- Oswald JD, Machado JP (2018) Biodiversity of the Neuropterida (Insecta: Neuroptera, Megaloptera, and Raphidioptera). In: Footit RG, Adler PH (Eds) Insect Biodiversity: Science and Society (Vol. II, 1st Edn.). Wiley-Blackwell, Oxford, Chichester, Hoboken, 627–671. https://doi.org/10.1002/9781118945582.ch21
- Oswald JD, Penny ND (1991) Genus-group names of the Neuroptera, Megaloptera and Raphidioptera of the world. Occasional Papers of the California Academy of Sciences 147: 1–94. https://doi.org/10.5962/bhl.part.3428
- Pantaleoni RA (1990a) I Neurotteri (Neuropteroidea) della Valle del Bidente-Ronco (Appennino Romagnolo). Bollettino dell’Istituto di Entomologia “Guido Grandi” della Universitä degli studi di Bologna 44: 89–122.
- Pantaleoni RA (1990b) Neurotteri e fasce de vegetazione in Romagna. Bollettino dell'Istituto di Entomologia "Guido Grandi" della Università degli Studi di Bologna 44: 143–154.
- Pantaleoni RA (1990c) I Neurotteri (Insecta Neuropteroidea) della collezione dell’Istituto di Entomologia Agraria dell’Università di Padova. Bollettino dell’Istituto di Entomologia “Guido Grandi” della Università degli Studi di Bologna 45: 73–99.
- Pantaleoni RA (1990d) [1993] I neurotteri (Insecta Neuropteroidea) delle collezioni “Zangheri” (Museo di Storia Naturale della Romagna) e “Malmerendi” (Museo Civico di Scienze Naturali di Faenza). Bollettino del Museo Civico di Storia Naturale di Verona 17: 277–292.
- Pantaleoni RA (1999) Neuropterida described by A. Costa with type designation. Deutsche Entomologische Zeitschrift 46: 249–261. https://doi.org/10.1002/mmnd.4800460211
- Pantaleoni RA (2005) Interpretation of Achille Costa’s data on Neuropterida. Bulletin of Insectology 58: 71–92.
- Pantaleoni RA (2007) Perspectivas del uso de Raphidioptera y Neuroptera Coniopterygidae como agentes de control biológico. In: Rodríguez-Del-Bosque ELA, Arredondo-Bernal HC (Eds) Teoría y Aplicación del Control Biológico. Sociedad Mexicana de Control Biológico, México, 106–126.
- Pantaleoni RA, Campadelli G, Crudele G (1994) Nuovi dati sui Neurotteri dell’alto Appennino romagnolo. Bollettino dell’Istituto di Entomologia “Guido Grandi” della Università degli Studi di Bologna 48: 171–183.
- Pantaleoni RA, Aspöck U, Cao OV, Aspöck H (2005) Subilla principiae n. sp., a new spectacular snakefly from Sardinia (Raphidioptera Raphidiidae). Redia 87(2004): 3–6.
- Pantaleoni RA, Cocco A, Floris I, Letardi A, Loru L (2019) Going overseas: From island to continent colonization in the Mediterranean snakefly Fibla maclachlani (Albarda, 1891). BioInvasions Records 8(2): 442–451. https://doi.org/10.3391/bir.2019.8.2.27
- Plant CW (1992) A working list of the lacewings of the London Area. The London Naturalist 71: 117–136.
- Plant CW (1994) Provisional atlas of the lacewings and allied insects (Neuroptera, Megaloptera, Raphidioptera and Mecoptera) of Britain and Ireland. Biological Records Centre. NERC Institute of Terrestrial Ecology, Monks Wood, Huntingdon, England, 203 pp.
- Plant CW (2001) Snake-flies, alderflies and lacewings (Neuropterida) and scorpion flies (Mecoptera): A provisional review of their status and distribution in Essex with notes on their etymology. Essex Naturalist. New Series 18: 177–218.
- Poivre C (1991) Subilla balesdenti n.sp. (Raphidioptera, Raphidiidae). Neuroptera International 6: 125–129.
- Popov A (1992) Zoogeographical analysis of Neuropteroidea (Insecta) of the Balkan Peninsula. In: Canard M, Aspöck H, Mansell MW (Eds) Current Research in Neuropterology. Proceedings of the Fourth International Symposium on Neuropterology. Bagnères-de-Luchon, Haute-Garonne (France), 24–27 June 1991. (Insecta: Neuroptera, Megaloptera, Raphidioptera), 319–330.
- Popov A (1993) Raphidiopteren und Neuropteren aus Bulgarien in den Sammlungen des Nationalmuseums in Prag. Historia Naturalis Bulgarica 4: 16–28.
