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The snakeies of the Mediterranean islands: review and biogeographical
analysis (Neuropterida, Raphidioptera)
Horst Aspöck1, Ulrike Aspöck2,3
1 InstituteofSpecicProphylaxisandTropicalMedicine,MedicalParasitology,MedicalUniversityofVienna,Kinderspitalgasse15,1090Vienna,
Austria
2 NaturalHistoryMuseumVienna,DepartmentofEntomology,Burgring7,1010Vienna,Austria
3 DepartmentofEvolutionaryBiology,UniversityofVienna,Djerassiplatz1,1030Vienna,Austria
https://zoobank.org/9E52FBF7-700E-4FC3-A62E-0334CE3DE926
Corresponding author: Ulrike Aspöck (ulrike.aspoeck@nhm-wien.ac.at; ulrike.aspoeck@univie.ac.at)
Academic editor: D. Zimmermann ♦
Received
6 February 2023 ♦
Accepted
17 March 2023 ♦
Published
3 May 2023
Abstract
The Mediterranean region represents one 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), Subillaprincipiae (Sardinia), S.colossea (Rhodes), Raphidia (R.) peter-
ressli (Chios), R. (R.) ariadne (Crete), Fibla (F. ) maclachlani (Corsica, Sardinia, Sicily), F. (Reisserella) pasiphae (Crete).
In the Aegean Sea there is a remarkable dierence between the eastern and the western islands marked by the so-called Rechinger
line based on results of the analysis of the ora. The Raphidioptera conrm this line impressively.
It is rather unlikely that further Raphidioptera species endemic to Mediterranean islands are still to be detected – with one ex-
ception: 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 snakeies 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 (snakeies) is the smallest order of ho-
lometabolous insects. So far ca. 250 described valid species
are known, and although several species are still to be detect-
ed, it is very unlikely that the real number of snakey spe-
cies harboring our planet presently exceeds 300. The order
comprises two families: Raphidiidae (with ca. 210 known
species) and Inocelliidae (with > 40 species). The two fami-
lies have a similar general appearance, but are easily distin-
guished by several striking features. All snakeies are char-
acterized by an elongated prothorax, by translucent wings
Dtsch. Entomol. Z. 70 (1) 2023, 175–218|DOI 10.3897/dez.70.101559
Copyright 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 unre-
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Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
176
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. 2a, b).
Both families are characterized by long developmental
periods. Most species need two years with at least one hi-
bernation of the mature larva (or rarely pupa). The larvae
of the majority of species (many Raphidiidae and proba-
bly all Inocelliidae) (e.g. Figs 2c, 6h) are corticolous, but
larvae of many species of Raphidiidae live in the detritus
around roots of bushes or in crevices of rocks. Larvae
of all snakeies and adults of the Raphidiidae are pred-
ators of various soft-bodied arthropods, while adults of
Inocelliidae feed (at least sometimes) on pollen. All adult
snakeies are active in the day-time; very rarely single
specimens are exceptionally attracted by articial light.
Today, the distribution of both families of the order is
conned to arboreal parts of the Northern Hemisphere
(Fig. 1), which is to be traced back to the fact that a de-
crease of temperature during winter is a precondition for a
normal development (H. Aspöck and U. Aspöck 1991; H.
Aspöck 2002; H. Aspöck et al. 2018, 2019). In the Mesozo-
ic, particularly in the Cretaceous, snakeies had their Gold-
en Age. Probably the planet was inhabited by thousands of
species, and Raphidioptera occurred also in tropical zones
and the Southern Hemisphere. At the end of the Cretaceous,
about 66 million years ago (mya), an asteroid of more than
10 km diameter crashed into the earth causing a worldwide
darkness and a decrease of temperature for several years.
This dramatic change in all terrestrial ecosystems probably
led to the near extinction of the snakeies; however, at least
two lines survived in the Northern Hemisphere representing
the two families of our times. Their general appearance re-
sembles largely that of the Mesozoic Raphidioptera (H. As-
pöck 1998; Lu et al. 2020). The extant Raphidioptera may
be justiably apostrophized as living fossils.
Most snakey species are characterized by small, in
many cases extremely small distribution areas largely
restricted to glacial or postglacial refugial areas. The ca-
pacity to disperse and thus their expansivity are usually
very low; there are only very few species with large dis-
tributions, 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 characteris-
tics are also of striking signicance for an understanding
of the Raphidioptera fauna of the Mediterranean islands.
Monographs and overviews on Raphidioptera:
H. Aspöck et al. (1991), H. Aspöck et al. (2001), H. Aspöck
and U. Aspöck (2009), U. Aspöck and H. Aspöck (2009).
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 tecton-
ic 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 (Hofrichter 2020).
Presently, there are more than 4,300 islands in the Medi-
terranean Sea, most of them small and uninhabited, in many
cases only rocks. However, some islands are large (e.g.
Figure 1. World distribution of Raphidioptera.
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
dez.pensoft.net
177
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). Sev-
eral islands have high elevations (e.g. Sicily >3300 m, Sar-
dinia >1800 m, Corsica >2700 m, Euboea >1700 m, Crete
>2450 m, Rhodes 1250 m, Cyprus >1950 m) with large for-
ests and thus excellent ecological conditions for snakeies.
The genesis of the islands is very dierent. Some of
them represent fragments broken o from the margins of
the continent and drifted into the sea, e.g. Sardinia, Corsi-
ca, and the Baleares broke o from Iberia in the Oligocene,
between 35 and 28 mya. Others represent the exposed el-
evations of otherwise overooded parts of adjacent con-
tinental 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 (Simaiakis and Mylonas 2008).
Sicily is an island composed of three parts of dier-
ent origin. The northeasternmost part comes from the
Apennine Peninsula (which consists of the Apennines on
one hand and the Apulian Plate on the other hand). The
northwestern part of Sicily (including central parts) de-
rives from Iberia, namely from the land broken o 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 sub-
marine separation occurred probably in the Miocene. This
separated part drifted to the west and emerged from the sea
later (Rögl, pers. comm., Badano and Makris 2020).
Rhodes was once a part of Anatolia and became sepa-
rated possibly in the Pliocene due to tectonic events and
subsequent transgressions (Rögl et al. 1991; Simaiakis
and Mylonas 2008).
Most of the islands near the mainland had connections
to the mainland during the glacial periods of the Pleis-
tocene, 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 con-
nected with neighboring islands, but not with the main-
land. Sardinia and Corsica (Corsardinia) were sometimes
connected to each other, but never to the mainland since
their separation from Iberia. Crete was aected by several
transgressions, but at no time was the whole island ood-
ed; 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 im-
portant precondition for the evolution of endemic species
restricted to a certain island. As regards the Raphidiop-
tera of Mediterranean islands, Sardinia, Corsica, Crete,
Karpathos, and Rhodes do harbor endemic snakeies,
and none of these islands had any connection to conti-
nental regions at least throughout the whole Pleistocene.
Euboea also harbors at least one endemic snakey, but
this is due to a refugial center in high mountains (see Dis-
cussion). Interestingly, Cyprus has no endemic snakey.
So far, the order Raphidioptera has not yet been recorded
from the Baleares. However, it is out of the question for
snakeies 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 rst record of a snakey from Mediterranean islands
dates back to Dominicus Scina (1818), who listed “Ra-
phidiaophiopsis” among the insects of Palermo. Raphid-
iaophiopsis does not occur in Sicily, so we do not know
which species he had really found. At that time almost
every snakey was called “Raphidiaophiopsis”. Hagen
(1867) was the rst one who published reliable records
of Raphidioptera from Mediterranean islands. He de-
scribed Raphidiacorsica (today: Xanthostigma corsica)
from Corsica and Raphidiacyprica (today: Phaeostigma
(Crassoraphidia) cyprica) from Cyprus. This is a small
and short paper; nevertheless it is a milestone in raphid-
iopterology (and even in entomology in general) as it
contains descriptions of ve new species solely based
on characters of the male genitalia. More than 20 years
passed until the next record of a snakey on a Mediter-
ranean island was published: Novak (1891) recorded Ra-
phidia anis Schneider (today: Dichrostigma avipes)
from the Adriatic island Hvar. The rst record of a spe-
cies of the family Inocelliidae was published by Albarda
(1891) in his admirable “Révision des Rhaphidides”. He
described Inocellia maclachlani (today: Fibla (F. ) ma-
clachlani) from Corsica. The rst record of a snakey
on Crete was published by Navás (1915a), when he de-
scribed Lesnabiroi (today: Phaeostigma (Aegeoraphid-
ia) biroi), a species endemic to Crete. No further discov-
eries of snakeies on Mediterranean islands were made
during the following decades. In 1960, only three spe-
cies of Raphidiidae and one species of Inocelliidae were
known from a total of four Mediterranean islands: Corsi-
ca, Hvar, Crete, and Cyprus. In the rst half of the 1960s
we began an intensive work on Raphidioptera. A study of
the large collection of unidentied snakeies of the Natu-
ral History Museum Vienna (and of other collections) led
to descriptions of a second snakey species from Crete
– Raphidiaariadne – and of the rst snakey species –
Phaeostigma (Aegeoraphidia) prophetica – from Rhodes
(H. Aspöck and U. Aspöck 1964a, 1965a). Moreover, a
female of an unknown species of Inocelliidae from Crete
was described and gured (but not named), which rep-
resented the rst record of this family on an Aegean is-
land. A few years later a male of this species was found
so that the species could adequately be described (H.
Aspöck and U. Aspöck 1971b): Fibla (Reisserella) pa-
siphae. From 1966 onwards we explored regularly and
systematically the Raphidioptera fauna of the Mediter-
ranean region, and from 1973 onwards particularly also
that of the islands (Naxos, Syros, Chios, Samos, Ikaria,
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Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
178
Euboea, Thasos, Samothraki, Skopelos, Skyros, Levkas,
Rhodes, Sicily, Crete, Karpathos, Lesbos, Limnos, Cy-
prus, Sardinia, Corsica; for details see Gusenleitner 2004;
H. Rausch and R. Rausch 2004; H. Aspöck 2012). These
eld trips led to discoveries of new species or subspe-
cies: Phaeostigma (Ph.) euboica, Ph. (Graecoraphidia)
divinaretsinata,Phaeostigma (Magnoraphidia) ammi,
Ph. (Pontoraphidia) setulosa aegea, Ph. (Aegeoraphid-
ia) karpathana, Ph. (Superboraphidia) minois, Subilla
prinicpiae, S.colossea, Raphidia (R.) mediterranea, Ra-
phidia (R.) peterressli, Fibla (Reisserella) pasiphae (H.
Aspöck and U. Aspöck 1971b, 1973, 1976; H. Aspöck
et al. 1977, 1979, 1991; U. Aspöck and H. Aspöck 1989,
1990; Pantaleoni et al. 2005).
In addition, several species known from mainland re-
gions were found on Mediterranean islands for the rst
time: Venustoraphidia nigricollis, Parvoraphidia mi-
crostigma, Ornatorahpidiaavilabris, O.christianodag-
mara, Phaeostigma (Magnoraphidia) major, Ph. (M.)
wewalkai, Ph. (Aegeoraphidia) raddai, Subilla connis,
S.artemis, Ulrikesyriaca, R. (R.) beieri, R. (R.) mysia,
R. (R.) ambigua, Parainocelliaressli.
During the past fteen 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 snakey fauna of the
Mediterranean islands.
Materials and methods
This comprehensive review and overview is based on
several thousand specimens of Raphidioptera from Med-
iterranean islands identied 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 (Aspöck et al. 1991), and except original
descriptions these papers are not dealt with here again.
However, all publications from 1991 onwards dealing
with species on Mediterranean islands are considered here
and are listed in the synonymy lists of each taxon. Large
parts of specimens of Raphidioptera of the Mediterranean
regions including the islands are deposited in the follow-
ing collections: Naturhistorisches Museum Wien, Biol-
ogiezentrum Linz, private collection Horst and Ulrike
Aspöck (Vienna), private collection Hubert and Renate
Rausch (Scheibbs, Lower Austria). Further specimens of
Raphidioptera from Mediterranean islands are deposited
in most European museums; we have seen and identied
by far the largest part of these (see Aspöck et al. 1991).
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 reect National
Geographic’s current map policy. Sources: National Geo-
graphic, ESRI, DeLorme, HERE, UNEP-WCMC, USGS,
NASA, ESA, METI, NRCAN, GEBCO, NOAA, incre-
ment P Corp.
The systematic order follows H. Aspöck and U. As-
pöck (2014). For details see also H. Aspöck et al. (1991,
2001), U. Aspöck and H. Aspöck (1989, 1990), Panta-
leoni et al. (2005).
Abbreviations used in the synonymy lists: anat = anat-
omy; anncat = annotated catalogue; asl. = above sea lev-
el; bibliogr = bibliography; biogeogr = biogeography;
biol = biology; cat = catalogue; charact = character-
ization; chorol = chorology; com = comment; comp-
morph = comparative morphology; descr = description;
distr = distribution; distrmap = distribution map;
ecol = ecology; ethol: cop = ethology, copulation; ety-
mol = etymology; FD! = false determination; gs = gen-
ital segments; ill = illustration; imag = imago (adult);
la = larva; list = listed; molecsyst = molecular sys-
tematics; mon = monograph; nom = nomenclature;
odescr = original description; overv = overview; pa-
ras = parasites, parasitoids; phyl = phylogeny; phylo-
geogr = phylogeography; phyltree = phylogenetic tree;
pu = pupa; rec = record; s.l. = sensu lato; s. str. = sen-
su 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 record-
ed from Mediterranean islands are treated with respect to
their synonymies, taxonomy, systematics, biology, ecolo-
gy, distribution, and biogeography (Tables 1, 2).
Family Raphidiidae Latreille, 1810
Family Raphidiidae Latreille, 1810: H. Aspöck et al. 1991 (mon); H.
Aspöck and U. Aspöck 1991 (overv); H. Aspöck et al. 2001 (ann-
cat); H. Aspöck 2002 (biol); H. Aspöck and U. Aspöck 2009 (overv);
U. Aspöck and H. Aspöck 2009 (overv); Haring et al. 2011 (phylog-
eny); Oswald and Machado 2018 (overv); Gruppe et al. 2023 (biol).
Venustoraphidia H. Aspöck & U. Aspöck, 1968
Venustoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidianigricollis Albarda, 1891]: H. Aspöck et al. 1991 (mon);
H. Aspöck et al. 1989 (biogeogr, distr, ill: distrmap); Oswald and
Penny 1991 (cat, nom); H. Aspöck et al. 2001 (anncat); H. Aspöck
2002 (biol, paras); Haring et al. 2011 (phyl); H. Aspöck 2012 (cat);
H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat); Monserrat
and Papenberg 2012, 2015 (mon).
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
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179
Table 1. Records of Raphidioptera on Mediterranean islands I.
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 xxxxx
X.corsica xxxxx
X.aloysiana x
Parvoraphidia x
P.microstigma x
Ornatoraphidia x x
O.avilabris x
O.christianodagmara x
Phaeostigma xxx x x xxxxxxxx
Phaeostigmas.str. x
Ph. (Ph.) euboica x
Graecoraphidia x
Ph. (G.) divinaretsinata x
Crassoraphidia x
Ph. (C.) cyprica x
Magnoraphidia x x x x
Ph. (M.) major x x
Ph. (M.) ammi x x
Ph. (M.) wewalkai x
Pontoraphidia x
Ph. (P. ) setulosaaegea x
Aegeoraphidia xxxxxxx
Ph. (Ae.) raddai xxxx
Ph. (Ae.) prophetica x
Ph. (Ae.) karpathana x
Ph. (Ae.) biroi x
Superboraphidia x
Ph. (S.) minois x
Table 2. Records of Raphidioptera on Mediterranean islands II.
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.connis x
S.artemis x
S.principiae x
S.colossea x
Ulrike x
U.syriaca x
Raphidia xx xxxxxxxxxxxxxx
Raphidias.str. xx xxxxxxxxxxxxxx
R.(R.)mediterranea xxxxxxx 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 xxx
D.avipes xxx
Inocelliidae
Fibla x x x x
Fiblas.str. xxx
F.(F.)maclachlani xxx
Reisserella x
F.(R.)pasiphae x
Parainocellia x
P.ressli x
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Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
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Taxonomy and systematics. H. Aspöck et al. (1991), Har-
ing et al. (2011), Monserrat and Papenberg (2012, 2015).
The genus is markedly dierentiated by morphological
characters of adults and larvae and also by molecular
phylogeny. It has emerged as the sister group of (Puncha
+ Calabroraphidia) + Xanthostigma. Venustoraphidia
comprises three geographically vicariant species.
Biology and ecology. Larvae corticolous on a great
variety of deciduous and coniferous trees. Development
usually two years. Last hibernation stage: full-grown lar-
va. Adults: (EIV)V–VI(BVII).
Distribution. Apennine Peninsula, Balkan Peninsu-
la, northern part of Iberian Peninsula, Central and East-
ern Europe.
Venustoraphidia nigricollis (Albarda, 1891)
Raphidia nigricollis Albarda, 1891 (odescr): H. Aspöck et al. 1991
(mon).Venustoraphidia nigricollis (Albarda 1891): H. Aspöck et
al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); Pantaleoni
1990b (ecol); Pantaleoni 1990d [1993] (rec); Kielhorn 1991 (ecol,
rec); Saure and Gerstberger 1991 (ecol, rec); Devetak 1992a (rec);
Devetak 1995 (rec); Pröse 1995 (distr, rec); H. Aspöck and Hölzel
1996 (distr); Saure 1996 (biogeogr, ecol, rec); Sziráki 1996 (ecol,
rec); Achtelig 1997 (ecol, rec); Gruppe 1997 (ecol, rec); Schubert
and Gruppe 1999 (ecol, rec); Sziráki 1999 (rec); H. Aspöck et al.