- Popov A (1997) Neuroptera, Raphidioptera and Mecoptera from Macedonia with two new records of Chrysopidae. Historia Naturalis Bulgarica 7: 31–33.
- Popov A (2000a) Entomofaunistic diversity of the Central Balkan National Park. Orthopteroidea, Neuropteroidea, Mecoptera. In: Sakalian M (Ed.) Biological Diversity of the Central Balkan National Park. Pensoft, Sofia, 540–542. [571–573.]
- Popov A (2000b) Entomofaunistic diversity of the Rila National Park. Orthopteroidea, Neuropteroidea, Mecoptera. In: Sakalian M (Ed.) Biological diversity of the Rila National Park. Pensoft, Sofia, 527–531. [581–584.]
- Popov A (2001) The snakeflies and the lacewing insects (Insecta: Raphidioptera and Neuroptera) of the Kresna Gorge (SW Bulgaria). [Bulgarisch.] In: Beron P (Ed.) Biodiversity of Kresna Gorge (SW Bulgaria), 131–143.
- Popov A (2002) Neuropterida of Northern Europe. In: Sziráki G (Ed.) Neuropterology 2000. Proceedings of the Seventh International Symposium on Neuropterology. Acta Zoologica Academiae Scientiarum Hungaricae 48(Supplementum 2), 281–291.
- Popov A (2004) List of Raphidioptera, Neuroptera and Mecoptera published from the Eastern Rhodopes (Bulgaria and Greece). In: Beron P, Popov A (Eds) Biodiversity of Bulgaria. 2. Biodiversity of Eastern Rhodopes (Bulgaria and Greece). Pensoft & National Museum of Natural History, Sofia, 371–378.
- Principi MM (1960) Contributi alio studio dei Neurotteri italiani. XV. Descrizione di una nuova specie di Raphidia Linn., R. grandii, (Neuroptera-Raphidiidae) e considerazioni generali sulla morfologia degli ultimi uriti dei Neurotteri. Bollettino dell‘Istituto di Entomologia della Università di Bologna 24: 325–337.
- Pröse H (1995) Kommentierte Artenliste der Netzflügler Bayerns (Insecta: Neuropteroidea). Beiträge zur bayerischen Entomofaunistik 1: 151–158.
- Rambur MP (1842) Histoire naturelle des insectes. Névroptères. Roret, Paris, 534 pp.
- Ratzeburg JTC (1844) Die Forst-Insecten, oder Abbildung und Beschreibung der in den Wäldern Preussens und der Nachbarstaaten als schädlich oder nützlich bekannt gewordenen Insekten; in systematischer Folge und mit besonderer Rücksicht auf die Vertilgung der Schädlichen. Die Ader-, Zwei-, Halb-, Netz- u. Geradflügler. Bd. 3. Nicolai, Berlin, 248–254.
- Rausch H, Aspöck H (1991) Phaeostigma (Graecoraphidia) albarda n. sp. – eine neue Kamelhalsfliege von der Peloponnes (Griechenland) (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 43(1/2): 17–24.
- Rausch H, Aspöck H (1992) Zur Kenntnis der Larven, der Biologie und Ökologie und der Verbreitung von drei für die südliche Balkan-Halbinsel endemischen Raphidiiden-Spezies (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 44(1/2): 35–41.
- Rausch H, Aspöck H (1993) Phaeostigma holzingeri n. sp. – eine neue Kamelhalsfliege aus Ipiros (Griechenland) (Neuropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 45(1/2): 19–26.
- Rausch H, Rausch R (2004) 1000 Tage auf entomologischen Forschungsreisen. Unsere gemeinsamen Unternehmungen zur Erforschung der Neuropterida von 1969 bis 2000. In: Aspöck U (Ed.) Entomologie und Parasitologie. Festschrift zum 65. Geburtstag von Horst Aspöck, 640 pp. Denisia 13: 79–105.
- Rausch H, Aspöck H, Aspöck U (2016) Rätselhaftes Massenauftreten einer mediterranen Kamelhalsfliege im Mühlviertel, Oberösterreich (Insecta: Neuropterida: Raphidioptera: Raphidiidae). Linzer Biologische Beitrage 48(1): 523–534.
- Rechinger KH (1950) Grundzüge der Pflanzenverbreitung in der Ägäis I–III. Vegetatio 2(2–3): 55–119. [239–308, 365–386.] https://doi.org/10.1007/BF00151672
- Rechinger KH, Rechinger-Moser F (1951) Phytogeographia Aegaea. Akademie der Wissenschaften Wien, Mathematisch-Naturwissenschaftliche Klasse. Denkschrift 105(2): 3–208.
- Rögl F, Bernor RL, Dermitzakis MD, Müller C, Stancheva M (1991) On the Pontian correlation in the Aegean (Aegina Island). Newsletters on Stratigraphy 24(3): 137–158. https://doi.org/10.1127/nos/24/1991/137
- Röhricht W (1996) Netzflügler und Schnabelfliegen aus Mitteldeutschland. Naturwissenschaftliche Beiträge des Museum Dessau 9: 135–156.
- Röhricht W (2000) Netzflügler und Schnabelfliegen aus Mitteldeutschland. (1. Nachtrag). Naturwissenschaftliche Beiträge des Museums Dessau 12: 82–92.
- Saure C (1996) Die Kamelhalsfliegen (Raphidioptera) der Mark – Beitrag zur Tierwelt von Brandenburg und Berlin. Entomologische Nachrichten und Berichte 40: 75–82.
- Saure C, Gerstberger M (1991) Standardliste und Rote Liste der Neuropteroidea (Netzflügler s.l.) von Berlin. In: Auhagen A, Platen R, Sukopp H (Eds) Rote Listen der gefährdeten Pflanzen und Tiere in Berlin. Landschaftsentwicklung und Umweltforschung S 6: 237–241.
- Schmitz O (1992) Beitrag zur Netzflüglerfauna (Insecta: Neuropteroidea) von Köln und Umgebung. Decheniana Beihefte 31: 165–180.
- Schneider WG (1843) Monographia generis Rhaphidiae Linnaei. Continens et novas de huius generis singulis speciebus institutes observationes, et inegram omnium, quae hucusque inventae sunt, specierum descriptionem. Grass, Barth & Co., Vratislaviae, 96 pp. https://doi.org/10.5962/bhl.title.59533
- Schubert H, Gruppe A (1999) Netzflügler der Kronenregion – Bemerkenswerte Funde und Habitatpräferenzen (Neuropteroidea). Nachrichtenblatt der Bayerischen Entomologen 48: 91–96.
- Schummel TE (1832) Versuch einer genauen Beschreibung der in Schlesien einheimischen Arten der Gattung Raphidia. Linnaeus – E. Pelz, Breslau, 16 pp.
- Scinà D (1818) La Topografia di Palermo e de‘suoi contorni. Dalla Reale Stamperia Palermo, 98 pp.
- Shen R, Aspöck H, Aspöck U, Plant J, Dai Y, Liu XY (2022) Unraveling the evolutionary history of the snakefly family Inocelliidae (Insecta: Raphidioptera) through integrative phylogenetics. Cladistics 38(5): 515–537. https://doi.org/10.1111/cla.12503
- Simaiakis S, Mylonas M (2008) The Scolopendra species (Chilopoda: Scolopendromorpha: Scolopendridae) of Greece (E-Mediterranean): a theoretical approach on the effect of geography and palaeogeography on their distribution. Zootaxa 1792(1): 39–53. https://doi.org/10.11646/zootaxa.1792.1.3
- Stein JPEF (1863) Beitrag zur Neuropteren-Fauna Griechenlands mit Berücksichtigung dalmatinischer Arten. Berliner Entomologische Zeitschrift 7: 411–422. https://doi.org/10.1002/mmnd.47918630310
- Steinmann H (1963) Raphidiopterological studies I. Navasana gen. n. from Hungary, Harraphidia gen. n. from Morocco, new Lesna Navás und Subilla Navás species from Europe. Acta Zoologica Hungarica 9: 183–198.
- Steinmann H (1964) Raphidiopterological studies II. New Raphidia L. and Raphidilla Nav. species from Europe and Asia. Acta Zoologica Hungarica 10: 199–227.
- Stephens JF (1836) Illustrations of British entomology; or, a synopsis of indigenious insects: containing their generic and specific distinctions; with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable (Vol. VI). Baldwin & Cradock, Mandibulata 240 pp.
- Strid A (1996) Phytogeographia Aegaea and the flora Hellenica database. Annalen des Naturhistorischen Museums in Wien. Serie B, Fur Botanik und Zoologie 98(B Suppl.): 279–289.
- Suntrup A (1990) Untersuchungen zur Faunistik und Autökologie von Netzflüglern (Insecta: Neuropteroidea) in Norddeutschland. Diplomarbeit. II. Zoologisches Institut, Georg-August-Universität, Göttingen: 65 pp. [+ 76 pp. (Karten).]