2001 (anncat); Gruppe and Schubert 2001 (ecol, rec); Tröger 2002
(rec); Popov 2004 (chorol); Gruppe 2006a (ecol); Gruppe 2006b
(ecol, rec); H. Aspöck and U. Aspöck 2007 (biogeogr, distr); Panta-
leoni 2007 (biol); Gruppe 2008 (ecol, rec); Klokočovnik et al. 2010
(rec); Letardi et al. 2010 (rec); H. Aspöck and U. Aspöck 2013 (cat,
etymol); H. Aspöck and U. Aspöck 2014 (cat); Weissmair et al. 2021
(biol, ecol); H. Aspöck and U. Aspöck 2021 (overv, ill: la, pu, ♂, ♀
imag), 2022a (overv, ill: la, pu, ♂, ♀ imag), 2022b (overv, ill: la, pu,
♂, ♀ imag); Tillier et al. 2022b (rec).
Taxonomy. H. Aspöck et al. (1991). Adults can easily be
identied by the entirely black pronotum, usually already
by the small size (Fig. 2a, b). Larvae are readily recogniz-
able by the characteristic pattern of the dorsal pigmenta-
tion (Fig. 2c).
Biology and ecology. Larvae exclusively corticolous
on Quercus, Malus, Pyrus, Acer and many other decid-
uous 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. 7b). Thasos
(Ipsarion, ca. 1000 m, on Pinus). Syntopic Raphidioptera
on Thasos: Phaeostigma (Pontoraphidia) setulosaaegea,
Raphidia (R.) beieri.
Continental distribution. Balkan Peninsula as far south
as to the gulf of Korinthos, Eastern Europe, Apennine Pen-
insula (Northern Italy, Calabria), Central Europe as far north
as to Northern Germany, Eastern and Southern France.
Biogeography. Polycentric Adriato-Balkanoponto-
mediterranean faunal element with high expansivity.
Xanthostigma Navás, 1909
Xanthostigma Navás, 1909 (described as a section of Raphidia L.) [type
species by absolute tautonymy: Raphidia xanthostigma Schummel,
1832]: H. Aspöck et al. 1991 (mon); Oswald and Penny 1991 (cat,
nom); H. Aspöck et al. 1999 (ecol, chorol); H. Aspöck 2002 (biol,
paras); H. Aspöck et al. 2001 (anncat); Haring et al. 2011 (phyl); U.
Aspöck et al. 2012 (g: phyltree, list); H. Aspöck and U. Aspöck 2013
(cat, etymol, phyl), 2014 (cat); Monserrat and Papenberg 2015 (overv).
Raphidilla Navás, 1915b [type species by original designation: Raphid-
ia xanthostigma Schummel, 1832]: H. Aspöck et al. 1991 (mon);
Oswald and Penny 1991 (cat, nom).
Rhaphidilla Navás, [1919] 1918b [unjustied emendation of Raphidilla
Navás, 1915]: H. Aspöck et al. 1991 (mon); Oswald and Penny 1991
(cat, nom); H. Aspöck and U. Aspöck 2014 (cat).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). Xanthostigma is a morphologically
clearly dierentiated genus of the Puncha clade and ac-
cording to molecular phylogeny results is the sister taxon
of Puncha + Calabroraphidia or of Italoraphidia. The
genus comprises ve known species with very dierent
distribution patterns including Eurosibirian, Mongolian,
Pontocaspian, and Mediterranean faunal elements.
Biology and ecology. Larvae of X. corsica proba-
bly 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 – usu-
ally two years. Last hibernation stage: full-grown larva.
Adults: IV–VII.
Distribution. Almost whole of Europe from southern-
most parts (X. corsica) to northern parts of Scandinavia
over the North of Asia as far as to Far East (X.xanthostig-
ma). Caucasus region, Mongolia, northern China.
Xanthostigma corsica (Hagen, 1867)
Raphidiacorsica Hagen, 1867 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiainsularis Albarda, 1891 (odescr): H. Aspöck et al. 1991 (mon).
Punchaitalica Navás, 1927b (odescr): H. Aspöck et al. 1991 (mon).
Xanthostigma corsica (Hagen, 1867): H. Aspöck et al. 1991 (mon);
Pantaleoni 1990c (com, rec); H. Aspöck and Hölzel 1996 (distr);
Letardi and Pantaleoni 1996 (rec); H. Aspöck et al. 2001 (anncat);
Letardi and Maltze 2001 (rec); Nicoli Aldini and Baviera 2001
(rec); H. Aspöck and U. Aspöck 2007 (biogeogr, distr); Letardi et al.
2008 (list, rec); Badano and Letardi 2010 (rec); Haring et al. 2011
(phyl, phyltree); Nicoli Aldini et al. 2012 (rec, distr); H. Aspöck and
U. Aspöck 2013 (cat, etymol; ill: imag, la), 2014 (cat).
Xanthostigmacorsicum (Hagen): Monserrat and Papenberg 2015 (mon,
rec, synlist); Tillier et al. 2022a (rec).
Taxonomy. H. Aspöck et al. (1991), Monserrat and Pa-
penberg (2015). X.corsica can always easily be identied
by morphological characters (e.g. three cells between R
and Rs in forewing) (Fig. 2d).
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Biology and ecology. Larvae mainly terricolous, how-
ever, several records also under bark of Pinus and of
Quercus. Many records of adults mainly on low vegeta-
tion and on bushes (particularly Genista) in a great variety
of dierent 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. 8a). X.cor-
sica is a frequent snakey in various biotopes in altitudes
from 0 to 1500 m asl on Sicily, Sardinia, Corsica, Giglio
and Elba. Probably it occurs also on several smaller Med-
iterranean islands west of the mainland of Italy. Syntopic
Raphidioptera: Xanthostigmaaloysiana (Sardinia), Sub-
illa connis (Sicily), Fibla maclachlani (Sicily, Sardin-
ia, Corsica).
Continental distribution. The species occurs on the
Apennine Peninsula from the south of Calabria to Tus-
cany 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 inva-
sion and might be dierentiated 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 from one island to another.
Biogeography. Polycentric Tyrrhenic-Adriato-Atlan-
tomediterranean faunal element.
Xanthostigma aloysiana (Costa, 1855)
Raphidiaaloysiana Costa, 1855 (odescr): H. Aspöck et al. 1991 (mon).
Raphidillapuella Navás, 1915c (odescr): H. Aspöck et al. 1991 (mon);
H. Aspöck and U. Aspöck 2014 (cat).
Raphidillasoror Navás, 1915c (odescr): H. Aspöck et al. 1991 (mon);
H. Aspöck and U. Aspöck 2014 (cat).
Xanthostigmaaloysiana (Costa): H. Aspöck et al. 1991 (mon); H. As-
pöck and Hölzel 1996 (distr); Letardi and Pantaleoni 1996 (rec);
Monserrat 1996 (distr); H. Aspöck et al. 2001 (anncat); U. Aspöck
and H. Aspöck 2005b (biogeogr); Pantaleoni 2005 (rec); H. Aspöck
and U. Aspöck 2007 (biogeogr, distr); Badano 2008 (distr, rec);
Badano and Letardi 2010 (rec); Monserrat and Papenberg 2010
(chorol); Haring et al. 2011 (phyl, phyltree); H. Aspöck and U. As-
pöck 2013 (cat, etymol), 2014 (cat).
Xanthostigmaaloysianum (Costa): Letardi and Maltze 2008; Letardi
2015 (rec); Monserrat and Papenberg 2015 (mon, rec, synlist); Thi-
erry et al. 2021 (rec, distr).
Taxonomy. H. Aspöck et al. 1991 (mon). X. aloysiana
resembles supercially X.corsica, but can easily be dif-
ferentiated by eidonomic characters (e.g. only two cells
between R and Rs in the forewing).
Biology and ecology. Poorly known. Larvae probably
terricolous. Adults in light pine forests or light mixed for-
ests with rich bush vegetation and in macchias of dier-
ent structure in altitudes of 600 to 1200 m, usually found
in single specimens only. Adults: V–VII.
Records on Mediterranean islands (Fig. 8a). So far
only in Central Sardinia (Gennargentu). These ndings,
so far not properly published, are from altitudes slightly
below 1000 m. Syntopic Raphidioptera in Sardinia: Xan-
thostigmacorsica, Fiblamaclachlani.
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 H. Aspöck & U. Aspöck, 1968
Parvoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as a
subgenus of Raphidia L.) [type species by original designation: Ra-
phidamicrostigma Stein, 1863]: H. Aspöck et al. 1991 (mon); Os-
wald and Penny 1991 (cat, nom); H. Aspöck et al. 2001 (anncat); H.
Aspöck 2002 (biol); U. Aspöck and H. Aspöck 2005b (com); Haring
et al. 2011 (phyl); H. Aspöck 2012 (cat); U. Aspöck et al. 2012 (g:
phyltree; list); H. Aspöck and U. Aspöck 2013 (cat, etymol, phyl);
H. Aspöck and U. Aspöck 2014 (cat).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). Parvoraphidia is markedly dieren-
tiated by morphological characters of adults (small size:
5–8,5 mm; short and very dark pterostigma) (Fig. 2f) and
larvae and also by molecular phylogeny. It has emerged
as the sister group of Ornatoraphidia + Turcoraphidia,
the genus comprises three closely related species (includ-
ing one subspecies), which can be reliably dierentiated
only by characters of the male genitalia.
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 conned to the south-
ern and southwestern parts of the Balkan Peninsula.
Biogeography. The three species represent Balkano-
pontomediterranean faunal elements with low expansivity.
Parvoraphidia microstigma (Stein, 1863)
Raphidia microstigma Stein, 1863 (odescr): H. Aspöck et al. 1991
(mon); H. Aspöck and U. Aspöck 2014 (cat).
Raphidia (Parvoraphidia) microstigma Stein: H. Aspöck et al.
1991 (mon).
Parvoraphidiamicrostigma (Stein, 1863): H. Aspöck et al. 1991 (mon);
H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat);
Haring et al. 2011 (phyl, phyltree); H. Aspöck and U. Aspöck 2013
(cat, etymol; ill: imag), 2014 (cat); Sziráki 2014 (rec).
Taxonomy. H. Aspöck et al. (1991). P.microstigma can
be morphologically dierentiated only by characters
of the male genitalia, however, the three species (and
one subspecies) of the genus show perfect geographi-
cal vicariance.
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Biology and ecology. Larvae probably exclusively ter-
ricolous. Numerous records of adults on low vegetation,
bushes and trees in various habitats: light oak and pine
forests, on clearings within r tree forests, on slopes with
Genista, on hedgerows at elds in altitudes of 650–1200 m.
Development possibly only one year. Last hibernation
stage: probably full-grown larva. Adults: V–VII.
Records on Mediterranean islands (Fig. 7b). Records
so far only on Levkas. It is, however, likely that the species
will also be found on other Ionian islands. Syntopic Ra-
phidioptera on Levkas: Ornatoraphidiaavilabris, Phae-
ostigma (Magnoraphidia) major, Subillaartemis.
Continental distribution. Greece north of the gulf of
Korinthos, Albania, North Macedonia.
Biogeography. Stationary monocentric Balkanopon-
tomediterranean faunal element.
Ornatoraphidia H. Aspöck & U. Aspöck, 1968
Ornatoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidiaetrusca Albarda, 1891]: H. Aspöck et al. 1991 (mon); Os-
wald and Penny 1991 (cat, nom); H. Aspöck et al. 2001 (anncat);
H. Aspöck 2002 (biol, paras); Haring et al. 2011 (phyl); H. Aspöck
2012 (cat); U. Aspöck et al. 2012 (g: phyltree); H. Aspöck and U.
Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). Ornatoraphidia is markedly dier-
entiated by morphological characters of adults and larvae.
Based upon molecular phylogeny the sister taxon is Turc-
oraphidia. The genus comprises two species, which show
very dierent patterns of distribution. Both species are
known from Mediterranean islands.
Biology and ecology. Larvae living in upper layers of
soil around roots of bushes. Development two years, see
under O.avilabris. Last hibernation stage: pupa or full-
grown larva. Adults: IV–VI.
Distribution. Balkan Peninsula, Apennine Peninsula,
southern parts of Central Europe.
Ornatoraphidia avilabris (Costa, 1855)
Rhaphidia ophiopsis var. avilabris Costa, 1855 (odescr): Pantaleoni
1999 (nom: earlier synonym of Raphidia etrusca), H. Aspöck and
U. Aspöck 2013 (cat, etymol).
Raphidiaetrusca Albarda, 1891 (odescr): H. Aspöck et al. 1991 (mon);
Sziráki et al. 1992 (rec).
Ornatoraphidiaetrusca (Albarda): Pantaleoni 1990a (com, rec); Panta-
leoni 1990d [1993] (rec); H. Aspöck et al. 1991 (mon); Rausch and
H. Aspöck 1991 (com, distr); Popov 1993 (rec); Letardi 1993 [1994]
(rec); Pantaleoni et al. 1994 (distr, rec); U. Aspöck et al. 1995 (ethol:
cop); H. Aspöck and Hölzel 1996 (distr); Letardi and Pantaleoni
1996 (rec); Popov 1997 (rec); Wachmann and Saure 1997 (charact,
g: imag); Popov 2000a (rec), 2000b (rec), 2001 (rec); Pantaleoni
2005 (nom).
Ornatoraphidiaavilabris (Costa): H. Aspöck et al. 1991 (mon); Szirá-
ki 1999 (rec); H. Aspöck et al. 2001 (anncat); Letardi and Migliaccio
2002 (list); U. Aspöck and H. Aspöck 2005b (ill: ♂ gs); U. Aspöck
and H. Aspöck 2005b (biogeogr); Pantaleoni 2005 (nom, rec); An-
derle and U. Aspöck 2007 (rec); H. Aspöck and U. Aspöck 2007
(biogeogr, distr); Badano 2008 (distr, rec); H. Aspöck and U. Aspöck
2009 (ill: ♂ pu); Aistleitner and Gruppe 2009 (rec); Badano and Le-
tardi 2010 (rec); Klokočovnik et al. 2010 (rec); Letardi et al. 2010
(rec); Haring et al. 2011 (phyl, phyltree); H. Aspöck and U. Aspöck
2013 (cat, etymol; ill: imag, la, ♂ imag, pu); H. Aspöck and U. As-
pöck 2014 (cat); Sziráki 2014 (rec); Letardi 2015 (rec); Tillier 2015
(rec); Devetak et al. 2015 (rec); Devetak and Rausch 2016 (rec); Hi-
ermann et al. 2018 (rec); H. Aspöck et al. 2021 (phylogeogr); Tillier
et al. 2022a (rec) ; Gruppe et al. 2023 (biol).
Raphidialuigionii Navás, 1927a (odescr): H. Aspöck et al. 1991 (mon);
H. Aspöck and U. Aspöck 2014 (cat).
Raphidia regisborisi Navás, 1929 (odescr): H. Aspöck et al. 1991
(mon); H. Aspöck and U. Aspöck 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). O. avilabris can
easily be dierentiated from the second species of the ge-
nus, O.christianodagmara, by morphological characters
(brown pterostigma with yellowish zone along the costa)
(Fig. 2g, h).
Biology and ecology. Larvae (Fig. 3a) soil-dwelling.
Development variable: Pupation may take place already
in the second year (i.e. after one hibernation of the larva)
and the last hibernation stage may be the pupa or the pre-
pupa. At least, under experimental conditions with more
or less natural temperatures, sometimes full-grown larvae
may pupate in winter, sometimes full-grown larvae over-
winter and pupate in spring. At any rate, adults always
appear in spring. In the eld, imagines can be found –
depending on the altitude – from IV to VI/VII. The spe-
cies is thermophilic, but euryoecious and occurs in a great
variety of habitats, preferably with light bush vegetation,
particularly also in light forests of various kinds and also
in macchia habitats in altitudes of 100 to 2200 m. O.a-
vilabris has repeatedly been found above the timberline.
Records on Mediterranean islands (Fig. 8b). O.a-
vilabris occurs on the island of Levkas (records at about
100 m asl); it will, however, probably be found also on
other Ionian islands and possibly also on islands in the
Adriatic Sea. Syntopic Raphidioptera on Levkas: Parv-
oraphidia microstigma, Phaeostigma (Magnoraphidia)
major, Subillaartemis.
Continental distribution. The distribution of O. a-
vilabris 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. avilabris represents a polycen-
tric Balkanopontomediterranean-Adriatomediterranean
element with moderate expansivity. The populations of
the southern Balkan Peninsula show considerable genetic
(but not morphological) dierences to populations from
the Apennine Peninsula and from Central Europe.
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Ornatoraphidia christianodagmara (H. Aspöck & U.
Aspöck, 1970)
Raphidia (Ornatoraphidia) christianodagmara H. Aspöck & U.
Aspöck, 1970 (odescr, ecol); H. Aspöck et al. 1991 (mon).
Ornatoraphidiachristianodagmara (H. Aspöck & U. Aspöck): H. As-
pöck et al. 1991 (mon); Popov 1992 (biogeogr); Rausch and H. As-
pöck 1992 (biol, distr, ecol, rec, tax, ill: la, map); H. Aspöck and
Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat); H. Aspöck 2012
(cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat);
Sziráki 2014 (rec).
Taxonomy. H. Aspöck et al. (1991), Rausch and Aspöck
(1992). The species can be easily dierentiated from
O.avilabris eidonomically by the yellow pterostigma.
Biology and ecology. Larvae are soil-dwelling (al-
though this needs conrmation). Development two years.
Pupation in late autumn or even in winter, in one case
pupation took place in March. Adults: V–VI. Findings of
adults in light Castaneasativa forests (Euboea) and in light
r tree (Abiescephalonica) forests (Parnis mountains).
Records on Mediterranean islands (Fig. 9a). Eu-
boea (Ochi mountains, 1100 m). Syntopic Raphidiop-
tera species on Euboea: Phaeostigma (Ph.) euboica,
Phaeostigma (Magnoraphidia) wewalkai, and Raphidia
(R.) mediterranea.
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 Balkanopon-
tomediterranean faunal element.
Phaeostigma Navás, 1909, s.l.
Phaeostigma Navás, 1909 (described as a section of Raphidia L.) [type
species by subsequent designation: Raphidia notata Fabricius,
1781]: U. Aspöck and H. Apöck 1989 (syst); U. Aspöck and H. As-
pöck 1990 (syst); H. Aspöck et al. 1991 (mon); Oswald and Penny
1991 (cat, nom); Wachmann and Saure 1997 (charact); H. Aspöck
et al. 2001 (anncat); H. Aspöck 2002 (biol, paras); U. Aspöck 2002
(compmorph); H. Aspöck and U. Aspöck 2007 (ill: map); Monserrat
and Papenberg 2010 (bibliogr, biogeogr, biol, chorol, descr, distr,
synlist, tax); Haring et al. 2011 (g:distrmap; phyl, phyltree); U.
Aspöck et al. 2012 (g: phyltree; list); H. Aspöck and U. Aspöck
2013 (cat, etymol, phyl), 2014 (cat); Monserrat and Papenberg
2015 (mon).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). With 41 species and three additional
subspecies, Phaeostigma is (after Mongoloraphidia) the
second largest genus of the family (and of the order). It
comprises nine subgenera, seven of these with altogether
12 species are also represented on Mediterranean islands.
Phaeostigma is morphologically clearly dierentiated.
Based on molecular phylogeny it is the sister group of the
rest of the Phaeostigma clade, i.e. (Subilla + Ulrike) +
[(Tjederiraphidia + Dichostigma) + Raphidia] (Haring et
al. 2011). The phylogeny of the subgenera of Phaeostig-
ma has not yet been studied; so far only considerations
based upon morphological criteria are available (H. As-
pöck et al. 1991).
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, Leb-
anon, Syria, northern Iraq, northern Iran. Several Med-
iterranean islands harbor at least one species: Levkas,
Kefalonia, Thasos, Skopelos, Euboea, Crete, Karpathos,
Ikaria, Samos, Lesbos, Chios, Rhodes, Cyprus.
Subgenus Phaeostigma Navás, 1909, s.str.
Phaeostigma Navás, 1909 (described as a section of Raphidia L.) [type
species by subsequent designation: Raphidia notata Fabricius,
1781]: H. Aspöck et al. 1991 (mon); H. Aspöck et al. 2001 (anncat);
H. Aspöck 2002 (biol); U. Aspöck 2002 (compmorph); H. Aspöck
and U. Aspöck 2013 (cat, etymol), 2014 (cat); Monserrat and Papen-
berg 2015 (synlist).
Erma Navás, 1918a (odescr) [type species by subsequent designation
and monotypy: Ermaabdita Navás, 1918a]: H. Aspöck et al. 1991
(mon); Oswald and Penny 1991 (cat, nom); H. Aspöck and U. As-
pöck 2014 (cat).
Navasana Steinmann, 1963 (odescr) [type species by original desig-
nation: Navasanaperumbrata Steinmann, 1963]: H. Aspöck et al.
1991 (mon); Oswald and Penny 1991 (cat, nom); H. Aspöck and U.
Aspöck 2014 (cat).
Note. The subgenus Phaeostigma comprises six known
species, one of these occurs on the island of Euboea.
Phaeostigma (Phaeostigma) euboica (H. Aspöck & U.
Aspöck, 1976)
Raphidia (Phaeostigma) euboica H. Aspöck & U. Aspöck, 1976 (ode-
scr): H. Aspöck et al. 1991 (mon).
Phaeostigma (Phaeostigma) euboica (H. Aspöck & U. Aspöck, 1976):
H. Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon);
Popov 1992 (biogeogr); H. Aspöck and Hölzel 1996 (distr); H. As-
pöck et al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck and U.
Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). The species belongs
to the Ph.notata complex; it can easily be identied by
the characters of the genital segments.
Biology and ecology. Larvae most probably exclu-
sively corticolous, preferably on conifers (Abies, Pinus).
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Figure 2. a. Venustoraphidianigricollis, ♂. Lower Austria, Eichkogel near Mödling. Foto: H. Bruckner; b. Venustoraphidianigricollis,
♀. Austria, Vienna. Foto: H. Bruckner; c. Venustoraphidianigricollis, larva. Austria, Vienna. Foto: H. Bruckner; d. Xanthostigmacor-
sica, ♀. Italy, Calabria, Aspromonte, Montalto. Foto: P. Sehnal; e. Xanthostigmacorsica, larva. Itlay, Calabria, Aspromonte, Montalto.
Foto: F. Anderle, now Denner; f. Parvoraphidiamicrostigma, ♀. Greece, Phokis, Lidorikion mts. Foto: P. Sehnal; g.Ornatoraphidiaa-
vilabris, ♀. Greece, Phokis, Lidorikion mts. Foto: P. Sehnal; h. Ornatoraphidiaavilabris, ♂. Greece, Viotia, Parnassos. Foto: P. Sehnal.
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
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Figure 3. a. Ornatoraphidiaavilabris, larva. Italy, Calabria, Sila Grande. Foto: F. Anderle, now Denner; b. Phaeostigma (Grae-
coraphidia) 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. Phaeostig-
ma (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) ammi, larva. Greece, Pilion. Foto: H. Bruckner.
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Figure 4. a. Phaeostigma (Magnoraphidia) wewalkai, ♂. Greece, Korinthia, Oros Onia. Foto: F. Anderle, now Denner; b. Phae-
ostigma (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 (Superbo-
raphidia) minois, ♂. Greece, Crete, Levka Ori, W Omalos. Foto: H. Aspöck; g. Subillaconnis, ♂. Lower Austria, Klosterneuburg.
Foto: H. Bruckner; h. Subillaprincipiae, ♂. Italy, Sardinia, Gennargentu, Fonni. Watercolour by Wilhelm Zelenka (1936–2011),
Vienna. In the possession of H. & U. Aspöck.
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Figure 5. a. Ulrikesyriaca, ♂. 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. Raphidiamediterranea, larva. Upper Austria,
Pelmberg near Helmonnsödt. Foto: H. Bruckner; e. Raphidiaariadne, ♂. Greece, Crete, near Males. Foto: H. Bruckner; f. Raphidia
ariadne, ♀. Greece, Crete, Anogia. Foto: H. Bruckner; g. Raphidiaariadne, larva. Greece, Crete, near Males. Foto: H. Bruckner;
h. Dichrostigmaavipes, ♂. Lower Austria, Eichkogel near Mödling. Foto: F. Anderle, now Denner.
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Figure 6. a. Dichrostigmaavipes, ♀. Lower Austria, Dürnstein. Foto: F. Anderle, now Denner; b. Dichrostigmaavipes, 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 (Re-
isserella) pasiphae, ♀. Crete, Triphti Forest. Foto: H. Paulus; h. Fibla (Reisserella) pasiphae, larva. Greece, Crete, Anogia > Axos.
Foto: F. Anderle, now Denner.
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Development two to three years. Last hibernation stage:
full-grown larva. Adults: V–VI in light coniferous forests.
Records on Mediterranean islands (Fig. 7b). Central
and southern parts of Euboea in altitudes of 700–1100 m
only. Syntopic Raphidioptera: Ornatoraphidiachristian-
odagmara, Phaeostigma (Graecoraphidia) divina retsi-
nata, Ph. (Magnoraphidia) ammi, Ph. (M.) wewalkai,
Raphidia (R.) mediterranea.
Biogeography. Ph.euboica is probably an endemism
of Euboea. It represents a monocentric, extremely sta-
tionary Balkanopontomediterranean faunal element.
Subgenus Graecoraphidia H. Aspöck & U. Aspöck, 1968
Graecoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidiadivina H. Aspöck & U. Aspöck, 1964a]: H. Aspöck et al.
1989 (biogeogr, distr; ill: distrmap); H. Aspöck et al. 1991 (mon);
Oswald and Penny 1991 (cat, nom); H. Aspöck et al. 2001 (anncat);
H. Aspöck 2002 (biol); H. Aspöck 2012 (cat); H. Aspöck and U.
Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Graecoraphidia is a
well dierentiated subgenus with three species, of which
one – Ph. (G.) divina – comprises three subspecies, one
of which occurs on Euboea (Fig. 3b).
Biology. Larvae (of all species and subspecies?) (Fig. 3c)
living in the soil, but can also be corticolous. Development
two to three years. Last hibernating stage: full-grown larva.
Adults: (IV)V–VI(VII) in various habitats with deciduous
trees and/or conifers in altitudes of 500–1200 m.
Distribution. The distribution of Graecoraphidia is
restricted to a small part of southern Greece.
Phaeostigma (Graecoraphidia) divina H. Aspöck & U.
Aspöck, 1964a
Raphidiadivina H. Aspöck & U. Aspöck, 1964a (odescr): H. Aspöck
et al. 1991 (mon); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck
2014 (cat).
Phaeostigma (Graecoraphidia) divina (H. Aspöck & U. Aspöck): H.
Aspöck et al. 1991 (mon); Rausch and Aspöck 1991 (map); Popov
1992 (biogeogr); H. Aspöck et al. 2001 (anncat); H. Aspöck 2012
(cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Note. Within this species three subspecies are dieren-
tiated, which occur in perfect geographic vicariance in
small parts of southern Greece. One of these occurs on
the island of Euboea.
Phaeostigma (Graeocoraphidia) divina retsinata
(H. Aspöck & U. Aspöck, 1973)
Raphidia (Graecoraphidia) divina retsinata H. Aspöck & U. Aspöck,
1973 (odescr): H. Aspöck et al. 1991 (mon).
Phaeostigma (Graecoraphidia) divina retsinata (H. Aspöck & U. As-
pöck): H. Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991
(mon); Rausch and H. Aspöck 1991 (ill: distrmap); H. Aspöck and
Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat); H. Aspöck 2012
(cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Ph. (G.) divinaretsi-
nata can be dierentiated from the other two subspecies
by characters of the male genitalia, but also by eidonom-
ic characters, such as the small size. The complicated
taxonomic and systematic situation of the subspecies of
Ph. (G.) divina needs an extensive phylogenomic study.
Biology and ecology. Larvae most probably soil-dwell-
ing. Development under experimental conditions three
years. Last hibernating stage: full-grown larva. Adults:
V–VI. Most specimens were collected from r trees in
altitudes of 850–1100 m.
Records on Mediterranean islands (Fig. 9b). The
subspecies was repeatedly found in a small area in Cen-
tral Euboea. Syntopic Raphidioptera on Euboea: Phae-
ostigma (Ph.) euboica, Ph. (Magnoraphidia) ammi,
Ph. (M.) wewalkai, Raphidia (R.) mediterranea.
Continental distribution. Greece (Parnis, Pat-
eras mountains).
Biogeography. Ph. (G.) divina retsinata is an ex-
tremely stationary Balkanopontomediterranean faunal el-
ement with a few refugial subcenters within a small area
in Greece.
Subgenus Crassoraphidia H. Aspöck & U.
Aspöck, 1968
Crassoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as a sub-
genus of Raphidia L.) [type species by original designation: Raphidia
cyprica Hagen, 1867]: H. Aspöck et al. 1989 (biogeogr, distr; ill: distr-
map); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991 (cat, nom);
H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol); H. Aspöck 2012
(cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Crassoraphidia is
a well dierentiated subgenus with three species one of
which occurring on the island of Cyprus.
Biology and ecology. Larvae are mainly soil-dwell-
ing, 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 al-
titudes of 600–1500 m.
Distribution. SW- and S-Anatolia, Cyprus, Lebanon.
Phaeostigma (Crassoraphidia) cyprica (Hagen, 1867)
Raphidiacyprica Hagen, 1867 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiaphoenicia H. Aspöck & U. Aspöck, 1964a (odescr): H. As-
pöck et al. 1991 (mon).
Phaeostigma (Crassoraphidia) cyprica (Hagen): H. Aspöck et al. 1989
(biogeogr, distr); H. Aspöck et al. 1991 (mon); H. Aspöck and Hölzel
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1996 (distr); H. Aspöck et al. 2001 (anncat); Dobosz 2007 (distr, rec;
ill: map); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Phaeostigmacyprica (Hagen): Haring et al. 2011 (phyl).
Taxonomy. Crassoraphidia cyprica (Fig. 3d) is close-
ly related to C. knappi, from which it can be separated
morphologically by slight dierences of male and fe-
male genitalia.
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. 10a). Cy-
prus. Ph. (C.) cyprica occurs in suitable habitats probably
all over the island. Syntopic Raphidioptera in Cyprus:
Ulrikesyriaca.
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 dierentiated morphologically from specimens from
Near East. One may assume that genomic studies will
lead to a better understanding.
Subgenus Magnoraphidia H. Aspöck & U. Aspöck,
1968
Magnoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidiamajor Burmeister, 1839]: H. Aspöck et al. 1989 (biogeogr,
distr; ill: distrmap); H. Aspöck et al. 1991 (mon); Oswald and Penny
1991 (cat, nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002
(biol); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck 2013 (cat,
etymol), 2014 (cat).
Note.Magnoraphidia is – at least on the basis of charac-
ters of the male genitalia – a well dierentiated subgenus
with six known species. Three of these species have been
found on Mediterranean islands.
Biology. Larvae of at least ve 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, Cen-
tral Europe.
Phaeostigma (Magnoraphidia) major (Burmeister, 1839)
Raphidiamajor Burmeister, 1839 (odescr): H. Aspöck et al. 1991 (mon).
Phaeostigma (Magnoraphidia) major (Burmeister 1839): H. Aspöck
et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); Deve-
tak 1992b (distr); H. Aspöck and Hölzel 1996 (distr); H. Aspöck
et al. 2001 (anncat); Tröger 2002 (rec); U. Aspöck and H. Aspöck
2005a (biogeogr); U. Aspöck and H. Aspöck 2005b (ill: pu); Lange
2010 (rec); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat);
Sziráki 2014 (rec); Weissmair et al. 2021 (biol, ecol).
Phaeostigma major (Burmeister): H. Aspöck et al. 1991 (mon);
Saure and Gerstberger 1991 (ecol, rec); Horstmann 1994 (paras);
Röhricht 1996 (rec); Saure 1996 (biogeogr, ecol, rec); Sziráki and
Popov 1996 (rec); Achtelig 1997 (ecol, rec); Sziráki 1999 (rec);
Röhricht 2000 (rec); Ábrahám 2001 (list, rec); Gruppe et al. 2004
(ecol, rec); Popov 2004 (chorol); Ábrahám 2006 (rec); Gruppe
2006b (ecol, rec); Gruppe and Müller 2007 (ecol, rec); Gruppe
2008 (ecol, rec).
Phaeostigma (Magnoraphidia) majus (Burmeister): Letardi et al.
2010 (rec).
Phaeostigmamajus (Burmeister): Tillier et al. 2022a (rec).
Taxonomy. H. Aspöck et al. (1991). Ph. (M.) major
(Fig. 3e, f) is the sister taxon of all other species of the
major complex (i.e. all species of the subgenus except
Ph. (M.) klimeschi (H. Aspöck & U. Aspöck) and can
easily be distinguished in both sexes by characters of the
genital segments.
Biology and ecology. Larvae (Fig. 3g) live under bark
of a great variety of deciduous trees, but may sometimes
(particularly in the southeast of Europe) be found on co-
nifers. Development two, three or more years. Last hiber-
nating stage: full-grown larva. Adults: (IV)V–VI(VII).
Ph. (M.) major occurs in a great variety of habitats with
trees in altitudes from 0 to ca. 1000 m.
Records on Mediterranean islands (Fig. 9a). Levkas,
Kefalonia. Probably the species can also be found on oth-
er Ionian islands. Syntopic species on Levkas: Parvora-
phidia microstigma, Ornatoraphidia avilabris, Subilla
artemis, Dichrostigmaavipes.
Continental distribution. Balkan Peninsula, Central
Europe (including eastern France and northern Germa-
ny), Eastern Europe.
Biogeography. Ph. (M.) major is a monocentric Bal-
kanopontomediterranean faunal element with consider-
able expansivity. Ph.major is the only monocentric Bal-
kanopontomediterranean snakey which has reached the
northern coast of Germany.
Phaeostigma (Magnoraphidia) ammi (H. Aspöck &
U. Aspöck, 1973)
Raphidia (Magnoraphidia) ammi H. Aspöck & U. Aspöck, 1973 (ode-
scr): H. Aspöck et al. 1991 (mon).
Phaeostigma (Magnoraphidia) ammi (H. Aspöck & U. Aspöck, 1973):
H. Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon);
Popov 1992 (biogeogr); H. Aspöck and Hölzel 1996 (distr); H. As-
pöck et al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck and U.