- Sziráki G (1993) Taxonomic status of Raphidia mediterranea H. Aspöck, U. Aspöck et Rausch, 1977 (Raphidioptera: Raphidiidae). Folia Entomologica Hungarica [= Rovartani Közlemények] 54: 147–150.
- Sziráki G (1996) Ecological investigations of the Neuropteroidea of oak forests in Hungary (Insecta: Raphidioptera, Neuroptera). In: Canard M, Aspöck H, Mansell MW (Eds) Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology. (Insecta: Neuroptera, Megaloptera, Raphidioptera) Cairo, Egypt, 2–7 May 1994, 229–232.
- Sziráki G (1999) Neuropteroidea of the Aggtelek National Park. The Fauna of the Aggtelek National Park: 143–149.
- Sziráki G (2010) Néhány kisebb fajszámú rovarrend szigetközi vizsgálata. In: Gubányi A, Mészáros F (Eds) A Szigetköz Állattani Értékei. Magyar Természettudományi Múzeum, Budapest, 49–56.
- Sziráki G (2014) Data to the Raphidioptera fauna of the Balkan Peninsula and Crete. Folia Historico-Naturalia Musei Matraensis 38: 87–90.
- Sziráki G, Popov A (1996) Neuropteroidea of the Bükk National Park. In: Mahunka S (Ed.) The Fauna of the Bükk National Park (Vol. 2). Hungarian Natural History Museum, Budapest, 389–396.
- Sziráki G, Ábrahám L, Szentkirályi F, Papp Z (1992) A check-list of the Hungarian Neuropteroidea (Megaloptera, Raphidioptera, Planipennia). Folia Entomologica Hungarica [= Rovartani Közlemények] 52: 113–118.
- Thierry D, Canard M, Cocquempot C (2021) Présence de Xanthostigma aloysianum (Costa, 1855) dans le Centre-Ouest de la France : Anjou et Touraine (Neuropterida Raphidioptera Raphidiidae). L'Entomologiste 77(3): 153–156.
- Tillier P (2015) Raphidioptera et Neuroptera (Insecta, Neuropterida) du Parc national du Mercantour (France). Zoosystema 37(4): 581–593. https://doi.org/10.5252/z2015n4a4
- Tillier P, Claude J, Danflous S, Decoin R, Touroult J, Vincent A (2022a) Contribution à la connaissance des Raphidioptères de France (Raphidioptera). –. Entomologiste 78(4): 249–268.
- Tillier P, Fleury J, Parmein G (2022b) Découverte dans le Loiret de la deuxième station mondiale d’une espèce énigmatique de raphidie: Venustoraphidia conviventibus Monserrat & Papenberg, 2012 (Raphidioptera, Raphidiidae). Bulletin de la Société Entomologique de France 127(1): 5–10. https://doi.org/10.32475/bsef_2227
- Tröger EJ (1993) Beitrag zur Kenntnis der Netzflügler in Franken (Neuropteroidea). Nachrichtenblatt der Bayerischen Entomologen 42: 33–46.
- Tröger EJ (2002) Netzflügler (Neuropterida) in Franken. Galathea, Nürnberg 13(Supplement): 37–69.
- Tröger EJ (2005a) Netzflügler (Neuropterida) im westlichen Kreta – eine Übersicht. Galathea, Nürnberg, 18. Supplement (7. Arbeitstagung deutsprachiger Neuropterologen, Rödelsee, 4.–6. April 2003): 33–42.
- Tröger EJ (2005b) Über Netzflügler (Neuropterida) in Baden-Württemberg. Galathea 21: 93–96. [Berichte des Kreises Nürnberger Entomologen]
- Vas J, Markó V, Ábrahám L, Mészáros Z (2001) Study of Neuropteroidea (Raphidioptera, Neuroptera) communities by using malaise traps in an untreated orchard and its environment. Acta Phytopathologica et Entomologica Hungarica 36(1–2): 115–122. https://doi.org/10.1556/APhyt.36.2001.1-2.15
- Wachmann E, Saure C (1997) Netzflügler, Schlamm- und Kamelhalsfliegen. Beobachtung – Lebensweise. Naturbuch Verlag, Augsburg, 159 pp.
- Weigelmeier S (2008) Raphidioptera-Larven als Totholzbewohner von Eichen im Spessart. Bachelorarbeit. Department für Biologie, Lehrstuhl für Entwicklungsbiologie der Friedrich-Alexander-Universität Erlangen-Nürnberg, 49 pp.
- Weissmair W, Rausch H, Rausch R (2021) Raphidioptera (Kamelhalsfliegen) in ausgewählten Streuobstkulturen in Oberösterreich. Entomologica Austriaca 28: 43–56.