Aspöck 2013 (cat, etymol), 2014 (cat).
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Taxonomy. H. Aspöck et al. (1991). Ph. (M.) ammi (Fig.
3h) forms together with Ph. (M.) horticola, Ph. (M.) ro-
busta, and Ph. (M.) wewalkai a group of closely related
species within the major complex. These species can be
dierentiated only by characters of the genitalia, which is
usually an easy task. However, on the island of Euboea
where two of the three species – Ph. (M.) ammi and Ph.
(M.) wewalkai – occur in a small area around the village
of Seta in the Dirphys mountains intermediate individu-
als between Ph.ammi and Ph.wewalkai can be found,
which we interpret as hybrids between the two species.
On all other places on Euboea as well as on the continent
the populations are homogenous.
Biology and ecology. Larvae (probably exclusively)
corticolous on a great variety of deciduous trees and coni-
fers in altitudes of 100 m (Skopelos) to 1100 m (Euboea).
Development two to three (or more) years. Last hibernat-
ing stage: full-grown larva. Adults: V-VI.
Records on Mediterranean islands (Fig. 9a). Sko-
pelos, Euboea.
Continental distribution. Conned to a small part of
Greece (Sterea Ellas: Othrys, Pilion mountains).
Biogeography. Extremely stationary, monocentric
Balkanopontomediterranean faunal element with several
isolated populations conned to single mountain ranges.
Phaeostigma (Magnoraphidia) wewalkai (H. Aspöck
& U. Aspöck, 1971)
Raphidia (Magnoraphidia) wewalkai H. Aspöck & U. Aspöck, 1971a
(odescr): H. Aspöck et al. 1991 (mon).
Phaeostigma (Magnoraphidia) wewalkai (H. Aspöck & U. Aspöck): H.
Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon);
Popov 1992 (biogeogr); H. Aspöck and Hölzel 1996 (distr); H. As-
pöck et al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck and U.
Aspöck 2013 (cat, etymol; ill: imag), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Ph. (M.) wewalkai
(Fig. 4a, b) can easily be distinguished from the spe-
cies of its sister group (= (Ph.ammi + Ph.horticola) +
Ph.robusta) by characters of the male and female geni-
talia. However, (at least) in a small area on the Dirphys
mountain range on Euboea, where it occurs sympatric
and syntopic with Ph. (M.) ammi, intermediate phaena
occur. We interpret these phaena as hybrids between the
two species. See also under Ph. (M.) ammi.
Biology and ecology. Larvae (probably exclusively)
corticolous on deciduous trees and on conifers in vari-
ous warmer habitats with rich vegetation from sea lev-
el 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. 8b). Euboea
(Dirphys and Ochi mountain ranges) and various spots in
the southwest of the island. Most probably the species
occurs in all parts of Euboea.
Continental distribution. Restricted to a small part of
eastern Sterea Ellas.
Biogeography. Extremely stationary, monocentric
Balkanopontomediterranean faunal element.
Subgenus Pontoraphidia H. Aspöck & U. Aspöck, 1968
Pontoraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidiapontica Albarda, 1891]: H. Aspöck et al. 1991 (mon); H.
Aspöck et al. 1989 (biogeogr, distr; ill: distrmap); Oswald and Pen-
ny 1991 (cat, nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002
(biol); H. Aspöck and U. Aspöck 2007 (ill: map); H. Aspöck 2012
(cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Phidiara U. Aspöck & H. Aspöck, 1968 (odescr) (described as a subge-
nus of Raphidia L.) [type species by original designation: Raphidia
grandii Principi, 1960]: H. Aspöck et al. 1991 (mon), H. Aspöck and
U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Pontoraphidia is
a well dierentiated subgenus with four species, one of
them with two subspecies. The sister taxon is possibly
Magnoraphidia.
Biology and ecology. Larvae of all species soil-dwell-
ing 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 alti-
tudes from 600–2200 m.
Distribution. The distribution comprises the south-
ern parts of the Apennine Peninsula, the northern Balkan
Peninsula, Eastern Europe and Anatolia.
Phaeostigma (Pontoraphidia) setulosa H. Aspöck & U.
Aspöck, 1967b
Raphidiasetulosa H. Aspöck & U. Aspöck, 1967b (odescr): H. Aspöck
et al. 1991 (mon).
Phaeostigma (Pontoraphidia) setulosa (H. Aspöck & U. Aspöck): H.
Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon);
Canbulat and Kiyak 2006 (distr, rec; ill: map); H. Aspöck 2012 (cat);
H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Phaeostigmasetulosa (H. Aspöck & U. Aspöck): H. Aspöck et al. 1991
(mon); Sziráki and Popov 1996 (com, rec).
Taxonomy. H. Aspöck et al. (1991). Ph. (P. ) setulosa is a
polytypic species with scattered records within a consid-
erably large distribution area: eastern Bulgaria (type local-
ity), two spots in western Anatolia, Athos mountain, the
island of Thasos, and northern Hungary. The populations
from Thasos and Athos show dierences compared to the
other populations so that it seems justied to separate them
at the level of subspecies. For a clarication of the some-
what complicated taxonomic situation within the species
molecular phylogeny studies will be necessary. The sister
species of Ph. (P. ) setulosa is probably Ph. (P. ) pontica.
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Phaeostigma (Pontoraphidia) setulosa aegea
H. Aspöck, U. Aspöck & Rausch, 1991
Phaeostigma (Pontoraphidia) setulosaaegea H. Aspöck, U. Aspöck &
Rausch, 1991 (odescr, mon); H. Aspöck and Hölzel 1996 (distr); H.
Aspöck 2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol),
2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Eidonomic charac-
ters with those of Ph. (P. ) s.setulosa agreeing, male and
female genitalia, however, dierent.
Biology and ecology. Larvae most probably exclu-
sively 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 is-
land of Thasos in glades within light pine forests at an alti-
tude of about 1000 m in large numbers on fruits of Aspho-
delus (on the Athos Mountain in 1600 m on pine trees).
Records on Mediterranean islands (Fig. 9a). Thasos
(Ipsarion). Syntopic Raphidioptera on Thasos: Venust-
oraphidianigricollis, Raphidia (R.) beieri.
Continental distribution. Athos mountain (Chalkidi-
ki, Greece).
Biogeography. Ph. (P. ) setulosaaegea is an extremely
stationary monocentric Balkanopontomediterranean fau-
nal element. The species – Ph. (P.) setulosa (s. l.) – is a
polycentric Balkanapontomediterranean-Anatoloponto-
mediterranean faunal element of moderate expansivity. It
will certainly be found in many parts of the Balkan Pen-
insula and in Anatolia.
Subgenus Aegeoraphidia H. Aspöck, U. Aspöck &
Rausch, 1991
Aegeoraphidia H. Aspöck & U. Aspöck & Rausch, 1991 (odescr, mon)
[type species by original designation: Raphidia (Phidiara) raddai U.
Aspöck & H. Aspöck, 1969]. H. Aspöck et al. 1991 (mon); H. As-
pöck 2002 (biol); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck
2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). This subgenus com-
prises two groups of species – the raddai-ressli group with
ve species (Ph. (Ae.) raddai, Ph. (Ae.) ressli, Ph. (Ae.)
porphetica, Ph. (Ae.) biroi, and Ph. (Ae.) vartianorum) and
the noane group with two species (Ph. (Ae.) noane and Ph.
(Ae.) remane). Of these, four species occur on Mediterra-
nean islands. Aegeoraphidia can be clearly dierentiated
from other subgenera particularly by characters of the male
genitalia. The sister group of the subgenus is still unknown.
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 com-
prises Crete, several islands in the eastern Aegean Sea, west-
ern, southern, and southeastern Anatolia, and northern Iraq.
Phaeostigma (Aegeoraphidia) raddai (U. Aspöck & H.
Aspöck, 1969)
Raphidia (Phidiara) raddai U. Aspöck & H. Aspöck, 1969 (odescr): H.
Aspöck et al. 1991 (mon).
Phaeostigma (Aegeoraphidia) raddai (U. Aspöck & H. Aspöck): H. Aspöck
et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); H. Aspöck
and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat); Canbulat and
Kiyak 2006 (distr, rec; ill: map); Dobosz 2007 (distr, rec); H. Aspöck
2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Ph. (Ae.) raddai can
be easily dierentiated from the other species of the sub-
genus even by eidonomic characters and particularly by
its typical male genitalia.
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. Devel-
opment two, three or more years. Last hibernating stage:
full-grown larva. Adults: IV–VI occurring in various hab-
itats from sea level to 800 m.
Records on Mediterranean islands (Fig. 9b). Les-
bos, Chios, Samos, Ikaria. The species will also be found
on other islands in the eastern Aegean Sea. Syntopic
Raphidioptera species: Raphidia (R.) mysia (Lesbos),
R.ambigua (Samos), R.peteressli (Chios), R.mediterra-
nea (Ikaria), Parainocelliaressli (Samos).
Continental distribution. Southwestern Anatolia.
Biogeography. Monocentric, stationary Anatolopon-
tomediterranean faunal element comprising a relatively
small area east of the Rechinger line.
Phaeostigma (Aegeoraphidia) prophetica (H. Aspöck
& U. Aspöck, 1964a)
Raphidiaprophetica H. Aspöck & U. Aspöck, 1964a (odescr): H. As-
pöck et al. 1991 (mon).
Phaeostigmaressliprophetica (H. Aspöck & U. Aspöck): H. Aspöck et
al. 1991 (mon).
Phaeostigma prophetica (H. Aspöck & U. Aspöck): H. Aspöck et al.
1991 (mon); U. Aspöck and H. Apöck 1989 (tax).
Phaeostigma (Aegeoraphidia) prophetica (H. Aspöck & U. Aspöck): H.
Aspöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon);
H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat);
H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol),
2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Closely related to
Ph. (Ae.) ressli, from which it can be dierentiated only
by ne, but always recognizable dierences in the male
genitalia. The main reason for treating Ph. (Ae.) prophet-
ica as a species and not a subspecies of Ph. (Ae.) ressli
is the fact that not Ph. (Ae.) ressli but another species of
this clade, Ph. (Ae.) vartianorum, occurs in those parts of
Anatolia opposite to Rhodes.
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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. 9b).
Rhodes, probably endemic to this island and apparently
occurring all over Rhodes. Syntopic Raphidioptera spe-
cies: Subillacolossea, Raphidia (R.) ambigua.
Biogeography. Monocentric, extremely stationary
Anatolopontomediterranean faunal element. Endemic
to Rhodes.
Phaeostigma (Aegeoraphidia) karpathana (U. Aspöck
& H. Aspöck, 1989)
Phaeostigma (Aegeoraphidia) karpathana U. Aspöck & H. Apöck,
1989 (odescr): H. Aspöck and Hölzel 1996 (distr); H. Aspöck et
al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck
2013 (cat, etymol), 2014 (cat).
Taxonomy. U. Aspöck and H. Aspöck (1989). In its eido-
nomic characters the species agrees with Ph. (Ae.) ressli
and Ph. (Ae.) prophetica, but it can easily be dierentiat-
ed by characters of the male and female genitalia.
Biology and ecology. Larvae soil-dwelling, but excep-
tionally also corticolous (ndings 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 (Pinushalepensis), open slopes
with Genista and Crataegus in altitudes from 50–750 m.
Adults in extraordinarily high numbers on Genistafasse-
lata and also on young pine trees.
Records on Mediterranean islands (Fig. 9b). Only
from Karpathos. Probably endemic to the island. Syn-
topic Raphidioptera species: Raphidia (R.) mediterranea.
It is possible that Ph. (Ae.) karpathana occurs also on the
near island of Kasos, but certainly not on other islands.
Biogeography. Monocentric, extremely stationary
species, endemic to Karpathos. The species can nei-
ther be classied 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 (Navás, 1915)
Lesnabiroi Navás, 1915a (odescr): H. Aspöck et al. 1991 (mon).
Raphidialabyrintha H. Aspöck & U. Aspöck, 1964a (odescr): H. As-
pöck et al. 1991 (mon).
Phaeostigmabiroi (Navás): H. Aspöck et al. 1991 (mon); U. Aspöck
and H. Apöck 1989 (tax); U. Aspöck and H. Aspöck 1990 (distr);
Ábrahám and Papp 1994 (com, FD!).
Phaeostigma (Aegeoraphidia) biroi (Navás): H. Aspöck et al. 1989
(biogeogr, distr); H. Aspöck et al. 1991 (mon); Popov 1992 (bio-
geogr); H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001
(anncat); Tröger 2005a (rec); H. Aspöck 2012 (cat); H. Aspöck and
U. Aspöck 2013 (cat, etymol; ill: ♂ imag, pu), 2014 (cat); Sziráki
2014 (rec).
Taxonomy. H. Aspöck et al. (1991). Ph. (Ae.) biroi
(Fig. 4c, d) is probably the sister taxon of the other species
of the biroi-ressli complex, from which it can be clearly
separated by characters of the male genitalia. Moreover,
in most cases even a dierentiation on the basis of eido-
nomic characters is possible: The pterostigma of Ph. (Ae.)
biroi is in its distal part paler than proximally, sometimes
even yellow. Moreover, Ph. (Ae.) biroi is the only Aegeo-
raphidia species occurring in Crete.
Biology and ecology. Larvae (Fig. 4e) probably cor-
ticolous (records from Pinus, Amygdalus, Quercus) and
soil-dwelling. Development two or three years. Last hi-
bernating stage: full-grown larva. Adults: IV–VI. Many
records on Genista, Ilex, Acer, Crataegus, Pinusbrutia
in various, mainly warm and relatively dry habitats from
sea level to 1500 m.
Records on Mediterranean islands (Fig. 9b). There
are many records from Crete, but none from other islands.
Syntopic Raphidioptera: Phaeostigma (Superboraphidia)
minois, Raphidia (R.) ariadne, Fibla (Reisserella) pasiphae.
Biogeography. Monocentric, stationary Cretan faunal
element. Endemic to Crete.
Subgenus Superboraphidia H. Aspöck &
U. Aspöck, 1968
Superboraphidia H. Aspöck & U. Aspöck, 1968 (odescr) (described as
a subgenus of Raphidia L.) [type species by original designation:
Raphidiaauberti H. Aspöck & U. Aspöck, 1966]: H. Aspöck et al.
1989 (biogeogr, distr; ill: distrmap); U. Aspöck and H. Aspöck 1990
(syst); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991 (cat,
nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol); H.
Aspöck 2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol),
2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Superboraphid-
ia comprises ve species (Phaeostigma (S.) auberti,
Ph. (S.). rauschi, Ph. (S.) mammaphila, Ph. (S.) turci-
ca, and Ph. (S.) minois), which are not closely related
but show a special synapomorphy in the male genitalia,
which justies to unite them in a subgenus. One of the
ve species occurs on a Mediterranean island.
Biology. Larvae of probably all ve species are
soil-dwelling. Development insuciently known, prob-
ably 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. Aspöck et al.
1991; Rausch and H. Aspöck 1992).
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Figure 7. a. Records of species of the family Raphidiidae, altogether 30 species on islands of the Mediterranean Sea; b. Records of
Venustoraphidianigricollis, Phaeostigma (Ph.) euboica, and of Parvoraphidiamicrostigma, on Mediterranean islands.
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Figure 8. a. Records of species of the genus Xanthostigma, X.corsica and X.aloysiana, on Mediterranean islands; b. Records of
Ornatoraphidiaavilabris, Phaeostigma (Magnoraphidia) wewalkai, and of Dichrostigmaavipes, on Mediterranean islands.
Phaeostigma (Superboraphidia) minois U. Aspöck &
H. Aspöck, 1990
Phaeostigma (Superboraphidia) minois U. Aspöck & H. Aspöck, 1990
(odescr, ecol, distr; ill: ♀ gs, ♂ gs, wings); Popov 1992 (biogeogr);
H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat);
Tröger 2005a (rec); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck
2013 (cat, etymol), 2014 (cat).
Taxonomy. U. Aspöck and H. Aspöck (1990). The spe-
cies (Fig. 4f) is eidonomically similar to Ph. (Aegeo-
raphidia) biroi, but can be easily dierentiated in both
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Figure 9. a. Records of Ornatoraphidiachristianodagmara, Phaeostigma (Magnoraphidia) major, Ph. (M.) ammi, Ph. (Pontora-
phidia) setulosaaegea, and of Ph. (Superboraphidia) minois, on Mediterranean islands; b. Records of Phaeostigma (Grecoraphid-
ia) divinaretsinata, and of species of Aegeoraphidia, Ph. (Aegeoraphidia) raddai, Ph. (Ae.) prophetica,Ph. (Ae.) karpathana, and
of Ph. (Ae.) biroi, on Mediterranean islands.
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Figure 10. a. Records of Phaeostigma (Crassoraphidia) cyprica, and of Ulrikesyriaca, on Mediterranean islands; b. Records of
species of the genus Subilla, S.connis, S.artemis, S.principiae, and S.colossea, on Mediterranean islands.
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Figure 11. 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. ) maclach-
lani, Fibla (Reisserella) pasiphae, and Parainocelliaressli, on Mediterranean islands.
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sexes by characters of the genitalia. The sister taxon is
unknown, and the species has a rather isolated position
within the subgenus.
Biology and ecology. Larvae probably soil-dwelling.
Larvae and development unknown. Adults: V, on Pinus
brutia and Cypressussempervivens, in light forests with
Quercus, Crataegus etc. in 830–1100 m.
Records on Mediterranean islands (Fig. 9a). Ph. (S.)
minois has so far only been found in a few spots near
the Omalos Plateau in the west of Crete (Chania). Syn-
topic Raphidioptera: Phaeostigma (Aegeoraphidia) biroi,
Raphidia (R.) ariadne, Fibla (Reisserella) pasiphae.
Biogeography. Endemic to Crete. Monocentric,
(extremely) stationary Cretan faunal element.
Subilla Navás, 1916
Subilla Navás, 1916 (odescr) [type species by original designation: Ra-
phidiasericea Albarda, 1891]: H. Aspöck et al. 1989 (biogeogr, distr;
ill: distrmap); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991
(cat, nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol,
paras); H. Aspöck and U. Aspöck 2007 (ill: map); Monserrat and Pa-
penberg 2010 (biogeogr, chorol); Haring et al. 2011 (phyl); U. Aspöck
et al. 2012 (g: phyltree; list); H. Aspöck and U. Aspöck 2013 (cat,
etymol, phyl), 2014 (cat); Monserrat and Papenberg 2015 (synlist).
Taxonomy and systematics. H. Aspöck et al. (1991), Har-
ing et al. (2011). Subilla is a very conspicuous genus clear-
ly dierentiated by imaginal as well as by larval characters
from all other genera of the family. The sister taxon is proba-
bly Tauroraphidia H. Aspöck & U. Aspöck & Rausch, 1982.
In the molecular phylogenetic analysis of the Raphidiidae
(Haring et al. 2011) the genus Ulrike H. Aspöck emerged as
the sister taxon of Subilla, however, Tauroraphidia was not
available for this analysis. The genus comprises altogether
ten species, ve of these (S.connis, S. aliena, S.artemis,
S. xylidiophila, S. walteri) form the connis-group, three
others (S.physodes, S.priapella, S.colossea) the physodes-
group, while S.fatma and S. principiae are somewhat iso-
lated, although they show anities to the physodes-group.
Biology and ecology. Larvae of all species cortico-
lous, 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, West-
ern and Eastern Europe, Anatolia. Altogether four species
have been found on Mediterranean islands: Sardinia, Sic-
ily, Levkas, Rhodes. Each of these islands harbors only
one Subilla species.
Subilla connis (Stephens, 1836)
Raphidiaconnis Stephens, 1836 (odescr): H. Aspöck et al. 1991 (mon);
Suntrup 1990 (ecol, paras, rec); Ábrahám 1992 (rec); Schmitz 1992
(ecol, rec).
Raphidiacognata Rambur, 1842 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiaschneideri Ratzeburg, 1844 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiacolubroides Costa, 1855 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiasericea Albarda, 1891 (odescrr): H. Aspöck et al. 1991 (mon).
Subillaconnis (Stephens): H. Aspöck et al. 1991 (mon); Kielhorn 1991
(ecol, rec); Saure and Gerstberger 1991 (ecol, rec); Ábrahám 1992
(rec); Plant 1992 (com, rec); Morgan 1993 (rec); Horstmann 1994
(paras); Plant 1994 (biol, distrmap, tax); H. Aspöck and Hölzel 1996
(distr); Röhricht 1996 (rec); Saure 1996 (biogeogr, ecol, rec); Achtelig
1997 (ecol, rec); Schubert and Gruppe 1999 (ecol, rec); Sziráki 1999
(rec); Röhricht 2000 (rec); Ábrahám 2001 (list, rec); H. Aspöck et
al. 2001 (anncat); Gruppe and Schubert 2001 (ecol, rec); Plant 2001
(com, rec); Vas et al. 2001 (ecol, rec); Popov 2002 (biogeogr); Gruppe
et al. 2004 (ecol, rec); Haring and U. Aspöck 2004 (phyl); Pantaleoni et
al. 2005 (tax); Pantaleoni 2005 (rec); Ábrahám 2006 (rec); H. Aspöck
and U. Aspöck 2007 (biogeogr, distr); Gruppe 2007a (rec); Gruppe
and Müller 2007 (ecol, rec); Gruppe 2008 (ecol, rec); Weigelmeier
2008 (biol); Letardi et al. 2010 (rec); Nicoli Aldini et al. 2012 (rec,
distr); Haring et al. 2011 (phyl, phyltree); H. Aspöck and U. Aspöck
2013 (cat, etymol), 2014 (cat); Morinière et al. 2014 (barcode, phyl);
Gruppe 2015 (ecol, rec); Weissmair et al. 2021 (biol, ecol).
Taxonomy. H. Aspöck et al. (1991). S.connis (Fig. 4g)
is eidonomically very similar to the other four species of
the connis-group (S.aliena, S.xylidiophila, S.artemins,
S.walteri), but can easily be dierentiated by characters
of the male genitalia. Moreover, no island harbors more
than one species.
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. 10b). There
are only a few records in the northeast of Sicily. Syntopic
other Raphidioptera in Sicily: X.corsica, F.maclachlani.
Continental distribution. Apennine Peninsula, Cen-
tral Europe, France, England, Denmark, Eastern Europe.
Biogeography. Expansive probably monocentric
Adriatomediterranean faunal element.
Subilla artemis (H. Aspöck & U. Aspöck, 1971)
Raphidia (Subilla) artemis H. Aspöck & U. Aspöck, 1971 (odescr): H.
Aspöck et al. 1991 (mon).
Subillaartemis (H. Aspöck & U. Aspöck, 1971): H. Aspöck et al. 1989
(biogeogr, distr); H. Aspöck et al. 1991 (mon); Popov 1992 (bio-
geogr); H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001
(anncat); Pantaleoni et al. 2005 (tax); H. Aspöck 2012 (cat); H. As-
pöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). S.artemis belongs to
the connis-group of the genus and is eidonomically sim-
ilar to the other species. Although the male genitalia show
a surprising variability, they always allow to identify the
species reliably.
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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. De-
velopment two to three years. Last hibernating stage: full-
grown larva. Adults: V–VI.
Records on Mediterranean islands (Fig. 10b).
Levkas. Syntopic Raphidioptera on Levkas: Parvora-
phidia microstigma, Ornatoraphidia avilabris, Phae-
ostigma (Magnoraphidia) major.
Continental distribution. Greece north of the Gulf of
Korinthos, North Macedonia.
Biogeography. S.artemis is a monocentric Balkanopon-
tomediterranean faunal element with moderate expansivity.
Subilla principiae Pantaleoni, U. Aspöck, Cao & H.
Aspöck, 2005
Subillaprincipiae Pantaleoni, U. Aspöck, Cao & H. Aspöck, 2005 (ode-
scr); H. Aspöck and U. Aspöck 2007a (biogeogr, distr); U. Aspöck
and H. Aspöck 2007 (ill: ♂ imag); H. Aspöck 2012 (cat); H. Aspöck
and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. Pantaleoni et al. (2005). S. principiae
(Fig. 4h) can clearly be dierentiated from all other spe-
cies of the genus by characters of the male and female
genitalia. Moreover, the species can be readily recognized
in the eld by the dark long pterostigma. There is no par-
ticularly close relationship to any other Subilla species.
Biology and ecology. So far larvae were found exclu-
sively under the bark of Quercuspubescens. Only known
from the type locality – a light oak forest – at 1050 m.
Adults: V–VI.
Records on Mediterranean islands (Fig. 10b).
Sardinia. The species is endemic to Sardinia, possibly
it occurs also in Corsica. Syntopic Raphidioptera:
Fiblamaclachlani.
Biogeography. Stationary Tyrrhenian faunal element.
Subilla colossea (H. Aspöck, U. Aspöck &
Rausch, 1979)
Raphidia (Subilla) colossea H. Aspöck, U. Aspöck & Rausch, 1979
(odescr): H. Aspöck et al. 1991 (mon).
Subillacolossea (H. Aspöck, U. Aspöck & Rausch, 1979): H. Aspöck et al.
1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); Horstmann 1993
(paras); Horstmann 1994 (paras); H. Aspöck and Hölzel 1996 (distr);
H. Aspöck et al. 2001 (anncat); Pantaleoni et al. 2005 (tax); H. Aspöck
2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). The species belongs
to the physodes-group of the genus. It can easily be iden-
tied by characters of the male and female genitalia.
Biology and ecology. Larvae exclusively corticolous
on Pinus and (rarely) on Quercusilex in light pine for-
ests 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. 10b). En-
demic to Rhodes, probably distributed all over the island.
Syntopic Raphidioptera: Phaeostigma (Aegeoraphidia)
prophetica, Raphidia (R.) ambigua.
Biogeography. Stationary Anatolopontomediterra-
nean faunal element.
Ulrike H. Aspöck, 1968
Ulrike H. Aspöck, 1968 (odescr) (described as subgenus of Raphidia
L.) [type species by original designation: Agullaattica H. Aspöck &
U. Aspöck, 1967a]: H. Aspöck et al. 1989 (biogeogr, distr; ill: dis-
trmap); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991 (cat,
nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol); Haring
et al. 2011 (phyl); U. Aspöck et al. 2012 (g: phyltree); H. Aspöck
and U. Aspöck 2013 (cat, etymol, phyl), 2014 (cat).
Taxonomy and systematics. H. Aspöck et al. (1991), Har-
ing et al. (2011). Ulrike is a conspicuous genus markedly
dierentiated from all other genera of the family by eido-
nomic characters as well as by those of the (particularly
male) genitalia. It comprises two closely related species
occurring in eastern parts of the Mediterranean region. In a
molecular phylogenetic analysis (Haring et al. 2011) Subil-
la has emerged as the sister taxon of Ulrike. Based on mor-
phological evidence, Subilla is probably the sister taxon
of Tauroraphidia, and both represent the sister of Ulrike.
Biology. See Ulrikesyriaca.
Distribution. Greece (Attica), Cyprus, Near East.
Ulrike syriaca (Steinmann, 1964)
Raphidilla syriaca Steinmann, 1964 (odescr): H. Aspöck et al.
1991 (mon).
Ulrikesyriaca (Steinmann, 1964): H. Aspöck et al. 1989 (biogeogr, dis-
tr); H. Aspöck et al. 1991 (mon); H. Aspöck and Hölzel 1996 (distr);
H. Aspöck et al. 2001 (anncat); Haring et al. 2011 (phyl, phyltree);
H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). U.syriaca (Fig. 5a)
is closely related to the second species of the genus,
U.attica (H. Aspöck & U. Aspöck), which is so far known
from the mainland of Greece only. The species can easily
be identied also by eidonomic characters.
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. 10a).
Records only from Cyprus. Syntopic Raphidioptera (in
Cyprus): Phaeostigma (Crassoraphidia) cyprica.
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).
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Raphidia Linnaeus, 1758 s.l.
Raphidia Linnaeus, 1758 (odescr) [type species by monotypy: Raphidia
ophiopsis Linnaeus, 1758]: H. Aspöck at al. 1989 (biogeogr, distr;
ill: distrmap); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991
(cat, nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol,
paras); Haring et al. 2011 (phyl); U. Aspöck et al. 2012 (g: phyltree;
list); H. Aspöck and U. Aspöck 2013 (cat, etymol, phyl), 2014 (cat).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). With 18 species in three subgenera
(Raphidia s. str.: 15 spp., Aserbeidshanoraphidia H. As-
pöck & U. Aspöck: 1 sp., Nigroraphidia H. Aspöck & U.
Aspöck: 2. spp.). Raphidia is – after Mongoloraphidia
and Phaeostigma – the third largest genus of the family
Raphidiidae. Only the subgenus Raphidia s. str. is repre-
sented on Mediterranean islands with altogether 6 species,
ve of these (R.mediterranea, R.beieri, R.peterressli, R.
mysia, R. ambigua) belong to a group of more closely
related species, while R. ariadne is somewhat isolated.
In a molecular phylogenetic study (Haring et al. 2011)
Raphidia s. l. has emerged as the sister of Dichrostigma +
Tjederiraphidia, but this needs conrmation.
All species can easily be dierentiated 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 mainly one 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.
Subgenus Raphidia Linnaeus, 1758, s. str.
Raphidia Linnaeus, 1758 (odescr) [type species by monotypy: Raphidia
ophiopsis Linnaeus, 1758]: H. Aspöck et al. 1989 (biogeogr, distr;
ill: distrmap); H. Aspöck et al. 1991 (mon); H. Aspöck 2002 (biol,
paras); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck 2013 (cat,
etymol), 2014 (cat).
Pretzmannia H. Aspöck & U. Aspöck, 1968 (described as a subgenus of
Raphidia L.) [type species by original designation: Raphidiaeuxina
Navás, 1915d]: H. Aspöck et al. 1991 (mon); Oswald and Penny
1991 (cat, nom); H. Aspöck and U. Aspöck 2014 (cat).
Raphidia (Raphidia) mediterranea H. Aspöck, U.
Aspöck & Rausch, 1977
Raphidia (Raphidia) ophiopsis mediterranea H. Aspöck, U.Aspöck &
Rausch, 1977 (odescr): H. Aspöck et al. 1989 (biogeogr, distr); H.
Aspöck et al. 1991 (mon); Popov 1993 (com); U. Aspöck et al. 1995
(ethol: cop); H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al.
2001 (anncat); H. Aspöck 2012 (cat).
Raphidia (Raphidia) mediterranea H. Aspöck, U. Aspöck & Rausch: H.
Aspöck et al. 1991 (mon); Letardi and Pantaleoni 1996 (com, rec);
H. Aspöck and U. Aspöck 2007 (biogeogr, distr); H. Aspöck and U.
Aspöck 2013 (cat, etymol; ill: imag, ♂ imag), 2014 (cat); Canbulat
2014 (biogeogr, distr); Sziráki 2014 (rec); Rausch et al. 2016 (tax,
rec, distr, ecol); Gruppe et al. 2017 (rec, ecol); H. Aspöck et al. 2017
(molecsyst, phyl, rec, distr, biogeogr).
Raphidiamediterranea H. Aspöck, U. Aspöck & Rausch: H. Aspöck et
al. 1991 (mon); Sziráki 1993 (nom, rec; ill: distrmap, gs); Güsten
1998 (rec); Pantaleoni 2005 (com, rec); Sziráki 2010 (ill: imag);
Tillier et al. 2022a (rec); Gruppe et al. 2023 (biol).
Taxonomy. H. Aspöck et al. (1991), Sziráki (1993), H.
Aspöck et al. (2017) (Fig. 5b, c). This species was origi-
nally described as a subspecies of Raphidiaophiopsis L.,
from which it diers morphologically slightly in charac-
ters of the male (and also female) genitalia. Based upon
considerable ecological characters R. mediterranea has
been elevated to species rank, and later this decision was
corroborated by a molecular systematic analysis (H. As-
pöck et al. 2017).
Biology and ecology. Larvae (Fig. 5d) soil-dwelling,
in high altitudes in Greece, at least partly, corticolous.
(Larvae of the isolated population in a farm house in Up-
per Austria develop in the straw of the fetched roof of
the old building; Gruppe et al. 2017.) Development one
or two years. Last hibernating stage: full-grown larva.
Adults: IV–VI(VII). The preferred habitats on altitudes
below 500 m are all kinds of maquis, often with mass
occurrence. In high altitudes (records at 1100 m) in light
forests, also in gardens.
Records on Mediterranean islands (Fig. 11a).
Skyros, Euboea, Andros, Hydra, Aegina, Naxos, Paros,
Ikaria, Karpathos. Syntopic Raphidioptera species
on Mediterranean islands: Phaeostigma (Ph.) eubo-
ica (Euboea), Ph. (Graecoraphidia) divina retsinata
(Euboea), Ph. (Magnoraphidia) wewalkai (Euboea),
Ph. (M.) ammi (Euboea), Ph. (Aegeoraphidia) raddai
(Ikaria), Ph. (Ae.) karpathana (Karpathos).
Continental distribution. Greece, Bulgaria, Roma-
nia, 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. Aspöck et al. 2017). The populations on Karpathos
show slight dierences compared to populations from
other islands as well as from the continent, maybe due to
an immigration long ago.
Raphidia (Raphidia) beieri H. Aspöck &
U. Aspöck, 1964a
Raphidiabeieri H. Aspöck & U. Aspöck, 1964a (odescr): H. Aspöck
et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); Popov
2004 (chorol).
Raphidiaschizurotergalis Bartoš, 1965 (odescr): H. Aspöck et al. 1991
(mon); H. Aspöck and U. Aspöck 2014 (cat).
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Raphidia (Raphidia) beieri H. Aspöck & U. Aspöck: H. Aspöck et al.
1991 (mon); H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al.
2001 (anncat); Popov 2001 (rec); Dobosz 2007 (distr, rec); H. Aspöck
2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Raphidia (R.) beieri
is closely related to R. (R.) kimminsi (which occurs in the
north of Anatolia) and R. (R.) grusinica (which occurs in
NE-Anatolia and in the western Caucasian regions) and
can be dierentiated only by characters of the male geni-
talia. Possibly these taxa form a superspecies. A clarica-
tion of the complicated taxonomic situation will only be
possible by molecular systematic analyses.
Biology and ecology. Euryoecious! Larvae cortico-
lous and soil-dwelling; many records under bark of Ab-
ies, Pinus, Quercus, Pyrus and Prunus, and in the litter
around roots of bushes. Development usually two years,
rarely one 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. 11a). Tha-
sos, Samothraki. Syntopic species of Raphidioptera on
Thasos: Phaeostigma (Pontoraphidia) setulosa aegea,
Venustoraphidianigricollis.
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 H. Aspöck &
U. Aspöck, 1973
Raphidia (Raphidia) peterressli H. Aspöck & U. Aspöck, 1973 (ode-
scr): H. Aspöck et al. 1991 (mon); H. Aspöck and Hölzel 1996 (dis-
tr); H. Aspöck et al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck
and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Raphidia peterressli H. Aspöck & U. Aspöck: H. Aspöck et al. 1989
(biogeogr, distr).
Taxonomy. H. Aspöck et al. (1991). Probably an insular
isolate of R.ambigua, clearly dierentiated by characters
of the male genitalia, a molecular systematic analysis in
comparison with R. ambigua and other species of Ra-
phidia s. str. is, however, needed.
Biology and ecology. Larvae not yet described, prob-
ably mainly soil-dwelling, but ndings 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. 11a). So far
only known from the island of Chios. Syntopic Raphidi-
optera: Phaeostigma (Aegeoraphidia) raddai.
Continental distribution. No records.
Biogeography. Probably endemic to Chios and thus
an extremely stationary Anatolopontomediterranean fau-
nal element.
Raphidia (Raphidia) mysia H. Aspöck, U. Aspöck &
Rausch, 1991
Raphidia (Raphidia) mysia H. Aspöck, U. Aspöck & Rausch, 1991
(mon, odescr): H. Aspöck and Hölzel 1996 (distr); H. Aspöck et
al. 2001 (anncat); H. Aspöck 2012 (cat); H. Aspöck and U. Aspöck
2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Eidonomically very
similar and closely related to several other species of the
genus (R.beieri, R.ambigua); by characters of the male
genitalia, however, easily to be identied.
Biology and ecology. Larvae at least also corticolous
(ndings on oak), but probably mainly soil-dwelling. De-
velopment two years. Last hibernating stage: full-grown
larva. Adults: V(–VI). Euryoecious! Often high popula-
tion densities in various habitats (e.g. light forests, pas-
tures with small areas of shrubs) and on dierent plants:
Pistaciaterebinthus, Pistacialentiscus, Phillyreatrifolia,
Pinushalepensis.
Records on Mediterranean islands (Fig. 11a). So
far only on Lesbos. Syntopic Raphidioptera (on Lesbos):
Phaeostigma (Aegeoraphidia) raddai.
Continental distribution. NW-Anatolia.
Biogeography. Stationary Anatolopontomediterra-
nean faunal element.
Raphidia (Raphidia) ambigua H. Aspöck &
U. Aspöck, 1964b
Raphidiaambigua H. Aspöck & U. Aspöck, 1964b (odescr): H. Aspöck
et al. 1991 (mon); H. Aspöck et al. 1989 (biogeogr, distr).
Raphidia (Raphidia) ambigua H. Aspöck & U. Aspöck: H. Aspöck et
al. 1991 (mon); H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al.
2001 (anncat); Canbulat and Özsara 2004 (distr, rec); Canbulat and
Kiyak 2006 (distr, rec; ill: map); Dobosz 2007 (distr, rec); H. Aspöck
2012 (cat); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). The species is closely
related to the other species of this group within the subge-
nus Raphidia s. str., but can easily be identied by char-
acters of the male genitalia. There is no island on which a
second Raphidia species besides R.ambigua occurs, there-
fore, it cannot be confused with other species of Raphidia.
Biology and ecology. Larvae corticolous on conifers
and deciduous trees as well as soil-dwelling. Develop-
ment one year. Last hibernating stage: full-grown larva.
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Adults: (IV)V–VI(VII). Euryoecious! In various habi-
tats: 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. 11a). Samos,
Rhodes. Syntopic Raphidioptera on islands: Phaeostig-
ma (Aegeoraphidia) raddai (Samos), Ph. (Ae.) prophet-
ica (Rhodes), Subilla colossea (Rhodes), Parainocellia
ressli (Samos).
Continental distribution. Most parts of Anatolia,
from the Mediterranean coast until the east.
Biogeography. Monocentric Anatolopontomediterra-
nean faunal element with high expansivity.
Raphidia (Raphidia) ariadne H. Aspöck &
U. Aspöck, 1964a
Raphidiaariadne H. Aspöck & U. Aspöck, 1964a (odescr): H. Aspöck
et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); U. Aspöck
et al. 2012 (g: imag).
Raphidia (Raphidia) ariadne H. Aspöck & U. Aspöck: H. Aspöck et
al. 1991 (mon); Popov 1992 (biogeogr); H. Aspöck et al. 2001 (an-
ncat); Tröger 2005a (rec); H. Aspöck 2012 (cat); H. Aspöck and U.
Aspöck 2013 (cat, etymol, ill: ♀ imag, ♂ imag), 2014 (cat); Sziráki
2014 (rec).
Taxonomy. H. Aspöck et al. (1991). A very conspicuous
species (Fig. 5e, f), somewhat isolated within the subge-
nus Raphidia s. str., it can hardly be confused with any
other species of the family.
Biology and ecology. Larvae (Fig. 5g) mainly
soil-dwelling, but also corticolous: several ndings of lar-
vae and pupae under bark of Pyrus on the Omalos Plateau
(1000–1100 m). Development one or two years. Last hi-
bernating stage: full-grown larva. Adults: IV–VI. Euryoe-
cus! In almost all habitats with coniferous or deciduous
trees or bushes all over the island of Crete from sea level
to (at least) 1550 m. Usually high population densities.
Records on Mediterranean islands (Fig. 11a). Endem-
ic to Crete. No records from elsewhere. Syntopic Raphid-
ioptera: Phaeostigma (Aegeoraphidia) biroi, Phaeostigma
(Superboraphidia) minois, Fibla (Reisserella) pasiphae.
Biogeography. Monocentric Cretan faunal element.
Dichrostigma Navás, 1909
Dichrostigma Navás, 1909 (described as a section of Raphidia L.) [type
species by subsequent designation: Raphidia avipes Stein, 1863]:
H. Aspöck et al. 1989 (biogeogr, distr; ill: distrmap); H. Aspöck et
al. 1991 (mon); Oswald and Penny 1991 (cat, nom); H. Aspöck et
al. 2001 (anncat); H. Aspöck 2002 (biol, paras); Haring et al. 2011
(phyl); U. Aspöck et al. 2012 (g: phyltree); H. Aspöck and U. As-
pöck 2013 (cat, etymol, phyl), 2014 (cat).
Lesna Navás, 1915b (odescr) [type species by original designation:
Raphidiaadanana Albarda, 1891]: H. Aspöck et al. 2001 (anncat).
Taxonomy and systematics. H. Aspöck et al. (1991).
Dichrostigma is a markedly dierentiated genus. In a
molecular phylogeny analysis (Haring et al. 2011) it has
emerged as the sister group of Tjederiraphidia and both
are the sister of Raphidia. Dichrostigma comprises four
species, which can easily be dierentiated also by eido-
nomic characters.
Biology and ecology. Larvae soil-dwelling. Develop-
ment 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 alti-
tudes from sea level to 1800 m.
Distribution. Central-, E-, SE-Europe, Anatolia, Near
East. One species occurs on Mediterranean islands.
Dichrostigma avipes (Stein, 1863)
Raphidiaophiopsis var. e: Schummel, 1832 (descr, distr): H. Aspöck et
al. 1991 (mon).
Raphidiaanis Schneider, 1843 (odescr, ecol, distr): Novak 1891 (rec);
H. Aspöck et al. 1991 (mon). – Homonym!
Raphidiaavipes Stein, 1863 (odescr): H. Aspöck et al. 1991 (mon);
Dobosz 1989 (rec); Dobosz 1994 (rec).
Raphidia (Dichrostigma) avipes Stein: H. Aspöck et al. 1991 (mon);
Dobosz 1991 (distr, ecol, rec; ill: map); Tröger 1993 (rec).
Subillasulfuricosta Steinmann, 1963 (odescr): H. Aspöck et al. 1991 (mon).
Raphidiasinica Steinmann, 1964 (odescr): H. Aspöck et al. 1991 (mon).
Raphidia maculicaput Steinmann, 1964 (odescr): H. Aspöck et al.
1991 (mon).
Raphidiadichroma Steinmann, 1964 (odescr): H. Aspöck et al. 1991 (mon).
Raphidia durmitorica Steinmann, 1964 (odescr): H. Aspöck et al.
1991 (mon).
Raphidiamonotona Steinmann, 1964 (odescr): H. Aspöck et al. 1991 (mon).
Subillabalesdenti Poivre, 1991 (odescr; ill: imag, gs, head, wings): H.
Aspöck et al. 2001 (anncat).
Raphidia (Lesna) avipes Stein: H. Aspöck et al. 1991 (mon).
Dichrostigmaavipes Stein: H. Aspöck et al. 1989 (biogeogr, distr, ill:
imag); Pantaleoni 1990a (com, rec); Pantaleoni 1990c (com, rec);
Pantaleoni 1990d [1993] (rec); H. Aspöck et al. 1991 (mon); De-
vetak 1991 (rec); Saure and Gerstberger 1991 (ecol, rec); Devetak
1992a (rec); Devetak 1992b (com); Rausch and H. Aspöck 1993
(rec, ecol); Horstmann 1994 (paras); Pantaleoni et al. 1994 (distr,
rec); U. Aspöck et al. 1995 (ethol: cop); H. Aspöck and Hölzel 1996
(distr); Hellrigl and Hölzel 1996 (list, rec); Letardi and Pantaleoni
1996 (rec); Saure 1996 (biogeogr, ecol, rec); Sziráki 1996 (ecol,
rec); Achtelig 1997 (ecol, rec); Wachmann and Saure 1997 (charact;
g: imag); U. Aspöck and H. Aspöck 1999 (ill: imag); Dobosz 1999
(rec); Sziráki 1999 (rec); Ábrahám 2001 (list, rec); H. Aspöck et
al. 2001 (anncat); Gruppe and Schubert 2001 (ecol, rec); Vas et al.
2001 (ecol, rec); Czechowska 2002 (ecol, rec); Tröger 2002 (rec);
Ábrahám et al. 2003 (rec); Dobosz 2003 (rec); Haring and U. As-
pöck 2004 (phyl); H. Aspöck and U. Aspöck 2005 (distr); U. Aspöck
and H. Aspöck 2005a (biogeogr); U. Aspöck and H. Aspöck 2005b
(com, anat); Nicoli Aldini 2005 (rec); Tröger 2005b (rec); Gruppe
2006c (ecol, rec); Anderle and U. Aspöck 2007 (rec); U. Aspöck and
H. Aspöck 2007 (ill: ♀ imag); Gruppe and Müller 2007 (ecol, rec);
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204
H. Aspöck and U. Aspöck 2007 (biogeogr, distr); Dobosz 2007 (dis-
tr, rec; ill: map); Badano 2008 (distr, rec); Gruppe 2008 (ecol, rec);
Klokočovnik et al. 2010 (rec); Letardi et al. 2010 (rec); Haring et al.
2011 (phyl, phyltree); H. Aspöck and U. Aspöck 2013 (cat, etymol;
ill: ♂, ♀ imag), 2014 (cat); Morinière et al. 2014 (barcode, phyl);
Sziráki 2014 (rec); Tillier et al. 2022а (rec).
Taxonomy. H. Aspöck et al. (1991). The species can eas-
ily be dierentiated also by eidonomic characters, partic-
ularly by the two-colored pterostigma (proximal brown,
distal yellowish) (Figs 5h, 6a).
Biology and ecology. Larvae (Fig. 6b) exclusive-
ly soil-dwelling in the litter around roots of trees and
bushes. Development two years. Last hibernating stage:
full-grown larva. Adults: (IV)V–VII(VIII). Euryoecious,
but preferring xerothermic habitats, e.g. light pine forests,
but also biotopes free of conifers.
Records on Mediterranean islands (Fig. 8b). Krk,
Hvar, Korfu. Syntopic Raphidioptera on Mediterranean
islands: No records.
Continental distribution. Balkan Peninsula (except
southern parts of Greece), eastern Europe (till the Ural),
Central Europe, northern Italy.
Biogeography. (Probably monocentric) Balkanopon-
tomediterranean faunal element with high expansivity.
Family Inocelliidae Navás, 1913
Family Inocelliidae Navás, 1913: H. Aspöck et al. 1991 (mon),
H. Aspöck and U. Aspöck 1991 (overv); H. Aspöck et al. 2001 (ann-
cat); H. Aspöck 2002 (biol); H. Aspöck and U. Aspöck 2009 (overv);
U. Aspöck and H. Aspöck 2009 (overv); H. Aspöck et al. 2012
(overv); Oswald and Machado 2018 (overv); Shen et al. 2022 (phyl).
Fibla Navás, 1915
Fibla Navás, 1915b (odescr) [type species by original designation: Fib-
lahesperica Navás, 1915]: H. Aspöck et al. 1989 (distr); H. Aspöck
1990 (biogeogr, distr, synlist; ill: map); H. Aspöck et al. 1991 (mon);
Oswald and Penny 1991 (cat, nom); U. Aspöck and H. Aspöck 1994
(biogeogr); H. Aspöck et al. 2001 (anncat); H. Aspöck 2002 (biol,
paras); Monserrat and Papenberg 2010 (chorol); Haring et al. 2011
(phyl); H. Aspöck et al. 2012 (com); H. Aspöck and U. Aspöck 2013
(cat, etymol), 2014 (cat); Monserrat and Papenberg 2015 (overv);
Shen et al. 2022 (phyl).
Taxonomy and systematics. H. Aspöck et al. (1991),
Haring et al. (2011). Systematic position within the fam-
ily not yet denitely resolved, however, see Shen et al.
(2022). It comprises two subgenera, Fibla s. str. (with 3
species) and Reisserella (with 1 species). Both subgenera
are represented on Mediterranean islands.
Biology and ecology. Larvae are exclusively cortico-
lous 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, Sic-
ily, Apennine Peninsula (sporadically), Crete, N-Africa.
Subgenus Fibla Navás, 1915, s. str.
Subgenus Fibla Navás, 1915b (odescr) [type species by original desig-
nation: Fibla hesperica Navás, 1915]: H. Aspöck et al. 1989 (bio-
geogr, distr; ill: distrmap); H. Aspöck 1990 (biogeogr, distr, synlist;
ill: map); H. Aspöck et al. 1991 (mon); H. Aspöck et al. 2001 (ann-
cat); H. Aspöck 2002 (biol, paras); H. Aspöck and U. Aspöck 2013
(cat, etymol), 2014 (cat); Monserrat and Papenberg 2015 (overv);
Shen et al. 2022 (phyl).
Burcha Navás, 1915b (odescr) [type species by original designation:
Inocelliamaclachlani Albarda, 1891]: H. Aspöck et al. 1991 (mon);
Oswald and Penny 1991 (cat, nom).
Estoca Navás, 1919 (odescr) [type species by monotypy: Estocapeyer-
imho Navás, 1919]: H. Aspöck et al. 1991 (mon).
Taxonomy. H. Aspöck et al. (1991). Fibla s. str. and Re-
isserella can easily be dierentiated by eidonomic char-
acters (and, moreover, by their geographic vicariance).
Biology and ecology. See Fibla s.l. and F.maclachlani.
Distribution. Iberian Peninsula, Tyrrhenian islands,
Sicily, Apennine Peninsula, N-Africa.
Fibla (Fibla) maclachlani (Albarda, 1891)
Inocelliacrassicornis auct. (nec Schummel!): H. Aspöck et al. 1991 (mon).
Inocellia maclachlani Albarda, 1891 (odescr): H. Aspöck et al.
1991 (mon).
Burchamaclachlani (Albarda): H. Aspöck et al. 1991 (mon).
Burcha sicula Navás, 1915a (odescr): H. Aspöck et al. 1991 (mon);
H. Aspöck & U. Aspöck 2014 (cat).
Fibla maclachlani (Albarda): H. Aspöck et al. 1991 (mon); Letardi
1994 (distrmap, ecol, rec); Pantaleoni 2005 (com, rec); Haring et al.
2011 (phyl); Tillier et al. 2022a (rec).
Fibla (Fibla) maclachlani (Albarda, 1891): H. Aspöck et al. 1991 (mon);
U. Aspöck and H. Aspöck 1994 (distr, biogeogr, ill: imag, la, map);
H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat);
Nicoli Aldini and Baviera 2001 (rec); Haring et al. 2011 (phyl);
H. Aspöck et al. 2012 (distr); Nicoli Aldini et al. 2012 (rec, distr);
H. Aspöck & U. Aspöck 2013 (cat, etymol; ill: la, ♂ imag), 2014
(cat); Pantaleoni et al. 2019 (distr, ecol, rec); Shen et al. 2022 (phyl).
Taxonomy. H. Aspöck et al. (1991). A very characteris-
tic species (Fig. 6c, d) and the only representative of the
family on the Tyrrhenian islands and in Sicily.
Biology and ecology. Larvae (Fig. 6e) exclusively cor-
ticolous, mainly on Pinus, but often also on deciduous trees
(Amygdalus, Quercus). Preferred biotopes: Light pine and
oak forests, but also in wild fruit gardens. Records from
150–1600 m; probably also occurring at sea level.
Records on Mediterranean islands (Fig. 11b). Cor-
sica, Sardinia, Sicily. Syntopic occurrence with other
Raphidioptera: Xanthostigmacorsica (Corsica, Sardinia,
Sicily), Xanthostigma aloysiana (Sardinia), Subillacon-
nis (Sicily), Subillaprincipiae (Sardinia). Development:
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Three or more years. Last hibernating stage: full-grown
larva. Adults: IV–VI.
Continental distribution. Apennine Peninsula, proba-
bly introduced from Sardinia with cork; old and recent re-
cords in Calabria need conrmation (Pantaleoni et al. 2019).
Biogeography. Polycentric (?) stationary Tyrrhe-
nian-(Adriatomediterranean?) faunal element.
Subgenus Reisserella H. Aspöck & U. Aspöck, 1971
Reisserella H. Aspöck & U. Aspöck, 1971 (odescr) (described as a sub-
genus of Inocellia Schneider) [type species by original designation:
Inocellia (Reisserella) pasiphae H. Aspöck & U. Aspöck, 1971]:
H. Aspöck et al. 1989 (distr); H. Aspöck 1990 (biogeogr, distr; ill:
map); H. Aspöck et al. 1991 (mon); Oswald and Penny 1991 (cat,
nom); H. Aspöck et al. 2001 (anncat); H. Aspöck 2012 (cat); H. As-
pöck et al. 2012 (distr); H. Aspöck and U. Aspöck 2013 (cat, ety-
mol), 2014 (cat); Shen et al. 2022 (phyl).
Taxonomy. H. Aspöck et al. (1991). Easily to be dier-
entiated from Fibla s.str. by eidonomic and by genitalic
characters. Only one species known.
Biology and ecology. See Fibla (Reisserella) pasiphae.
Distribution. Endemic to Crete.
Fibla (Reisserella) pasiphae (H. Aspöck &
U. Aspöck, 1971)
Fibla sp.: H. Aspöck & U. Aspöck, 1966 (tax, distr). H. Aspöck et al.
1991 (mon).
Inocellia (Reisserella) pasiphae H. Aspöck & U. Aspöck, 1971b (ode-
scr): H. Aspöck et al. 1991 (mon).
Filbaelkeweimarae Lauterbach, 1972 (odescr): H. Aspöck et al. 1991
(mon); H. Aspöck and U. Aspöck 2014 (cat).
Fibla (Reisserella) pasiphae (H. Aspöck & U. Aspöck): H. Aspöck et
al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); Popov 1992
(biogeogr); U. Aspöck and H. Aspöck 1994 (biogeogr; ill: distrmap);
H. Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat);
Tröger 2005a (rec); H. Aspöck 2012 (cat); H. Aspöck et al. 2012
(distr; g: imag, la, pu); H. Aspöck and U. Aspöck 2013 (cat, ety-
mol; ill: ♀ imag, la, ♂ imag, pu), 2014 (cat) ; Shen et al. 2022 (ill:
♀ imag, phyl).
Taxonomy. H. Aspöck et al. (1991). Easily recognized
(Fig. 6f, g), moreover, it is the only Inocelliid species oc-
curring in Crete.
Biology and ecology. Larvae (Fig. 6h) corticolous on
various deciduous trees (Amygdalus, Pyrus), probably
also on Pinus. Occurs in various habitats of light forests,
wild gardens, and in various types of maquis with sin-
gle trees in altitudes from sea level to (at least) 1200 m.
Development (at least) two years. Last hibernation stage:
full-grown larva. Adults: IV–VI.
Records on Mediterranean islands (Fig. 11b). Dis-
tributed all over Crete, but conned to this island. Syntopic
other Raphidioptera: Phaeostigma (Aegeoraphidia) biroi,
Ph. (Superboraphidia) minois, Raphidia (R.) ariadne.
Biogeography. Endemic to Crete. Monocentric Cretan
faunal element.
Parainocellia H. Aspöck & U. Aspöck, 1968
Parainocellia H. Aspöck & U. Aspöck, 1968 (odescr) (described as a
subgenus of Inocellia Schneider) [type species by original designa-
tion: Inocelliaressli H. Aspöck & U. Aspöck, 1965b]: H. Aspöck
et al. 1991 (mon); Oswald and Penny 1991 (cat, nom); H. Aspöck
et al. 2001 (anncat); H. Aspöck 2002 (paras); Liu et al. 2009 (syst);
Haring et al. 2011 (phyl); H. Aspöck 2012 (cat); H. Aspöck et al.
2012 (distr); H. Aspöck and U. Aspöck 2013 (cat, etymol), 2014
(cat); Shen et al. 2022 (phyl).
Taxonomy and systematics. H. Aspöck et al. (1991),
Liu et al. (2009). Parainocellia comprises presently three
species which are eidonomically similar to each other and
to other species of Inocelliidae presently in the genus Ino-
cellia. By morphological characters of the male genitalia
they can, however, easily be identied.
Biology and ecology. Larvae corticolous on many co-
niferous 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 (H. Aspöck & U. Aspöck, 1965b)
Inocelliaressli H. Aspöck & U. Aspöck, 1965b (odescr): H. Aspöck et
al. 1991 (mon); Shen et al. 2022 (phyl, ill: ♂ gs).
Inocellia (Parainocellia) ressli H. Aspöck & U. Aspöck: H. Aspöck et
al. 1991 (mon).
Parainocellia ressli (H. Aspöck & U. Aspöck): H. Aspöck et al. 1991
(mon); H. Aspöck and U. Aspöck 1991 (ill: head, wings, ♂ gs); H. As-
pöck et al. 2012 (distr); H. Aspöck and U. Aspöck 2013 (cat, etymol).
Parainocellia (Parainocellia) ressli (H. Aspöck & U. Aspöck): H. As-
pöck et al. 1989 (biogeogr, distr); H. Aspöck et al. 1991 (mon); H.
Aspöck and Hölzel 1996 (distr); H. Aspöck et al. 2001 (anncat); U.
Aspöck and H. Aspöck 2005b (ill: head, prothorax, pterothorax, tar-
salia); Canbulat and Kiyak 2006 (distr, rec; ill: map); U. Aspöck and
H. Aspöck 2007 (ill: head. la); Dobosz 2007 (distr, rec); H. Aspöck
2012 (cat); H. Aspöck and U. Aspöck 2014 (cat).
Taxonomy. H. Aspöck et al. (1991). Eidonomically very
similar to Parainocelliabraueri (Balkan Peninsula, south-
east of Central Europe, Eastern Europe) and to P.bicolor
(Apennine Peninsula, southwest of Central Europe), but
easily to be identied by characters of the male genitalia.
Biology and ecology. Larvae corticolous on Pinus,
Quercus (and probably many other trees). Typical hab-
itats: 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.
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Records on Mediterranean islands (Fig. 11b). Sa-
mos. Syntopic Raphidioptera on Samos: Phaeostigma
(Aegeoraphidia) raddai, Raphidia (R.) ambigua.
Continental distribution. South and east Anatolia.
Biogeography. Monocentric Anatoloponotmediterra-
nean 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. 12a, c shows the distribution
of these species in the world (see also Fig. 1). The Medi-
terranean region, as well as Central and Eastern Asia, are
known hotspots of biodiversity (Myers et al. 2000), which
is also reected by the distribution of Raphidioptera. By
far most snakey species occur in Europe and in the Mid-
dle East (115 species = 45,6% of the world fauna), and
of these 33 species = 13,1% of the world fauna inhabit
Mediterranean islands. 99 species occur in Central Asia
or Eastern Asia, i.e. 39,3% of the world fauna.
Only 5 species (= 2%) have so far been found in Af-
rica (restricted to the northwestern parts of the conti-
nent). America (a relatively small part of North Ameri-
ca only: see Fig. 1) harbors 32 species (= 12,7% of the
world fauna).
Fig. 12b, d shows the signicance of Raphidioptera
occurring on Mediterranean islands compared to the total
numbers of Raphidioptera, Raphidiidae and Inocelliidae
respectively within the whole Mediterranean region, i.e.
all countries of Europe, Asia, and Africa bordering the
Mediterranean Sea.
Geographical denitions: Middle East comprises Tur-
key, 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, Tai-
wan, Japan, Myanmar, and Thailand. In America Raphid-
ioptera occur in Canada, USA, and Mexico.
How have the Mediterranean islands been
colonized by Raphidioptera?
Tables 1, 2 give an overview of the 33 species of Raphid-
ioptera recorded from Mediterranean islands and of the
28 islands with conrmed occurrence of certain snakey
species. How and when have these 33 snakey species
come to the islands or have some of them ever been there?
Figure 12. The Raphidioptera of the Mediterranean region a. Raphidiidae of the world (206 spp.). Two species (Raphidiaophiopsis
and Xanthostigmaxanthostigma) 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 spe-
cies (Inocelliacrassicornis) 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.).
Africa
4
Europe,
Middle East
78
on islands
30
Central,
Eastern Asia
66
America
28
Europe,
Middle East
108
Africa
1
Europe,
Middle East
4
on islands
3
Central,
Eastern Asia
33
America
5
Europe,
Middle East
7
species not
recorded from
islands
74
endemic to
Mediterranean
islands
9
not endemic
21
species on
islands
30
species not
recorded from
islands
5
2
not endemic
1
species on islands
3
ab
dc
endemic to
Mediterranean
islands
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
dez.pensoft.net
207
The palaeogeography of the Mediterranean islands is
very diverse – a fact which is impressively reected also
by their diverse Raphidioptera fauna. The most important
factors are origin, emergence, age, and development of
the dierent islands on one 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 Mediter-
ranean Sea between 5 and 6 mya led to many land bridges
but these were of no major signicance for the dispersal
of the snakeies as the dried-up areas were largely cov-
ered by evaporates or had the characters of lagoons.
Here we try to analyze the origin of the Raphidioptera
fauna of those Mediterranean islands where snakeies
have been found:
Tyrrhenian islands
The Tyrrhenian islands Sardinia and Corsica were sep-
arated 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 snakey spe-
cies have been found (Xanthostigma corsica and Fibla
(F. ) maclachlani), in Sardinia four species (X. corsica,
X. aloysiana, Subilla principiae, and F. (F. ) maclach-
lani). 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. Aspöck et al. 1991). The most closely related
species are X.aloysiana and X.xanthostigma (Schummel,
1832), both occurring in large parts of the continent, X.
aloysiana in Italy, S-Switzerland, France and NE-Spain,
X.xanthostigma as a Eurosibirian species from Europe to
the Far East (H. Aspöck et al. 1991). It is thus a reason-
able assumption that X. corsica has reached the islands
by dispersal from the continent. When these invasions to
Sardinia, Corsica, and other islands may have occurred, is
unknown. The situation with X.aloysiana is very similar.
A third species of the family Raphidiidae occurring in
Sardinia was detected at the beginning of this century: Su-
billaprincipiae. The species is rather isolated within the
genus, and the sister taxon cannot be identied. S. prin-
cipiae 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 connis (Stephens, 1836),
which is an expansive Adriatomediterranean faunal el-
ement. Moreover, all other Subilla species are Ponto-
mediterranean 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.) maclach-
lani, is clearly of Iberian origin and thus a very old faunal
element in Corsardinia. It occurs also in Sicily, but pri-
marily not in the Italian mainland and can thus be regard-
ed 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 (Pantaleoni et al. 2019). Fibla (F. ) maclachlani
is closely related to Fibla (F. ) hesperica, which is widely
distributed on the Iberian Peninsula. Together with Fib-
la (F. ) peyerimho (Navás, 1919), which is known from
various parts of N-Morocco, N-Algeria, and N-Tunisia,
they form the subgenus Fibla Navás, 1915. One may as-
sume that in the Oligocene when the landmasses which
later formed Corsica and Sardinia got separated from
Iberia they took the ancestors of the extant Fibla (F.) ma-
clachlani with them into the insular isolation.
Adriatic and Ionian islands
The islands in the Adriatic Sea as well as those in the Io-
nian Sea had connections to the continent, i.e. the Balkan
Peninsula, during the Pleistocene. This led to an immigra-
tion of species from the continent into the former islands.
The Raphidioptera fauna of the islands of Krk, Hvar,
Korfu, Levkas, and Kefalonia comprises only snake-
y species which occur also in the adjacent continent:
Dichrostigmaavipes (Krk, Hvar, Korfu), Parvoraphid-
ia microstigma (Levkas), Ornatoraphidia avilabris
(Levkas), Subilla artemis (Levkas), Phaeostigma
(Magnoraphidia) major (Levkas, Kefalonia).
Northern and western Aegean islands
Also the islands of the Northern Aegean Sea harbor only
snakey species of the adjacent continental regions, to
which they were connected by land bridges in the Pleisto-
cene: Phaeostigma (Pontoraphidia) setulosaaegea (Tha-
sos), Raphidia (R.) beieri (Thasos, Samothraki), Venust-
oraphidianigricollis (Thasos).
Among the Sporades only Skopelos and Skyros have
been investigated for Raphidioptera. Skopelos harbors
Phaeostigma (Magnoraphidia) ammi, 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
to one land mass in the Pleistocene.
On the island of Skyros so far only one snakey species
has been found: Raphidiamediterranea, 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 snakey species: Ornatoraphid-
ia christianodagmara, Phaeostigma (Ph.) euboica, Ph.
(Graecoraphidia) divinaretsinata, Ph. (Magnoraphidia)
ammi, Ph. (M.) wewalkai, and Raphidia (R.) mediterra-
nea. During the Pleistocene Euboea was repeatedly widely
connected with continental Greece for long periods so that
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there must have been an intensive faunal exchange. It is
thus surprising that one of the six species of the snakeies
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 iso-
lated populations in two mountain ranges in Attika (Par-
nis, Pateras). The other two subspecies of Ph. (G.) divina,
Ph.d. divina and Ph. d. simillima, are conned 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 con-
ned to Greece probably throughout the whole Pleistocene.
Euboea harbors two species of Magnoraphidia: Phae-
ostigma (M.) ammi and Ph. (M.) wewalkai. Both species
occur in restricted areas of the adjacent continental region,
from where they may have colonized Euboea in the Pleisto-
cene. 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 ad-
jacent continental regions in one of the glacial periods.
The genus Ornatoraphidia comprises only one further
species: O. avilabris, which occurs in many parts of
Greece, however, neither in Euboea nor in Attica.
Raphidia (R.) mediterranea is a Balkanopontomed-
iterranean faunal element, which has been probably in-
troduced 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. Aspöck et al. 2017).
The only snakey species so far recorded from An-
dros, from Aegina, and from Hydra is Raphidia (R.)
mediterranea (see above). As regards the Kyklades,
snakeies have been found on the islands of Naxos and
Paros. Naxos has been intensively investigated. Only
one snakey species has been found on each of these is-
lands: 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 snakeies, possibly only R. mediterra-
nea. Since the formation of the Aegean Sea due to tec-
tonic events about 10 mya the islands had repeated con-
nection to one another or were overooded and emerged
again, and none of the Kyklades remained isolated for
a suciently long time to allow the formation of a sta-
ble snakey fauna. Moreover, all these islands lack high
mountains, which are an important precondition for the
formation of refugial subcenters.
Crete
A totally dierent 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
overooded, 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 mil-
lion years. This is why Crete harbors so many endemic
animals and plants. Crete harbors four snakey species –
three species of the family Raphidiidae and one 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 nd out the
nearest relatives of these species which is always a chal-
lenging task of phylogeography.
Phaeostigma (Aegeoraphidia) biroi is related to Ph.
(Ae.) karpathana (endemic to Karpathos), Ph. (Ae.) pro-
phetica (endemic to Rhodes), Ph. (Ae.) ressli (SW-Anato-
lia), and Ph. (Ae.) vartianorum (SW-Anatolia). The whole
subgenus is of distinct Anatolian origin, but it remains un-
known how and when (the ancestors of) the species have
come to Crete. Before the development of the Aegean Sea
Greece and Anatolia formed one large land mass. Phae-
ostigma (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.mammaph-
ila (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 isolat-
ed within the subgenus. Ph. (S.) minois may be a descen-
dent of the old fauna of Crete. The fourth snakey 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 iso-
lated? At rst one may think of a possible land-bridge to
the north of Africa in a glacial period, but according to pa-
laeogeological 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
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
dez.pensoft.net
209
by swimming wood from the western part of the Mediter-
ranean 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 Karpa-
thos emerged as an island. Later, in the Messinian, Kar-
pathos and Rhodes became united, but from the late Plio-
cene onwards Karpathos became again isolated, while
Rhodes was connected with the Anatolian mainland. In
the Pleistocene both islands remained isolated. This pa-
laeogeographical history explains the snakey fauna of
Karpathos and Rhodes. Karpathos (the island has been
investigated intensively by us for Raphidioptera) har-
bors two species of Raphidioptera, both Raphidiidae:
Phaeostigma (Aegeoraphidia) karpathana and Raphidia
(R.) mediterranea. Ph. (Ae.) karpathana is either the de-
scendant 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 dierent
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 mo-
lecular systematic study. However, presently there is no
adequately preserved material available. R.mediterranea
has been most probably introduced from its original re-
fugial area in Greece into many other parts of its present
distribution area (H. Aspöck et al. 2017), mainly by hu-
man activities, but probably also by natural ways (e.g.
swimming wood) so that the species may have reached
also Karpathos at any time in the past.
Rhodos
Rhodes (Rhodos) is inhabited by three species of Raphid-
ioptera, all Raphidiidae: Phaeostigma (Ae.) prophetica,
Subillacolossea, and Raphidia (R.) ambigua. It is out of
the question that all three species are of Anatolian ori-
gin. 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 in-
vestigated thoroughly have to be especially considered:
Lesbos, Chios, Samos, Ikaria. All four islands were reg-
ularly 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 Phaeostig-
ma (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 distrib-
uted 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, rep-
resented 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 dierent species of
Raphidia s.str., and all four species occur on “their” is-
land from coastal areas to the highest elevations. On the
island of Samos Parainocelliaressli was found. The spe-
cies 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 phytogeo-
graphical studies in the Aegean area (Rechinger 1950;
Rechinger and Rechinger-Moser 1951) and could show
that there is a biogeographical borderline between the
Cyclades and the East Aegean Islands (Lesbos, Chios, Sa-
mos, Ikaria, Rhodes and the other islands of this region)
separating the ora of Europe from that of Asia. Strid
(1996) proposed the name “Rechinger´s Line” for this
boundary. This line can convincingly be conrmed by the
distribution of the Raphidioptera of the Aegean Islands.
Cyprus
Cyprus is inhabited by two snakey species, both Raphi-
diidae: Phaeostigma (Crassoraphidia) cyprica and Ulrike
syriaca. Both species occur in Asian countries bordering
the Mediterranean Sea (Syria, Lebanon, Israel). It is sur-
prising that none of the many Raphidioptera species oc-
curring in the south of Anatolia have been found on the
island of Cyprus which is a strong argument for the as-
sumption that Cyprus was never connected to Anatolia.
Cyprus is of Levantine origin by a submarine break o, 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
(Jolivet et al. 2006; Bache et al. 2012). In this case Cyprus
could have easily received terrestrial organisms from the
Levante. Otherwise, the island must have been colonized
by introductions by wind or via wood drifted in the sea. At
any rate, the two snakey species presently occurring in
Cyprus cannot be dierentiated from individuals from the
Levante morphologically. This supports the assumption of
a late introduction, in the Pleistocene or even in the Holo-
cene, possibly by human activities. The Neuropterida fau-
na of Cyprus is – compared to other large Mediterranean
islands – moderate and comprises only three endemisms
(Badano and Makris 2020), which is in agreement with
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Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
210
the comparatively young age of the island and the highly
reduced ways of invasions by species from the continent.
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 armed. The
majority of the islands of the Mediterranean Sea are small,
frequently rocks of low altitude and uninhabited. General-
ly speaking, snakeies 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 snakeies are
fullled in large islands with elevations of 500 to 1500 m.
Of the 28 islands with records of Raphidioptera the
following islands have been specically investigated in-
tensively 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 Raphid-
ioptera, 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
snakeies, however none were found. The small islands
o the southern coast of Anatolia, in particular Kastelorizo
(highest elevation <300 m), may harbor one or two snake-
y 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. Prob-
ably no Raphidioptera will be found on any of the islands
of Malta. The highest elevations of Malta are less than
250 m asl. Raphidiamediterranea could occur, if it has
been introduced. There is an entirely dierent 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 o from the mainland and having given rise to
the Balearic Islands harbors the remnants of the old snake-
y 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 snakey 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 (Subillaprincipiae)
• Crete (Phaeostigma (Aegeoraphidia) biroi,
Ph. (Superboraphidia) minois, Raphidia (R.) ari-
adne, Fibla (Reisserella) pasiphae)
• Rhodes (Phaeostigma (Aegeoraphidia) prophetica,
Subillacolossea)
• 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 Phaeostig-
ma on Euboea there is, however, a quite dierent situation:
The species is conned to small areas at high elevations.
Possibly these areas are more or less (with shifts of ver-
tical 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 phylogeograph-
ical studies
So far all identications of snakeies 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 moun-
tains. Other subspecies – Ph. (G.) d. divina and
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
dez.pensoft.net
211
Ph. (G.) d. simillima – have been described from
other mountains north of the Gulf of Korinthos. It
would be of interest to investigate various popula-
tions 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 Ana-
tolian 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 spe-
cies 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 po-
sition 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 har-
bors an endemic species. Comparative molecular
studies with other species of the subgenus occur-
ring 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 East-
ern Europe and even in an old farmhouse in Central
Europe. Comparative molecular studies of speci-
mens 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, partic-
ularly also with respect to the slight morphological
dierences found in the population of Karpathos.
It is unlikely that cryptic species will be detected on
Mediterranean islands by detecting signicant genomic
dierences compared to populations of continental re-
gions. All these species have colonized the islands quite
recently. Moreover, genetic dierences without morpho-
logical correlations can hardly justify the dierentiation
of separate species (H. Aspöck et al. 2021).
Endangered species of Raphidioptera on Mediterra-
nean islands
Finally, we would like to raise the question whether any
of the snakey species occurring on Mediterranean is-
lands 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 conned to such
small areas that they could possibly be eradicated if their
habitats are destroyed: Phaeostigma (Ph.) euboica (Eu-
boea), Ph. (Graecoraphidia) divina retsinata (Euboea),
Ornatoraphidia christianodagmara (Euboea), Ph. (Su-
perboraphidia) minois (Crete), and particularly Subilla
principiae (Sardinia). This latter species has so far only
been found in a small light forest of Quercuspubescens.
Acknowledgements
The photographs of living specimens were taken by Mag.
Franziska Anderle, now Denner (Lower Austria, Hörers-
dorf), Mag. Harald Bruckner and Peter Sehnal (Natural
History Museum Vienna). The photograph 6g was kindly
provided by Prof. Dr. Hannes Paulus (University of Vi-
enna). Moreover, H. Bruckner provided all distribution
maps. Grateful thanks for their patience, skill and consci-
entiousness. Dr. Alexandra R. Szewczyk, B.A., Dr. Ed-
win 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 (Geologi-
cal-Paleontological Department, Natural History Muse-
um Vienna) for many stimulating hours of fruitful discus-
sion and for providing literature on the geological history
of the Mediterranean Sea. Moreover, we gratefully ac-
knowledge further information on the Mediterranean Sea
in the course of the Pleistocene received from Univ. Prof.
Dr. Mathias Harzhauser (Geological-Paleontological De-
partment, 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 situa-
tions and – last but not least – for unforgettable weeks of
joint eld work in wonderful biotopes harboring snake-
ies: 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 (Uni-
versity of Maribor, Slovenia), Prof. Dr. Peter Duelli
(Swiss Federal Institute for Forest, Snow and Landscape
Research, Birmensdorf, Switzerland), and Prof. Dr. Mi-
chael Ohl (Museum für Naturkunde Berlin, Germany) for
thoroughly reviewing and improving the manuscript, and
to Dr. Dominique Zimmermann (Natural History Muse-
um Vienna), Editor in Chief of DEZ and Subject Editor,
dez.pensoft.net
Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
212
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 aldery, snakey 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 Somo-
gy county (Megaloptera, Raphidioptera, Neuroptera).] https://doi.
org/10.24394/NatSom.2001.1.253
Ábrahám L (2006) A Mecsek recésszárnyú-alkatú faunája és újabb fau-
nisztikai adatai (Neuropterida: Megaloptera, Raphidioptera, Neu-
roptera). [= 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, Raphidiop-
tera, 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 Dierent 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) Kamelhalsiegen (Ins. Raphidioptera) aus der
Umgebung von Würzburg. Galathea, Nürnberg 3(Suppl.): 48–51.
Aistleitner U, Gruppe A (2009) Nachweise von Netzüglern aus dem
Fürstentum Liechtenstein (Insecta: Neuropterida: Raphidioptera,
Megaloptera, Neuroptera). Entomofauna 30(13): 209–220.
Albarda H (1891) Révision des Rhaphidides. Tijdschrift voor Entomol-
ogie 34: 65–184.
Anderle F, Aspöck U (2007) Neuropterida (Insecta Endopterygota) of
the Nature Reserve Eichkogel (Lower Austria): arguments for pro-
tecting an insular biocoenosis in the South of Vienna. In: Pantaleoni
RA, Letardi A, Corazza C (Eds) Proceedings of the Ninth Interna-
tional 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 Inter-
national Symposium on Neuropterology. Bergen Dal, Kruger Na-
tional Park (R.S.A.). Pretoria, 1988: 47–61.
Aspöck H (1998) Distribution and biogeography of the order Raphid-
ioptera: updated facts and a new hypothesis. In: Panelius S (Ed.)
Neuropterology 1997. Proceedings of the Sixth International Sym-
posium 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 wissenschaft-
shistorischer 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. (Vorläuge Beschreibung). Entomol-
ogisches 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 Südos-
teuropa und Kleinasien. (Mit kritischen Bemerkungen zur Klassi-
kation der Familie). Annalen des Naturhistorischen Museums in
Wien 68: 309–364.
Aspöck H, Aspöck U (1965b) Vorläuge Mitteilung über Untersuchun-
gen an europäischen Inocelliidae (Neuroptera, Raphidiodea). Ento-
mologisches 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) Agullaattica nov. spec. – eine neue Ra-
phidiiden-Art aus Griechenland (Insecta, Neuroptera). Entomolo-
gisches Nachrichtenblatt, Wien 14: 7–11.
Aspöck H, Aspöck U (1967b) Raphidia setulosa nov. spec. aus dem Bal-
kan-Gebirge. Mit Bemerkungen über die Raphidiiden (Insecta, Neurop-
tera) Bulgariens.–. Entomologisches Nachrichtenblatt, Wien 14: 17–20.
Aspöck H, Aspöck U (1968) Vorläuge Mitteilung zur generischen
Klassizierung der Raphidiodea (Insecta, Neuroptera). Entomolo-
gisches Nachrichtenblatt, Wien 15(7–8): 53–64.
Aspöck H, Aspöck U (1970) Raphidia (Ornatoraphidia) christianodag-
mara n. sp. – eine neue europäische Raphidiiden-Spezies (Insecta,
Raphidioptera). Entomologische Berichten (Amsterdam) 30: 99–102.
Aspöck H, Aspöck U (1971a) Drei neue europäische Raphidi-
iden-Spezies (Neuropteroidea – Raphidioptera). Nachrichtenblatt
der Bayerischen Entomologen 20: 86–88.
Aspöck H, Aspöck U (1971b) Inocellia (Reisserella n. subgen.) pasi-
phae n. sp. aus Kreta (Ins., Raphidioptera, Inocelliidae). Entomolo-
gische 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 Auösung des Raphidianotata-Kom-
plexes (Neur., Raphidioptera, Raphidiidae). Zeitschrift der Arbeits-
gemeinschaft Österreichischer Entomologen 27(3/4): 57–70.
Aspöck H, Aspöck U (1991) Raphidioptera (Snake-ies, camel-
neck-ies). In: Naumann ID et al. (Eds) The Insects of Australia.
A textbook for students and research workers (2nd edn. Vol. I). Mel-
bourne University Press, 521–524.
Aspöck H, Aspöck U (2007) The Raphidioptera of the Apennine Pen-
insula: a biogeographical analysis. In: Pantaleoni RA, Letardi A,
Corazza C (Eds) Proceedings of the 9th International Symposium on
Dtsch. Entomol. Z. 70 (1) 2023, 175–218
dez.pensoft.net
213
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 – Kamelhalsiegen. 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 Kamelhalsiegen der Erde: Systematisches Ver-
zeichnis und Etymologie (Insecta: Endopterygota: Neuropterida:
Raphidioptera). Entomologica Austriaca: Zeitschrift der Osterre-
ichischen Entomologischen Gesellschaft 20: 9–155.
Aspöck H, Aspöck U (2014) Die Autoren der Taxa der rezenten Ra-
phidiopteren (Insecta: Endopterygota). Entomologica Austriaca:
Zeitschrift der Osterreichischen Entomologischen Gesellschaft 21:
9–152.
Aspöck H, Aspöck U (2021) Die Schwarzhalsige Kamelhalsiege – Ve-
nustoraphidia nigricollis. Insekt des Jahres 2022 Deutschland Öster-
reich Schweiz. Herausgeber: Kuratorium Insekt des Jahres, Senck-
enberg Deutsches Entomologisches Institut. Achtseitiges Faltblatt.
Aspöck H, Aspöck U (2022a) Die Schwarzhalsige Kamelhalsiege,
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, Med-
iterranean Asia and of Europe: a comparative review (Insecta). In:
Canard M, Aspöck H, Mansell MW (Eds) Pure and Applied Re-
search 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: Neuropteri-
da). In: Weihrauch F, Frank O, Gruppe A, Jepson JE, Kirschey L, Ohl M
(Eds) Proceedings of the XIII International Symposium of Neuropterol-
ogy, 17–22 June 2018, Osmylus Scientic Publishers, Germany, 79–93.
Aspöck H, Aspöck U, Rausch H (1977) Polyzentrische Ausbreitung
eines „sibirisch-mediterranen“ Faunenelements am Beispiel der
polytypischen Kamelhalsiege Raphidiaophiopsis L. (Neuroptera,
Raphidioptera, Raphidiidae). Zeitschrift der Arbeitsgemeinschaft
Österreichischer Entomologen 28(4): 89–105.
Aspöck H, Aspöck U, Rausch H (1979) Beschreibung von Raphid-
ia (Subilla) colossea n. sp., einer neuen Raphidiiden-Spezies von
Rhodes, und Bemerkungen über Raphidia (Subilla) physodes Navas
(Neuropteroidea: Raphidioptera). Zeitschrift der Arbeitsgemein-
schaft 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 (Neu-
ropteroidea: Raphidioptera). Entomologische Zeitschrift mit Insek-
tenbörse, 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 East-
ern Mediterranean: A Zoogeographical Analysis. Biologia Gal-
lo-Hellenica 15: 67–111.
Aspöck H, Aspöck U, Rausch H (1991) Die Raphidiopteren der Erde.
Eine monographische Darstellung der Systematik, Taxonomie, Bi-
ologie, Ökologie und Chorologie der rezenten Raphidiopteren der
Erde, mit einer zusammenfassenden Übersicht der fossilen Raphid-
iopteren (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 Raphidiop-
teren: Eine zusammenfassende Übersicht des gegenwärtigen Wissens-
standes (Insecta: Neuropteroidea). Mitteilungen der Deutschen Ge-
sellschaft fur Allgemeine und Angewandte Entomologie 9: 393–402.
Aspöck H, Aspöck U, Rausch H (1999) Biologische und chorolo-
gische Charakterisierung der Raphidiiden der östlichen Paläarktis
und Verbreitungskarten der in Kasachstan, Kirgisistan, Usbekistan,
Turkmenistan und Tadschikistan nachgewiesenen Arten der Fami-
lie (Neuropterida: Raphidioptera: Raphidiidae). In: Aspöck H (Ed.)
Neuropterida: Raphidioptera, Megaloptera, Neuroptera. Kamel-
hälse, Schlammiegen, Ameisenlöwen. Stapa 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 (Neurop-
terida: Raphidioptera). A review of present knowledge. Mitteilungen
der Deutschen Gesellschaft fur Allgemeine und Angewandte Ento-
mologie 18: 565–573.
Aspöck H, Abbt V, Aspöck U, Gruppe A (2018) The Phenomenon of
Metathetely, formerly known as Prothetely, in Raphidioptera (Insec-
ta: 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 Cen-
tral European populations of Ornatoraphidiaavilabris (Costa) come
from? (Neuropterida: Raphidioptera: Raphidiidae). Deutsche Ento-
mologische 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 snakey (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 Arbeit-
sgemeinschaft Österreichischer Entomologen 21(4): 109–120.
Aspöck U, Aspöck H (1989) Phaeostigmakarpathana n. sp. – eine neue
Kamelhalsiege von der südostägäischen Insel Karpathos (Neu-
ropteroidea: Raphidioptera: Raphidiidae). Zeitschrift der Arbeits-
gemeinschaft Österreichischer Entomologen 41(1/2): 25–31.
Aspöck U, Aspöck H (1990) Phaeostigma (Superboraphidia) minois n.
sp. – eine neue Kamelhalsiege aus Kreta (Neuropteroidea: Raphid-
ioptera: Raphidiidae). Zeitschrift der Arbeitsgemeinschaft Österre-
ichischer Entomologen 41(3/4): 71–76.
Aspöck U, Aspöck H (1994) Paradoxe Verbreitungsbilder bei Neurop-
teroidea (Insecta: Raphidioptera, Neuroptera). Zeitschrift der Arbe-
itsgemeinschaft Österreichischer Entomologen 46(1/2): 30–44.
Aspöck U, Aspöck H (1999) Kamelhälse, Schlammiegen, Ameisen-
löwen ... Wer sind sie? (Insecta: Neuropterida: Raphidioptera, Meg-
dez.pensoft.net
Horst Aspöck & Ulrike Aspöck: Snakeflies of the Mediterranean islands
214
aloptera, Neuroptera). In: Aspöck H (Ed.) Neuropterida: Raphidi-
optera, Megaloptera, Neuroptera. Kamelhälse, Schlammiegen,
Ameisenlöwen... Stapa 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 Mit-
teleuropa [SIEEC]) 37: 29–38.
Aspöck U, Aspöck H (2005b) Neuropterida (Neuropteroidea, Neurop-
tera sensu lato), Ordnungen 28–30. In: Dathe HH (Ed.) 5. Teil: In-
secta. Lehrbuch der Speziellen Zoologie. Begründet von A. Kaest-
ner. Zweite Auage. 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 oberösterre-
ichischen Landesmuseen. Neue Serie 66: 451–516.
Aspöck U, Aspöck H (2009) Raphidioptera (Snakeies). 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 Neuropteri-
da: long lasting and current controversies and challenges (Insecta: En-
dopterygota). Arthropod Systematics & Phylogeny 70(2): 119–129.
Bache F, Popescu SM, Rabineau M, Gorini C, Suc JP, Clauzon G, Oliv-
et JL, Rubino JL, Melinte-Dobrinescu MC,